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respiratory complex assembly/ respirasome assembly #3

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ValWood opened this issue Oct 17, 2024 · 31 comments
Open

respiratory complex assembly/ respirasome assembly #3

ValWood opened this issue Oct 17, 2024 · 31 comments
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ValWood commented Oct 17, 2024 •
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Systematic ID Gene name Product description Budding yeast orthologs Human orthologs Targets
SPBC115.02c afg1 AFG1 family mitochondrial ATPase/chaperone Afg1 AFG1 AFG1L
SPBC21D10.07 cmc1 copper-binding protein of the mitochondrial intermembrane space Cmc1 CMC1,CMC2 CMC1,CMC2
SPBC16E9.03c coa1 cytochrome c oxidase assembly protein Coa1 COA1 COA1
SPAC15A10.17 coa2 cytochrome c oxidase assembly protein Coa2 COA2 NDUFA3
SPAC1B3.21 coa3 cytochrome c oxidase assembly protein Coa3 COA3 COA3
SPCC550.01c coa4 cytochrome c oxidase assembly protein Coa4 COA4 COA4
SPBC16A3.16 coa5 cytochrome c oxidase assembly protein Coa5 PET191 COA5
SPBC24C6.13 coa6 cytochrome c oxidase assembly protein Coa6 COA6 COA6
SPAC1420.04c cox1101 cytochrome c oxidase assembly protein Cox1101/ mitochondrial ribosome assembly protein Rsm22 fusion protein COX11,RSM22 COX11,METTL17
SPAC19B12.13 cox1102 cytochrome c oxidase assembly protein Cox1102/ mitochondrial ribosome assembly protein Rsm2202, fusion protein COX11,RSM22 COX11,METTL17
SPCC757.15 cox14 cytochrome c oxidase assembly protein Cox14 COX14 COX14
SPBC25H2.18 cox20 cytochrome c oxidase assembly protein Cox20 COX20 COX20
SPBC1604.25 pet117 cytochrome c oxidase assembly protein Pet117 PET117 PET117
SPAC25B8.07c rcf1 cytochrome c oxidase assembly protein Rcf1 RCF1 HIGD1A,​HIGD1B,​HIGD1C,​HIGD2A,​HIGD2B
SPBC1215.01 shy1 cytochrome c oxidase assembly protein Shy1 SHY1 SURF1
SPCC4B3.20 cmc2 cytochrome c oxidase assembly protein, copper-binding, Cmc2 CMC1,CMC2 CMC1,CMC2
SPAC1565.01 rcf2 cytochrome c oxidase subunit Rcf2 RCF2 HIGD1A,​HIGD1B,​HIGD1C,​HIGD2A,​HIGD2B
SPAC2C4.09   mitochondrial calcium uniporter regulator FMP32 CCDC90B,MCUR1
SPAC3H1.08c   mitochondrial calcium uniporter regulator FMP32 CCDC90B,MCUR1
SPBC947.14c cbp6 mitochondrial complex III assembly protein Cbp6 CBP6 UQCC2
SPAC1486.08 cox16 mitochondrial copper chaperone for cytochrome c oxidase Cox16 COX16 COX16
SPBC26H8.14c cox17 mitochondrial copper chaperone for cytochrome c oxidase Cox17 COX17 COX17
SPCC1672.04c cox19 mitochondrial copper chaperone for cytochrome c oxidase Cox19 COX19 COX19
SPBC119.06 sco1 mitochondrial copper chaperone for cytochrome c oxidase Sco1 SCO1,SCO2 SCO1,SCO2
SPAC25B8.04c mss51 mitochondrial cytochrome C oxidase assembly protein Mss51 MSS51 MSS51
SPAC4G8.11c atp10 mitochondrial F1-FO ATP synthase chaperone Atp10 ATP10  
SPAC3A12.12 atp11 mitochondrial F1-FO ATP synthase chaperone Atp11 ATP11 ATPAF1
SPAC9.12c atp12 mitochondrial F1-FO ATP synthase chaperone Atp12 ATP12 ATPAF2
SPBC3B8.10 ina17 mitochondrial F1-FO ATP synthase peripheral stalk assembly factor Ina17 INA17  
SPAC926.10 ina22 mitochondrial F1-FO ATP synthase peripheral stalk assembly factor Ina22 INA22  
SPCC1442.15c cox18 mitochondrial inner membrane insertase Cox18 COX18 COX18
SPCC320.12 atp23 mitochondrial inner membrane peptidase Atp23 ATP23 ATP23
SPAC1486.11 fmc1 mitochondrial proton-transporting ATP synthase complex assembly factor, LYR family, Fmc1 FMC1 FMC1
SPAC1834.10c   mitochondrial proton-transporting ATP synthase complex assembly protein   TMEM70
SPAC25A8.03c   mitochondrial repliratory complex assembly factro/arginine methyltransferase, human NDUFAF7 ortholog YKL162C NDUFAF7
SPBC26H8.16 sdh8 mitochondrial respiratory chain complex II assembly factor 4 Sdh8 SDH8 SDHAF4 sdh1 (chaperone)
SPCC4B3.17 cbp3 mitochondrial respiratory chain complex III assembly protein Cbp3 CBP3 UQCC1
SPBC30D10.21 mzm1 mitochondrial respiratory chain complex III assembly protein, LYR family Mzm1 MZM1 LYRM7
SPBC27B12.14 cbp4 mitochondrial respiratory complex III assembly protein Cbp4 CBP4 UQCC3
SPBC3H7.08c mra2 mitochondrial respiratory complex/ ATP synthase assembly protein Mra2 MRA1 C15orf48,NDUFA4,NDUFA4L2
SPAPB8E5.10 min8 mitochondrial respiratory complex/ATP synthase complex assembly protein Mra1, meiosis specific splicing in fission yeast MRA1 C15orf48,NDUFA4,NDUFA4L2
SPAC644.07 bcs1 mitochondrial Rieske ISP assembly ATPase Bcs1 BCS1 BCS1L
SPAC664.12c sdh6 mitochondrial succinate dehydrogenase assembly factor, LYR family Sdh6 SDH6 SDHAF1 sdh2
SPBP23A10.03c sdh7 mitochondrial respiratory chain complex II assembly LYR family protein Sdh7 SDH7 SDHAF3 sdh2
SPAC12B10.06c emi5 succinate dehydrogenase complex assembly protein Emi5 SDH5 SDHAF2
SPBC26H8.01 thi2 thiazole biosynthetic enzyme THI4  
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This is great. I have a rough but incomplete list of assembly factors. I will start putting them in some order.
The HGNC have a list, if you want to check it against the human orthos: https://www.genenames.org/data/genegroup/#!/group/645

