Thermodynamic studies of the core histones: ionic strength and pH dependence of H2A-H2B dimer stability.


Abstract

The thermal stability of the core histone dimer H2A-H2B has been studied by high-sensitivity differential scanning calorimetry and circular dichroism spectroscopy. The unfolding transition temperature of the 28 kDa H2A-H2B dimer increases as a function of both the ionic strength of the solvent and the total protein concentration. At neutral pH and physiological ionic strength, the thermal denaturation is centered at about 50 degrees C with a corresponding enthalpy change of about 40 kcal/mol of 14 kDa monomer unit and an excess heat capacity of about 1.4 kcal/(K.mol) of 14 kDa monomer unit. The H2A-H2B dimer is stable mainly between pH 5.5 and 10.5. Below pH 4.0, the system is unfolded at all temperatures. The thermodynamic analysis is performed at low ionic strength where almost complete reversibility is attained, since higher salt conditions seem to promote aggregation and irreversibility of the transitions. Analysis of the data shows that at low ionic strength and pH values between 6.5 and 8.5, the H2A-H2B dimer behaves as a highly cooperative system, melting as a single unit without any detectable intermediates of dissociated, yet folded, H2A and H2B monomers. This is consistent with the observed protein concentration dependence of the midpoint of the thermal denaturation. The two-state unfolding process can be described by the general scheme AB-->2U, indicating that the individual H2A and H2B polypeptides are folded, stable entities only when complexed as the H2A-H2B dimer and that the major contribution to the stabilization of the dimer derives from the coupling between the H2A and H2B interfaces. Study holds ProTherm entries: 5382, 5383, 5384, 5385, 5386, 5387, 5388, 5389, 5390, 5391, 5392, 5393, 5394, 5395, 5396, 5397, 5398 Extra Details: ionic strength; pH dependence; protein concentration;,H2A-H2B dimer; heat capacity

Submission Details

ID: v3sYWqzr3

Submitter: Connie Wang

Submission Date: April 24, 2018, 8:30 p.m.

Version: 1

Publication Details
Karantza V;Baxevanis AD;Freire E;Moudrianakis EN,Biochemistry (1995) Thermodynamic studies of the core histones: ionic strength and pH dependence of H2A-H2B dimer stability. PMID:7727455
Additional Information

