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BMRB: 25443

2MY3

Snu17p-Pml1p structure intermediate during RES complex assembly

Authors

Wysoczanski, P., Becker, S., Zweckstetter, M.

Assembly

Heterodimeric complex composed of Snu17p/Ist3p(25-138) and Pml1p(22-42)

Entity

1. Snu17p (polymer, Thiol state: not present), 118 monomers, 13478.84 Da Detail

GAMGNEYKDN AYIYIGNLNR ELTEGDILTV FSEYGVPVDV ILSRDENTGE SQGFAYLKYE DQRSTILAVD NLNGFKIGGR ALKIDHTFYR PKRSLQKYYE AVKEELDRDI VSKNNAEK

2. Pml1p (polymer, Thiol state: not present), 23 monomers, 2513.772 Da Detail

GSKSQYIDIM PDFSPSGLLE LES

Total weight

15992.611 Da

Max. entity weight

13478.84 Da

Source organism

Saccharomyces cerevisiae S288C

Exptl. method

Refine. method

simulated annealing

Data set

assigned_chemical_shifts

Chem. Shift Complete

Sequence coverage: 97.9 %, Completeness: 90.9 %, Completeness (bb): 91.5 % Detail

Polymer type: polypeptide(L)

	Total	¹H	¹³C	¹⁵N
All	90.9 % (1500 of 1650)	90.7 % (781 of 861)	91.1 % (582 of 639)	91.3 % (137 of 150)
Backbone	91.5 % (767 of 838)	92.1 % (267 of 290)	90.3 % (371 of 411)	94.2 % (129 of 137)
Sidechain	91.0 % (856 of 941)	90.0 % (514 of 571)	93.6 % (334 of 357)	61.5 % (8 of 13)
Aromatic	94.0 % (126 of 134)	94.0 % (63 of 67)	94.0 % (63 of 67)
Methyl	96.5 % (139 of 144)	95.8 % (69 of 72)	97.2 % (70 of 72)

1. Snu17p

GAMGNEYKDN AYIYIGNLNR ELTEGDILTV FSEYGVPVDV ILSRDENTGE SQGFAYLKYE DQRSTILAVD NLNGFKIGGR ALKIDHTFYR PKRSLQKYYE AVKEELDRDI VSKNNAEK

2. Pml1p

GSKSQYIDIM PDFSPSGLLE LES

Show sample condition

Sample #1

Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

Sample #2

Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
8	sodium phosphate	natural abundance		25 mM
9	sodium chloride	natural abundance		250 mM
10	Snu17p	natural abundance		1.0 ~ 1.2 mM
11	Pml1p	[U-13C; U-15N]		0.8 ~ 1.0 mM
12	sodium azide	natural abundance		1 mM
13	D2O	natural abundance		10 %
14	H2O	natural abundance		90 %

Protein Blocks Logo

Calculated from 20 models in PDB: 2MY3, Strand ID: A, B Detail

Release date

2015-08-09

Citation

Structures of intermediates during RES complex assembly
Wysoczanski, P., Becker, S., Zweckstetter, M.
Sci. Rep. (2015), 5, 12545-12545, PubMed 26212312 , DOI 10.1038/srep12545 , Abstract

Related entities 1. Snu17p, : 15 entities Detail

Related entities 2. Pml1p, : 1 : 1 : 5 : 1 entities Detail

Interaction partners 2. Pml1p, : 9 interactors Detail

More details for 9 interactors identified by 5 citations

Experiments performed 21 experiments Detail

1 2D 1H-15N HSQC

Instrument

Bruker Avance - 900 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

2 2D 1H-15N HSQC

Instrument

Bruker Avance - 900 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
8	sodium phosphate	natural abundance		25 mM
9	sodium chloride	natural abundance		250 mM
10	Snu17p	natural abundance		1.0 ~ 1.2 mM
11	Pml1p	[U-13C; U-15N]		0.8 ~ 1.0 mM
12	sodium azide	natural abundance		1 mM
13	D2O	natural abundance		10 %
14	H2O	natural abundance		90 %

3 2D 1H-15N HSQC

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

4 2D 1H-15N HSQC

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
8	sodium phosphate	natural abundance		25 mM
9	sodium chloride	natural abundance		250 mM
10	Snu17p	natural abundance		1.0 ~ 1.2 mM
11	Pml1p	[U-13C; U-15N]		0.8 ~ 1.0 mM
12	sodium azide	natural abundance		1 mM
13	D2O	natural abundance		10 %
14	H2O	natural abundance		90 %

5 2D 1H-15N HSQC

Instrument

Bruker Avance - 700 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

6 2D 1H-15N HSQC

Instrument

Bruker Avance - 700 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
8	sodium phosphate	natural abundance		25 mM
9	sodium chloride	natural abundance		250 mM
10	Snu17p	natural abundance		1.0 ~ 1.2 mM
11	Pml1p	[U-13C; U-15N]		0.8 ~ 1.0 mM
12	sodium azide	natural abundance		1 mM
13	D2O	natural abundance		10 %
14	H2O	natural abundance		90 %