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ValWood commented Oct 18, 2024

need to deal with this too, is it a transferase? (flavination sounds like an activity)
Screenshot 2024-10-18 at 09 13 11

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ValWood commented Oct 24, 2024 •
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Notes

  • anything which is an LRR repeat protein is upstream of assembly. These are all involved in various aspects of gene expression (transcription, translation and splicing), but historically got annotated to complex assembly because the complexes don't for. I see a few of those in there.
  • also cox10 protoheme IX farnesyltransferase, catalyzes conversion of protoheme to heme A in heme A, so its required upstream of cyc1, cox1, cyt1 and tim18 (all have heme centres), was originally isolated as an assembly defect in cerevisiae
  • There do appear to be genes specifically involved in the assembly of supercomplexes (aka respirasomes) i.e The levels of supercomplexes III2IV2 and III2IV were found to be lower in ∆shy1 cells compared to WT cells, while the abundance of COA complexes increased (Fig. 7a), indicating that the formation of supercomplexes involving complex IV is influenced by deletion of shy1. In contrast, the abundance of III2 and V complexes was largely unchanged in ∆shy1 cells (Fig. 7a).
  • UQCRB is ubiquinol-cytochrome-c reductase complex subunit 6 (so a respiratory complex III subunit rather than an assembly factor)
  • TACO1 is a translational activator, I plan to include in this it in this model http://noctua.geneontology.org/workbench/noctua-visual-pathway-editor/?model_id=gomodel%3A67086be200000519
  • TIMM21 upstream, import into matrix
  • respiratory chain complex IV

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ValWood commented Oct 24, 2024

@hattrill should "NOT SEEN" be in the final column for the ones below NDUFAB1 (I am assuming this always means that it isn't in my list?)