Study Summary

Number of data points 32
Proteins Histone H2B 1/2/3/4/6 ; Histone H2A-IV
Unique complexes 1
Assays/Quantities/Protocols Experimental Assay: Tm buffers:CAPS: 7.5 mM, ionic:NaCl: 100 mM, pH:11.5 ; Experimental Assay: Tm buffers:CAPS: 7.5 mM, ionic:NaCl: 100 mM, pH:10.5 ; Experimental Assay: Tm buffers:bicine: 10 mM, ionic:NaCl: 100 mM, pH:9.5 ; Experimental Assay: dHvH buffers:bicine: 10 mM, pH:9.5 ; Experimental Assay: Tm buffers:bicine: 10 mM, ionic:NaCl: 100 mM, pH:8.5 ; Experimental Assay: dHvH buffers:bicine: 10 mM, pH:8.5 ; Experimental Assay: Tm pH:7.5, ionic:NaCl: 100 mM, buffers:HEPES: 10 mM ; Experimental Assay: dHvH pH:7.5, buffers:HEPES: 10 mM ; Experimental Assay: Tm ionic:NaCl: 100 mM, buffers:HEPES: 10 mM, pH:6.5 ; Experimental Assay: dHvH buffers:HEPES: 10 mM, pH:6.5 ; Experimental Assay: Tm buffers:MES: 10 mM, pH:5.5, ionic:NaCl: 100 mM ; Experimental Assay: dHvH buffers:MES: 10 mM, pH:5.5 ; Experimental Assay: Tm pH:4.5, ionic:NaCl: 100 mM, buffers:MES: 10 mM ; Experimental Assay: Tm buffers:glycine: 10 mM, pH:4.0, ionic:NaCl: 100 mM ; Experimental Assay: Tm ionic:NaCl: 50 mM, pH:11.5, buffers:glycine: 10 mM ; Experimental Assay: Tm ionic:NaCl: 50 mM, pH:10.5, buffers:glycine: 10 mM ; Experimental Assay: dCp buffers:glycine: 10 mM, pH:9.5 ; Experimental Assay: dHcal buffers:glycine: 10 mM, pH:9.5 ; Experimental Assay: Tm ionic:NaCl: 50 mM, pH:9.5, buffers:glycine: 10 mM ; Experimental Assay: dCp buffers:glycylglycine: 10 mM, pH:8.5 ; Experimental Assay: dHcal buffers:glycylglycine: 10 mM, pH:8.5 ; Experimental Assay: Tm ionic:NaCl: 50 mM, pH:8.5, buffers:glycylglycine: 10 mM ; Experimental Assay: dCp pH:7.5, buffers:imidazole: 10 mM ; Experimental Assay: dHcal pH:7.5, buffers:imidazole: 10 mM ; Experimental Assay: Tm pH:7.5, buffers:imidazole: 10 mM, ionic:NaCl: 50 mM ; Experimental Assay: dCp buffers:imidazole: 10 mM, pH:6.5 ; Experimental Assay: dHcal buffers:imidazole: 10 mM, pH:6.5 ; Experimental Assay: Tm buffers:imidazole: 10 mM, pH:6.5, ionic:NaCl: 50 mM ; Experimental Assay: dCp pH:5.5, buffers:imidazole: 10 mM ; Experimental Assay: dHcal pH:5.5, buffers:imidazole: 10 mM ; Experimental Assay: Tm buffers:imidazole: 10 mM, pH:5.5, ionic:NaCl: 50 mM ; Experimental Assay: Tm pH:4.5, buffers:glycylglycine: 10 mM, ionic:NaCl: 50 mM
Libraries Mutations for sequence A:KSRSSRAGLQFPVGRVHRLLRKGNYAERVGAGAPVYLAAVLEYLTAEILELAGNAARDNKKTRIIPRHLQLAIRNDEELNKLLGKVTIAQGGVLP/B:SYSIYVYKVLKQVHPDTGISSKAMGSMNSFVNDIFERIAGLASRLAHYNKRSTITSREIQTAVRLLLPGELAKHAVSEGTKAVTKHTSSK/C:PGTVALREIRRYQKSTELLIRKLPFQRLVREIAQDFKTDLRFQSSAVMALQEASEAYLVGLFEDTNLCAIHAKRVTIMPKDIELARRIRGERA/D:QGITKPAIRRLARRGGVKRISGLIYEETRGVLKVFLENVIRDAVTYTEHAKRKTVTAMDVVYALKRQGRTLYGFGG

Structure view and single mutant data analysis

Study data

No weblogo for data of varying length.
Colors: D E R H K S T N Q A V I L M F Y W C G P
 

Data Distribution

Studies with similar sequences (approximate matches)

Correlation with other assays (exact sequence matches)