7 2D 1H-15N HSQC

Instrument

Bruker Avance - 600 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

8 2D 1H-15N HSQC

Instrument

Bruker Avance - 600 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
8	sodium phosphate	natural abundance		25 mM
9	sodium chloride	natural abundance		250 mM
10	Snu17p	natural abundance		1.0 ~ 1.2 mM
11	Pml1p	[U-13C; U-15N]		0.8 ~ 1.0 mM
12	sodium azide	natural abundance		1 mM
13	D2O	natural abundance		10 %
14	H2O	natural abundance		90 %

9 2D 1H-13C HSQC aliphatic

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

10 3D 1H-13C NOESY aromatic

Instrument

Bruker Avance - 900 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

11 3D 1H-13C NOESY aromatic

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
8	sodium phosphate	natural abundance		25 mM
9	sodium chloride	natural abundance		250 mM
10	Snu17p	natural abundance		1.0 ~ 1.2 mM
11	Pml1p	[U-13C; U-15N]		0.8 ~ 1.0 mM
12	sodium azide	natural abundance		1 mM
13	D2O	natural abundance		10 %
14	H2O	natural abundance		90 %

12 3D 1H-13C NOESY aliphatic

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
8	sodium phosphate	natural abundance		25 mM
9	sodium chloride	natural abundance		250 mM
10	Snu17p	natural abundance		1.0 ~ 1.2 mM
11	Pml1p	[U-13C; U-15N]		0.8 ~ 1.0 mM
12	sodium azide	natural abundance		1 mM
13	D2O	natural abundance		10 %
14	H2O	natural abundance		90 %

13 3D HNCO

Instrument

Bruker Avance - 600 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

14 3D HNCA

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

15 3D HNCA

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
8	sodium phosphate	natural abundance		25 mM
9	sodium chloride	natural abundance		250 mM
10	Snu17p	natural abundance		1.0 ~ 1.2 mM
11	Pml1p	[U-13C; U-15N]		0.8 ~ 1.0 mM
12	sodium azide	natural abundance		1 mM
13	D2O	natural abundance		10 %
14	H2O	natural abundance		90 %

16 3D HN(CO)CA

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

17 3D HN(CO)CA

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
8	sodium phosphate	natural abundance		25 mM
9	sodium chloride	natural abundance		250 mM
10	Snu17p	natural abundance		1.0 ~ 1.2 mM
11	Pml1p	[U-13C; U-15N]		0.8 ~ 1.0 mM
12	sodium azide	natural abundance		1 mM
13	D2O	natural abundance		10 %
14	H2O	natural abundance		90 %

18 3D HCCH-TOCSY

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

19 3D 1H-15N NOESY

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

20 3D 1H-15N NOESY

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
8	sodium phosphate	natural abundance		25 mM
9	sodium chloride	natural abundance		250 mM
10	Snu17p	natural abundance		1.0 ~ 1.2 mM
11	Pml1p	[U-13C; U-15N]		0.8 ~ 1.0 mM
12	sodium azide	natural abundance		1 mM
13	D2O	natural abundance		10 %
14	H2O	natural abundance		90 %

21 HBCBCGCDHD

Instrument

Bruker Avance - 800 MHz HCN cryoprobe

Sample

State isotropic, Solvent system 90% H2O/10% D2O, Pressure 1 atm, Temperature 308 K, pH 6.8, Details 160-180 uL, 3 mm tube

#	Name	Isotope labeling	Type	Concentration
1	sodium phosphate	natural abundance		25 mM
2	sodium chloride	natural abundance		250 mM
3	Snu17p	[U-13C; U-15N]		0.8 ~ 1.0 mM
4	Pml1p_Fmoc	natural abundance		1.2 ~ 1.5 mM
5	sodium azide	natural abundance		1 mM
6	D2O	natural abundance		10 %
7	H2O	natural abundance		90 %

NMR combined restraints 5 contents Detail

Properties

Input source #1: NMR data (NEF) - Assigned chemical shifts, Distance restraints, Dihedral angle restraints	combined_25443_2my3.nef
Input source #2: Coordindates	2my3.cif
Diamagnetism of the molecular assembly	True (excluding Oxygen atoms)
Whether the assembly has a disulfide bond	None
Whether the assembly has a other bond	None
Whether the assembly contains a cyclic polymer	None
Overall data processing status	Warning

Disulfide bonds

None

Other bonds (neither disulfide, covalent nor hydrogen bonds, e.g. Zinc–sulphur bond)