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Hi @ValWood there are quite a few that I have doubts about - some are modifiers and protein maturation factors that have made their way onto HGNC, MitoCarta or via GO - IBA, ISS, etc. As this is a mash-up of HGNC calls, MitoCarta calls and what we have in the GO, think there will be a lot of removal from this.

We have COX6B1 as a MITOCHONDRIAL COMPLEX IV - SUPERNUMERARY SUBUNIT gene groups. - I've made a note about those as many of the supernumerary/accessory subunits have assembly GO terms - I think that this is because they are required for complex stability rather than assembly and it's 'cellular phenotype that's been annotated. That's why need to align it against reviews.

"NOT SEEN" means that I found an ortholog via DIOPT for Pombe but not on your list (or I could have missed when I did the cross-search.

Human ATPSCKMT, FDXR, FDX2, HCCS, LRPPRC, OXA1L, SCO1 - come from HGNC or MitoCarta are ones on my current 'dump' list as they seem upstream to assembly.

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Hopefully fixed now - some columns got merged a messed the layout up.

I will be removing a lot from the list....just trying to work out where to start - think that ones that are classed as accessory/supernumerary subunits will be first stop there are a lot.

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ValWood commented Oct 24, 2024 •
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OK once you have cleaned up, I'll review the pombe ones and make sure all of mine are in your table.
I have some definite orthologs for some:
COA2 is SPBC16E9.03c(coa2)
TTC19 is SPAP27G11.02(mgr3) equivalent complex member
FMC1 is SPAC1486.11(fmc1)
AIFM1 is SPAC26F1.14c(aif1)
SAMM50 is SPAC17C9.06(sam50)
pombe has no NDUFA2 (the human ortholog of pombe mrpl51 is human MRPL51)
So I will add these (with the evidence once I dig it out).

I'll also have a good look for anything that is missing from pombe (although I am not expecting to find the Complex I assembly - because we don't have it). We do have 3 (I think ) of the subunits but these might really be assembly factors that are also used by other complexes (or something like that).

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This looks like a nice review publication to start with for me:
https://doi.org/10.1042/EBC20170098....complex I assembly looks fun! Starting to wish we had no complex I ;-)

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ValWood commented Oct 24, 2024

The Wikipedia page isn't half bad, I was looking at it the other week when I found some of the assembly factors in pombe
https://en.wikipedia.org/wiki/Respiratory_complex_I

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Think I've found the paper that 'poisoned' the complex I assembly well - A lot of annotations originate from PMID:27626371 in which the KO of different accessory CI SUs were examined for their role in assembly. However, although the absence of SUs may impact assembly, I think that this is because they have a structural role in the complex/complex intermediates rather than being factors that assist in the assembly of the complex.

I like this as a definition of assembly "This multistep process involves transiently associated assembly factors that integrate core and accessory/supernumerary subunits as well as cofactors into the final holoenzyme." (PMID: 38086790).

I will have a bit more of a dig and then ask Alex Whitworth as if this works need to get a few places to revise their lists as well as GO.

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hattrill commented Nov 7, 2024 •
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Think that list is ok now - will take a bit for GO fixes to filter down.
(validating that no orthologs in Pombe for CI assembly factors)
(Editted to remove COX7AL and COX7AL2. COX7AL is a CIV subunit and COX7AL2 is more similar to mammalian COX7A1 than COX7AL2, so more likely to be CIV SU.)