Relevant PDB Entries

Structure ID Release Date Resolution Structure Title
2ARO 2005-08-20T00:00:00+0000 2.1 Crystal Structure Of The Native Histone Octamer To 2.1 Angstrom Resolution, Crystalised In The Presence Of S-Nitrosoglutathione
4EDU 2012-03-27T00:00:00+0000 2.58 The MBT repeats of human SCML2 in a complex with histone H2A peptide
6K60 2019-05-31T00:00:00+0000 3.15 Structural and functional basis for HLA-G isoform recognition of immune checkpoint receptor LILRBs
6KVD 2019-09-04T00:00:00+0000 2.21 Crystal structure of human nucleosome containing H2A.J
1EQZ 2000-04-06T00:00:00+0000 2.5 X-RAY STRUCTURE OF THE NUCLEOSOME CORE PARTICLE AT 2.5 A RESOLUTION
1HIO 1991-09-19T00:00:00+0000 3.1 HISTONE OCTAMER (CHICKEN), CHROMOSOMAL PROTEIN, ALPHA CARBONS ONLY
1HQ3 2000-12-14T00:00:00+0000 2.15 CRYSTAL STRUCTURE OF THE HISTONE-CORE-OCTAMER IN KCL/PHOSPHATE
1TZY 2004-07-12T00:00:00+0000 1.9 Crystal Structure of the Core-Histone Octamer to 1.90 Angstrom Resolution
2ARO 2005-08-20T00:00:00+0000 2.1 Crystal Structure Of The Native Histone Octamer To 2.1 Angstrom Resolution, Crystalised In The Presence Of S-Nitrosoglutathione
2HIO 1999-06-15T00:00:00+0000 3.1 HISTONE OCTAMER (CHICKEN), CHROMOSOMAL PROTEIN