None

Non-standard residues

None

Sequence alignments

Entity_assembly_ID (NMR): A vs Auth_asym_ID (model): A

--------10--------20--------30--------40--------50--------60--------70--------80--------90-------100
GAMGNEYKDNAYIYIGNLNRELTEGDILTVFSEYGVPVDVILSRDENTGESQGFAYLKYEDQRSTILAVDNLNGFKIGGRALKIDHTFYRPKRSLQKYYE
||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
GAMGNEYKDNAYIYIGNLNRELTEGDILTVFSEYGVPVDVILSRDENTGESQGFAYLKYEDQRSTILAVDNLNGFKIGGRALKIDHTFYRPKRSLQKYYE

-------110--------
AVKEELDRDIVSKNNAEK
||||||||||||||||||
AVKEELDRDIVSKNNAEK

Entity_assembly_ID (NMR): B vs Auth_asym_ID (model): B

200-----210-------220--
GSKSQYIDIMPDFSPSGLLELES
|||||||||||||||||||||||
GSKSQYIDIMPDFSPSGLLELES
--------10--------20---

Chain assignments

Entity_assembly_ID (NMR)	Auth_asym_ID (model)	Length	Unmapped	Conflict	Sequence coverage (%)
A	A	118	0	0	100.0
B	B	23	0	0	100.0

Content subtype: combined_25443_2my3.nef

#	Content subtype	Saveframe	Status	# of rows (sets)	Experiment type	Sequence coverage (%)
1	Assigned chemical shifts	SHIFT_LIST	Warning	1557		99.2 (chain: A, length: 118), 91.3 (chain: B, length: 23)
1	Distance restraints	CNS/XPLOR_distance_constraints_3	OK	3088 (1)	noe	98.3 (chain: A, length: 118), 87.0 (chain: B, length: 23)
2	Distance restraints	CNS/XPLOR_distance_constraints_4	OK	50 (1)	hbond	32.2 (chain: A, length: 118)
1	Dihedral angle restraints	CNS/XPLOR_dihedral_2	OK	230 (230)	.	93.2 (chain: A, length: 118), 91.3 (chain: B, length: 23)

Assigned chemical shifts

Saveframe: SHIFT_LIST
- Status: Warning
- Sequence coverage of experimental data
  - Entity_assembly_ID: A, Chain length: 118, Sequence coverage: 99.2%
  - Entity_assembly_ID: B, Chain length: 23, Sequence coverage: 91.3%
- Total number of assignments
  - ¹H chemical shifts: 850
  - ¹³C chemical shifts: 572
  - ¹⁵N chemical shifts: 135
- Completeness of assignments
  - Entity_assemble_ID: A
  - Entity_assemble_ID: B
- Peptide bond of PRO
- Tautomeric state of HIS
- Rotameric state of ILE , LEU, and VAL
- Histogram of normalized assigned chemical shifts
- Random coil index and derived parameters (S² and NMR RMSD)
  - Chain_ID: A
  - Chain_ID: B

Distance restraints

Saveframe: CNS/XPLOR_distance_constraints_3
- Status: OK
- Experiment type: noe
- Sequence coverage of experimental data
  - Entity_assembly_ID: A, Chain length: 118, Sequence coverage: 98.3%
  - Entity_assembly_ID: B, Chain length: 23, Sequence coverage: 87.0%
- Total number of restraints: 3088
- Weight of restraints: 1.0 (3088)
- Potential type of restraints: square-well-parabolic (3088)
- Range of distance target values: 2.25, 4.48 (Å)
- Histogram of distance target values
- Histogram of discrepancy in distance target values
- Distance restraints per residue
  - Chain_ID: A
  - Chain_ID: B
- Distance restraints on contact map
- Saveframe: CNS/XPLOR_distance_constraints_4
  - Status: OK
  - Experiment type: hbond
  - Sequence coverage of experimental data
    - Entity_assembly_ID: A, Chain length: 118, Sequence coverage: 32.2%
  - Total number of restraints: 50
  - Weight of restraints: 1.0 (50)
  - Potential type of restraints: square-well-parabolic (50)
  - Range of distance target values: 2.0, 3.0 (Å)
  - Histogram of distance target values
  - Distance restraints per residue
    - Chain_ID: A
    - Chain_ID: B
      None
  - Distance restraints on contact map

Dihedral angle restraints

Saveframe: CNS/XPLOR_dihedral_2
- Status: OK
- Experiment type: .
- Sequence coverage of experimental data
  - Entity_assembly_ID: A, Chain length: 118, Sequence coverage: 93.2%
  - Entity_assembly_ID: B, Chain length: 23, Sequence coverage: 91.3%
- Total number of restraints: 230
- Total number of restraints per polymer type: 230
- Weight of restraints: 1.0 (230)
- Potential type of restraints: square-well-parabolic (230)
- Plot of Phi-Psi angle restraints
- Dihedral angle restraints per residue
  - Chain_ID: A
  - Chain_ID: B

Keywords Ist3p, Pml1p, RES, RRM, Snu17p, cooperativity, heterodimer, spliceosome, splicing