Assembly GO term annotation FBgn SYMBOL NAME Human Ortholog (best) S.Pombe Fly ortholog on PomBase list
mitochondrial respiratory chain complex I assembly FBgn0086712 Egm Enigma ACAD8|ACAD9*|ACADL (ACAD9 is the Mitochondrial Complex I Assembly complex component) no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0034578 CG15653 - COA1 coa1 SPBC16E9.03c
mitochondrial respiratory chain complex I assembly FBgn0051913 CG31913 - COA1 coa1 SPBC16E9.03c
mitochondrial respiratory chain complex I assembly FBgn0037018 Dmac2 Distal membrane arm assembly component 2 DMAC2 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0028436 ECSIT ECSIT ECSIT no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0033093 CG3270 - FOXRED1 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0002021 l(2)37Bb lethal (2) 37Bb FOXRED1 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0259832 CG34229 - LYRM2 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0039689 NdufAF1 NADH:ubiquinone oxidoreductase complex assembly factor 1 NDUFAF1 SPAC25A8.03c SPAC25A8.03c
mitochondrial respiratory chain complex I assembly FBgn0263051 CG43346 - NDUFAF2 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0034919 NdufAF3 NADH:ubiquinone oxidoreductase complex assembly factor 3 NDUFAF3 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0037777 NdufAF4 NADH:ubiquinone oxidoreductase complex assembly factor 4 NDUFAF4 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0033891 CG8067 - NDUFAF5 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0030352 sicily severe impairment of CI with lengthened youth NDUFAF6 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0037880 CG17726 - NDUFAF7 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0259726 CG42380 - NDUFAF8 (only called by eggNOG, Panther) no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0032986 Nubpl NUBP iron-sulfur cluster assembly factor, mitochondrial NUBPL No ortholog found (SPAC637.08 is similar to NUBPL but NUBP1 as cytosolic Fe-S protein is the ortholog for SPAC637.08 than NUBPL) no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0010340 Timmdc1 Translocase of inner mitochondrial membrane domain containing 1 TIMMDC1 no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0032033 Tmem126 Transmembrane protein 126 TMEM126A|TMEM126B* (TMEM126B is a member of the Mitochondrial Complex I Assembly complex, TMEM126A is not but is involved in the assembly of the ND4 distal membrane module of complex I) no orthologs found no orthologs found
mitochondrial respiratory chain complex I assembly FBgn0033808 Tmem186 Transmembrane protein 186 TMEM186 no orthologs found no orthologs found
mitochondrial respiratory chain complex II assembly FBgn0033274 CG14757 - SDHAF2 emi5 SPAC12B10.06c
mitochondrial respiratory chain complex II assembly FBgn0033523 CG12895 - SDHAF2 emi5 SPAC12B10.06c
mitochondrial respiratory chain complex II assembly FBgn0038437 Sdhaf3 Succinate dehydrogenase assembly factor 3 SDHAF3 SPBP23A10.03c SPBP23A10.03c
mitochondrial respiratory chain complex II assembly FBgn0031971 Sirup Starvation-upregulated protein SDHAF4 SPBC26H8.16 SPBC26H8.16
mitochondrial respiratory chain complex II assembly FBgn0028844 CG15283 - SDHAF4 SPBC26H8.16 SPBC26H8.16
mitochondrial respiratory chain complex III assembly FBgn0032195 Bcs1 Bcs1 chaperone BCS1L SPAC644.07 SPAC644.07
mitochondrial respiratory chain complex III assembly FBgn0050373 CG30373 - C16orf91 no orthologs found no orthologs found
mitochondrial respiratory chain complex III assembly FBgn0083986 CG34150 - LYRM7 new18 SPBC30D10.21
mitochondrial respiratory chain complex III assembly FBgn0032744 Ttc19 Tetratricopeptide repeat domain 19 TTC19 mgr3 SPAP27G11.02
mitochondrial respiratory chain complex III assembly FBgn0035722 Uqcc1 Ubiquinol-cytochrome c reductase complex assembly factor 1 UQCC1 cbp3 SPCC4B3.17
mitochondrial respiratory chain complex III assembly FBgn0287589 CG46459 - UQCC2 cbp6 (via sc orthology) SPBC947.14c