Relevant UniProtKB Entries

Percent Identity Matching Chains Protein Accession Entry Name
100.0 Histone H2B 1/2/3/4/6 P0C1H3 H2B1_CHICK
100.0 Histone H2B 1/2/3/4/6 Q16778 H2B2E_HUMAN
96.8 Histone H2B 1/2/3/4/6 P0C1H4 H2B5_CHICK
99.2 Histone H2B 1/2/3/4/6 P62807 H2B1C_HUMAN
99.2 Histone H2B 1/2/3/4/6 Q6ZWY9 H2B1C_MOUSE
99.2 Histone H2B 1/2/3/4/6 Q64478 H2B1H_MOUSE
99.2 Histone H2B 1/2/3/4/6 P62808 H2B1_BOVIN
99.2 Histone H2B 1/2/3/4/6 Q5R893 H2B1_PONAB
96.7 B Histone H2A-IV P33778 H2B1B_HUMAN
99.2 Histone H2B 1/2/3/4/6 P06899 H2B1J_HUMAN
99.2 Histone H2B 1/2/3/4/6 P23527 H2B1O_HUMAN
98.4 Histone H2B 1/2/3/4/6 Q93079 H2B1H_HUMAN
98.4 Histone H2B 1/2/3/4/6 O60814 H2B1K_HUMAN
98.4 Histone H2B 1/2/3/4/6 Q99877 H2B1N_HUMAN
98.4 Histone H2B 1/2/3/4/6 P58876 H2B1D_HUMAN
98.4 Histone H2B 1/2/3/4/6 P10854 H2B1M_MOUSE
99.2 Histone H2B 1/2/3/4/6 P14001 H2B_CAIMO
98.4 Histone H2B 1/2/3/4/6 P10853 H2B1F_MOUSE
93.3 B Histone H2A-IV P06900 H2B12_XENLA
97.6 Histone H2B 1/2/3/4/6 Q2PFX4 H2B1K_MACFA
96.0 Histone H2B 1/2/3/4/6 Q99879 H2B1M_HUMAN
96.8 Histone H2B 1/2/3/4/6 P57053 H2BFS_HUMAN
97.6 Histone H2B 1/2/3/4/6 Q64475 H2B1B_MOUSE
97.6 Histone H2B 1/2/3/4/6 Q8CGP1 H2B1K_MOUSE
97.6 Histone H2B 1/2/3/4/6 Q32L48 H2B1N_BOVIN
92.2 B Histone H2A-IV P02281 H2B11_XENLA
96.0 Histone H2B 1/2/3/4/6 Q8CGP2 H2B1P_MOUSE
97.6 Histone H2B 1/2/3/4/6 Q99880 H2B1L_HUMAN
96.0 Histone H2B 1/2/3/4/6 Q64524 H2B2E_MOUSE
94.4 Histone H2B 1/2/3/4/6 Q75VN4 H2B_RHASC
96.6 Histone H2B 1/2/3/4/6 P83863 H2B_PENVA
96.6 Histone H2B 1/2/3/4/6 P19374 H2B_PLADU
95.5 Histone H2B 1/2/3/4/6 P35068 H2B1_TIGCA
95.5 Histone H2B 1/2/3/4/6 P35069 H2B3_TIGCA
95.5 Histone H2B 1/2/3/4/6 Q27442 H2B_ANOGA
95.5 Histone H2B 1/2/3/4/6 P17271 H2B_DROHY
95.5 Histone H2B 1/2/3/4/6 Q76FD7 H2B_DROSE
95.5 Histone H2B 1/2/3/4/6 Q8I1N0 H2B_DROYA
94.4 Histone H2B 1/2/3/4/6 P27326 H2B_URECA
93.3 Histone H2B 1/2/3/4/6 P21897 H2B_CHITH
91.0 Histone H2B 1/2/3/4/6 P07794 H2BL1_PSAMI
96.6 Histone H2B 1/2/3/4/6 P59781 H2B_DROER
96.6 Histone H2B 1/2/3/4/6 Q76FE5 H2B_DROMA
96.6 Histone H2B 1/2/3/4/6 P02283 H2B_DROME
96.6 Histone H2B 1/2/3/4/6 Q76FE9 H2B_DROOR
96.6 Histone H2B 1/2/3/4/6 P59782 H2B_DROSI
96.6 Histone H2B 1/2/3/4/6 Q76FF3 H2B_DROTE
95.4 Histone H2B 1/2/3/4/6 P02284 H2B_PATGR
95.3 Histone H2B 1/2/3/4/6 P30757 H2B_SIPNU
92.9 Histone H2B 1/2/3/4/6 Q7M4G7 H2B_PATPE
91.4 Histone H2B 1/2/3/4/6 Q6DN03 H2B2C_HUMAN
91.4 Histone H2B 1/2/3/4/6 Q6DRA6 H2B2D_HUMAN
100.0 A Histone H2A-IV P02262 H2A1_RAT
100.0 A Histone H2A-IV P02263 H2A4_CHICK
100.0 A Histone H2A-IV Q3ZBX9 H2AJ_BOVIN
100.0 A Histone H2A-IV Q9BTM1 H2AJ_HUMAN
100.0 A Histone H2A-IV Q4R3X5 H2AJ_MACFA
100.0 A Histone H2A-IV P13912 H2A_CAIMO
98.9 A Histone H2A-IV Q6FI13 H2A2A_HUMAN
98.9 A Histone H2A-IV Q6GSS7 H2A2A_MOUSE
98.9 A Histone H2A-IV P0CC09 H2A2A_RAT
98.9 A Histone H2A-IV A1A4R1 H2A2C_BOVIN
98.9 A Histone H2A-IV Q16777 H2A2C_HUMAN
98.9 A Histone H2A-IV Q64523 H2A2C_MOUSE
98.9 A Histone H2A-IV P70082 H2AJ_CHICK
98.9 A Histone H2A-IV Q93077 H2A1C_HUMAN
98.9 A Histone H2A-IV P0C169 H2A1C_RAT
98.9 A Histone H2A-IV Q96KK5 H2A1H_HUMAN
98.9 A Histone H2A-IV Q99878 H2A1J_HUMAN
98.9 A Histone H2A-IV P0C0S9 H2A1_BOVIN
98.9 A Histone H2A-IV P0C0S8 H2A1_HUMAN
98.9 A Histone H2A-IV Q8R1M2 H2AJ_MOUSE
98.9 A Histone H2A-IV A9UMV8 H2AJ_RAT
98.9 A Histone H2A-IV P35062 H2A3_CHICK
97.9 A Histone H2A-IV P20671 H2A1D_HUMAN
97.9 A Histone H2A-IV P0C170 H2A1E_RAT
97.9 A Histone H2A-IV Q7L7L0 H2A3_HUMAN
97.9 A Histone H2A-IV Q8BFU2 H2A3_MOUSE
97.9 A Histone H2A-IV Q4FZT6 H2A3_RAT
97.9 A Histone H2A-IV Q8IUE6 H2A2B_HUMAN
96.8 A Histone H2A-IV P04908 H2A1B_HUMAN
96.8 A Histone H2A-IV C0HKE1 H2A1B_MOUSE
96.8 A Histone H2A-IV C0HKE2 H2A1C_MOUSE
96.8 A Histone H2A-IV C0HKE3 H2A1D_MOUSE
96.8 A Histone H2A-IV C0HKE4 H2A1E_MOUSE
96.8 A Histone H2A-IV Q8CGP5 H2A1F_MOUSE
96.8 A Histone H2A-IV C0HKE5 H2A1G_MOUSE
96.8 A Histone H2A-IV Q8CGP6 H2A1H_MOUSE
96.8 A Histone H2A-IV C0HKE6 H2A1I_MOUSE
96.8 A Histone H2A-IV Q8CGP7 H2A1K_MOUSE
96.8 A Histone H2A-IV C0HKE7 H2A1N_MOUSE
96.8 A Histone H2A-IV C0HKE8 H2A1O_MOUSE
96.8 A Histone H2A-IV C0HKE9 H2A1P_MOUSE
96.8 A Histone H2A-IV Q64522 H2A2B_MOUSE
96.8 A Histone H2A-IV P06897 H2A1_XENLA
96.8 A Histone H2A-IV Q7ZUY3 H2AX_DANRE
96.8 A Histone H2A-IV P02264 H2A_ONCMY
95.8 A Histone H2A-IV Q96QV6 H2A1A_HUMAN
97.9 A Histone H2A-IV P16104 H2AX_HUMAN
97.9 A Histone H2A-IV P27661 H2AX_MOUSE
95.8 A Histone H2A-IV Q64598 H2A1F_RAT
94.7 A Histone H2A-IV Q00728 H2A4_RAT
94.7 A Histone H2A-IV Q6GM86 H2AX_XENLA
92.6 A Histone H2A-IV Q6PV61 H2A_PENVA
92.6 A Histone H2A-IV P35061 H2A_ACRFO
92.6 A Histone H2A-IV P84052 H2A_DROER
92.6 A Histone H2A-IV P84053 H2A_DROHY
92.6 A Histone H2A-IV P84051 H2A_DROME
92.6 A Histone H2A-IV P84054 H2A_DROSI
92.6 A Histone H2A-IV P84055 H2A_DROYA
92.6 A Histone H2A-IV P19178 H2A_PLADU
92.6 A Histone H2A-IV P84056 H2A_RHYAM