mitochondrial respiratory chain complex III assembly FBgn0040715 CG15386 - UQCC3 (via InterPro; no orthologs found via blastp/DIOPT hit) cbp4 (no orthologs found via DIOPT; but is UQCC3= Cbp4 orthology via text in PMID:30030361) SPBC27B12.14
mitochondrial respiratory chain complex III assembly FBgn0052736 sloth1 sloth 1 UQCC5 no orthologs found no orthologs found
mitochondrial respiratory chain complex III assembly FBgn0259204 sloth2 sloth 2 UQCC6 no orthologs found no orthologs found
mitochondrial cytochrome c oxidase assembly FBgn0266708 Cep89 Centrosomal protein 89kDa CEP89 no orthologs found no orthologs found
mitochondrial cytochrome c oxidase assembly FBgn0040634 CG4186 - CHCHD7 no orthologs found (Cox23 in s.c) no orthologs found
mitochondrial cytochrome c oxidase assembly FBgn0032595 CG17996 - CMC1 cmc2 SPCC4B3.20
mitochondrial cytochrome c oxidase assembly FBgn0259721 CG42375 - CMC2 cmc1 SPBC21D10.07
mitochondrial cytochrome c oxidase assembly FBgn0034578 CG15653 - COA1 coa1 SPBC16E9.03c
mitochondrial cytochrome c oxidase assembly FBgn0051913 CG31913 - COA1 coa1 SPBC16E9.03c
mitochondrial cytochrome c oxidase assembly FBgn0261353 Ccdc56 Coiled-coil domain containing 56 COA3 tam3 SPAC1B3.21
mitochondrial cytochrome c oxidase assembly FBgn0265083 CG44194 - COA4 coa4 SPCC550.01c
mitochondrial cytochrome c oxidase assembly FBgn0040751 CG13018 - COA5 coa5 SPBC16A3.16
mitochondrial cytochrome c oxidase assembly FBgn0259722 CG42376 - COA6 new16 SPBC24C6.13
mitochondrial cytochrome c oxidase assembly FBgn0039965 Coa7 Cytochrome c oxidase assembly factor 7 COA7 No ortholog found no orthologs found
mitochondrial cytochrome c oxidase assembly FBgn0051648 Cox11 Cytochrome c oxidase copper chaperone COX11 COX11 cox1101/cox1102 SPAC1420.04c|SPAC19B12.13
mitochondrial cytochrome c oxidase assembly FBgn0040899 CG17776   COX14 cox14 SPCC757.15
mitochondrial cytochrome c oxidase assembly FBgn0011476 l(3)neo43 lethal (3) neo43 COX16 cox16 SPAC1486.08
mitochondrial cytochrome c oxidase assembly FBgn0030610 Cox17 Cytochrome c oxidase copper chaperone COX17 COX17 cox17 SPBC26H8.14c
mitochondrial cytochrome c oxidase assembly FBgn0035960 CG4942 - COX18 cox18 SPCC1442.15c
mitochondrial cytochrome c oxidase assembly FBgn0260222 CG42496 - COX19 cox19 SPCC1672.04c
mitochondrial cytochrome c oxidase assembly FBgn0002354 l(3)87Df lethal (3) 87Df COX20 cox20 SPBC25H2.18
mitochondrial cytochrome c oxidase assembly FBgn0034248 CG14483 - PET100 No ortholog found no orthologs found
mitochondrial cytochrome c oxidase assembly FBgn0031247 CG11562 - PET117 pet117 SPBC1604.25
mitochondrial cytochrome c oxidase assembly FBgn0262467 Scox Synthesis of cytochrome c oxidase SCO1/SCO2 sco1 SPBC119.06
mitochondrial cytochrome c oxidase assembly FBgn0085271 CG34242 - SMIM20 No ortholog found no orthologs found
mitochondrial cytochrome c oxidase assembly FBgn0029117 Surf1 Surfeit 1 SURF1 shy1 SPBC1215.01
mitochondrial cytochrome c oxidase assembly FBgn0033547 CG12935 - TMEM223 SPBP22H7.04 SPBP22H7.04
mitochondrial proton-transporting ATP synthase complex assembly FBgn0032644 CG5131 - ATP23 atp23 SPCC320.12
mitochondrial proton-transporting ATP synthase complex assembly FBgn0022344 CG10340 - ATPAF1 atp11 SPAC3A12.12
mitochondrial proton-transporting ATP synthase complex assembly FBgn0021856 l(2)k14505 lethal (2) k14505 ATPAF2 atp12 SPAC9.12c
mitochondrial proton-transporting ATP synthase complex assembly FBgn0083953 CG34117 - FMC1 fmc1 SPAC1486.11
mitochondrial proton-transporting ATP synthase complex assembly FBgn0030311 CG11699 - TMEM242 no orthologs found no orthologs found
mitochondrial proton-transporting ATP synthase complex assembly FBgn0035805 Tmem70 Transmembrane protein 70 TMEM70 SPAC1834.10c (only called by orthoMCL, Panther) SPAC1834.10c
mitochondrial respirasome assembly FBgn0037890 CG17734 - HIGD1A|HIGD1B|HIGD1C rcf1 SPAC25B8.07c
mitochondrial respirasome assembly FBgn0033519 CG11825 - HIGD1A|HIGD1C rcf1 SPAC25B8.07c
mitochondrial respirasome assembly FBgn0030743 CG9921 - HIGD2A|HIGD2B rcf1 SPAC25B8.07c
mitochondrial respirasome assembly FBgn0033209 CG12107 - RAB5IF no orthologs found no orthologs found