92.6 A Histone H2A-IV P84057 H2A_TIGCA
92.6 A Histone H2A-IV P13630 H2A_PARLI
93.7 A Histone H2A-IV A1CJ10 H2A_ASPCL
93.7 A Histone H2A-IV Q4WWC6 H2A_ASPFU
93.7 A Histone H2A-IV P0C953 H2A_ASPNC
93.7 A Histone H2A-IV P0C952 H2A_ASPNG
93.7 A Histone H2A-IV Q2U5A8 H2A_ASPOR
93.7 A Histone H2A-IV Q0CBD2 H2A_ASPTN
93.7 A Histone H2A-IV O74268 H2A_BOTFB
93.7 A Histone H2A-IV Q2HH37 H2A_CHAGB
93.7 A Histone H2A-IV P08844 H2A_EMENI
93.7 A Histone H2A-IV Q4HTT1 H2A_GIBZE
93.7 A Histone H2A-IV P0CT12 H2A_MAGO7
93.7 A Histone H2A-IV L7HZV6 H2A_MAGOY
93.7 A Histone H2A-IV A1D8G8 H2A_NEOFI
93.7 A Histone H2A-IV Q8X132 H2A_NEUCR
93.7 A Histone H2A-IV Q875B8 H2A_PODAS
91.6 A Histone H2A-IV P69142 H2AE_PSAMI
91.6 A Histone H2A-IV P69141 H2A_STRPU
93.7 A Histone H2A-IV P09589 H2A3_LYTPI
91.6 A Histone H2A-IV P04735 H2A1_PSAMI
90.5 A Histone H2A-IV P02268 H2A_SEPOF
91.6 A Histone H2A-IV P04909 H2A1_SCHPO
91.6 A Histone H2A-IV P04910 H2A2_SCHPO
90.5 A Histone H2A-IV P09588 H2A_CAEEL
92.5 A Histone H2A-IV P21896 H2A_CHITH
94.6 A Histone H2A-IV Q0U1A1 H2A_PHANO
92.5 A Histone H2A-IV Q07135 H2AO_CHITH
95.6 A Histone H2A-IV P69139 H2A3_PSAMI
95.6 A Histone H2A-IV P69140 H2A_PARAN
92.3 A Histone H2A-IV P02270 H2A_SIPNU
92.3 A Histone H2A-IV P27325 H2A_URECA
94.4 A Histone H2A-IV P07793 H2A4_PSAMI
93.3 A Histone H2A-IV Q6WV66 H2A_MYTCA
93.3 A Histone H2A-IV Q6WV69 H2A_MYTCH
93.3 A Histone H2A-IV Q8I0T3 H2A_MYTED
93.3 A Histone H2A-IV Q6WV88 H2A_MYTGA
91.2 A Histone H2A-IV Q4PEF9 H2A_USTMA
94.4 A Histone H2A-IV Q1E5N1 H2A_COCIM
93.4 A Histone H2A-IV P02269 H2A_ASTRU
92.2 A Histone H2A-IV P04736 H2A2_PSAMI
92.2 A Histone H2A-IV Q6WV67 H2A_MYTTR
93.2 A Histone H2A-IV P16886 H2AL_STRPU
95.1 A Histone H2A-IV P09590 H2A3_STRPU
94.6 A Histone H2A-IV P59890 H2A_HIPHI
96.0 A Histone H2A-IV P55897 H2A_BUFGR
98.9 C Histone H2A-IV P84227 H32_BOVIN
98.9 C Histone H2A-IV P84230 H32_CAIMO
98.9 C Histone H2A-IV P84229 H32_CHICK
98.9 C Histone H2A-IV Q4QRF4 H32_DANRE
98.9 C Histone H2A-IV Q71DI3 H32_HUMAN
98.9 C Histone H2A-IV P84231 H32_ICTBU
98.9 C Histone H2A-IV P84228 H32_MOUSE
98.9 C Histone H2A-IV Q6LBE8 H32_MUSPA
98.9 C Histone H2A-IV P84234 H32_ONCMY
98.9 C Histone H2A-IV P84232 H32_PORAF
98.9 C Histone H2A-IV P84233 H32_XENLA
98.9 C Histone H2A-IV Q28D37 H32_XENTR
98.9 C Histone H2A-IV P84238 H3_CHITH
98.9 C Histone H2A-IV P84236 H3_DROHY
98.9 C Histone H2A-IV P02299 H3_DROME
98.9 C Histone H2A-IV P84235 H3_PLADU