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ValWood commented Nov 7, 2024 •
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We do have bona fida orthologs of

COA1 (coa1 https://www.pombase.org/gene/SPBC16E9.03c

NDUFAF1 https://www.pombase.org/gene/SPAC9E9.15

TTC19 https://www.pombase.org/gene/SPAP27G11.02

These don't get picked up by the ortholog predictors but are clear from directed searches using Jackhmmer/FoldSeek/Pfam-N etc
I will look for the other missing ones but Jackhmmer has been temporarily decommissioned by EBI

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hattrill commented Nov 7, 2024

Also, looks like we don't have an ortholog for SDHAF1 (Human), which is SPAC664.12c in Pombe

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ValWood commented Nov 7, 2024 •
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FIXES

  • PomBAse afg1 mitochondrial cytochrome c oxidase assembly NAS deleted
  • UniProtKB:Q8WV93 | AFG1L GO:0006123 mitochondrial electron transport, cytochrome c to oxygen looks indirect disputed by helen =>"GO:0141164 mitochondrial protein quality control"
  • PomBAse thi2 | GO:0033108 | mitochondrial respiratory chain complex assembly | ISO with S000003376 deleted

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ValWood commented Nov 8, 2024

Woah, https://www.uniprot.org/uniprotkb/Q96E52/entry is over-annotated
did you query any of the others?

CC @Antonialock

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hattrill commented Nov 8, 2024

Disputed some assembly annotations to human ones...and a special request for a big batch from PMID:27626371

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hattrill commented Nov 8, 2024 •
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image

Do you think mss51 should be cytochrome c oxidase assembly factor or upstream of it - I am not sure. Could model it either way depending - in this fig they make it a preassembly step.

Don't have a good ortholog for thi2, but not so sure it is an assembly factor. SGD summary for THI4 "Thiazole synthase; abundant protein involved in the formation of the thiazole moiety of thiamine during thiamine biosynthesis; acts more as a co-substrate rather than an enzyme by providing the sulphur source for thiazole formation; undergoes a single turnover only; required for mitochondrial genome stability in response to DNA damaging agents. DELETED!

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ValWood commented Nov 8, 2024

INteresting.

  1. In S. cerevisiae Mss51 is supposedly "translational"
  2. In pombe it is supposedly "post translations" https://www.pombase.org/reference/PMID:22349564
    (so maybe it does both but is masked by other factors, or maybe parts of both are outdated)
  3. Ssc1 is also interesting. We have it as mitochondrial (2Fe-2S) cluster assembly chaperone Ssc1, and being part of the
    PAM complex, Tim23 associated import motor
    but nothing about a role in respiratory complex assembly

@pombase pombase deleted a comment from hattrill Nov 14, 2024
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ValWood commented Nov 14, 2024

Parking this here so I don't forget:

__COX assembly

@ValWood ValWood changed the title respiratory complex/ respirasome assembly respiratory complex assembly/ respirasome assembly Nov 15, 2024
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