98.9 C Histone H2A-IV P84237 H3_TIGCA
98.9 C Histone H2A-IV P84239 H3_URECA
97.8 C Histone H2A-IV P22843 H3_ACRFO
97.8 C Histone H2A-IV Q64400 H32_CRILO
97.8 C Histone H2A-IV P69071 H3_DERIM
97.8 C Histone H2A-IV P69072 H3_LYTPI
97.8 C Histone H2A-IV P69073 H3_PARLI
97.8 C Histone H2A-IV P69074 H3_PISBR
97.8 C Histone H2A-IV P69075 H3_PISOC
97.8 C Histone H2A-IV P69076 H3_PSAMI
97.8 C Histone H2A-IV P69077 H3_PYCHE
97.8 C Histone H2A-IV P69078 H3_SOLST
97.8 C Histone H2A-IV P69079 H3_STRDR
97.8 C Histone H2A-IV P68432 H31_BOVIN
97.8 C Histone H2A-IV P68431 H31_HUMAN
97.8 C Histone H2A-IV P68433 H31_MOUSE
97.8 C Histone H2A-IV Q6LBF0 H31_MUSPA
97.8 C Histone H2A-IV Q6LED0 H31_RAT
96.8 C Histone H2A-IV P08903 H32_ENCAL
96.8 C Histone H2A-IV P02302 H3C_XENLA
95.7 C Histone H2A-IV P59226 H32_ARATH
95.7 C Histone H2A-IV Q6LBE3 H32_ASPOF
95.7 C Histone H2A-IV Q6LCK1 H32_BRANA
95.7 C Histone H2A-IV Q5MYA4 H32_CICIN
95.7 C Histone H2A-IV Q71T45 H32_EUPES
95.7 C Histone H2A-IV P69246 H32_MAIZE
95.7 C Histone H2A-IV P68429 H32_MEDSA
95.7 C Histone H2A-IV P68430 H32_ONOVI
95.7 C Histone H2A-IV A2Y533 H32_ORYSI
95.7 C Histone H2A-IV Q2RAD9 H32_ORYSJ
95.7 C Histone H2A-IV P68427 H32_PEA
95.7 C Histone H2A-IV P69248 H32_PETCR
95.7 C Histone H2A-IV Q76MV0 H32_TOBAC
95.7 C Histone H2A-IV P68428 H32_WHEAT
95.7 C Histone H2A-IV Q10453 H331_CAEEL
95.7 C Histone H2A-IV C0HL66 H33A_DROME
95.7 C Histone H2A-IV C0HL67 H33B_DROME
95.7 C Histone H2A-IV Q5E9F8 H33_BOVIN
95.7 C Histone H2A-IV P84247 H33_CHICK
95.7 C Histone H2A-IV Q6PI20 H33_DANRE
95.7 C Histone H2A-IV P84250 H33_DROHY
95.7 C Histone H2A-IV P84243 H33_HUMAN
95.7 C Histone H2A-IV P84244 H33_MOUSE
95.7 C Histone H2A-IV Q71LE2 H33_PIG
95.7 C Histone H2A-IV P84246 H33_RABIT
95.7 C Histone H2A-IV P84245 H33_RAT
95.7 C Histone H2A-IV P84248 H33_SPISO
95.7 C Histone H2A-IV Q6PI79 H33_XENLA
95.7 C Histone H2A-IV Q6P823 H33_XENTR
95.7 C Histone H2A-IV P02301 H3C_MOUSE
96.8 C Histone H2A-IV P06352 H3_STRPU
94.6 C Histone H2A-IV Q6LCW8 H32_CHLRE
95.7 C Histone H2A-IV P08898 H3_CAEEL
94.6 C Histone H2A-IV Q9U7D1 H3_MASBA
94.6 C Histone H2A-IV P08437 H3_VOLCA
94.6 C Histone H2A-IV P59169 H33_ARATH
94.6 C Histone H2A-IV Q6RUR1 H33_CAPAN
94.6 C Histone H2A-IV Q71V89 H33_GOSHI
94.6 C Histone H2A-IV Q3C2E5 H33_LOLMU
94.6 C Histone H2A-IV P69245 H33_LOLTE
94.6 C Histone H2A-IV P69244 H33_MEDSA
94.6 C Histone H2A-IV Q71U98 H33_ORYCO
94.6 C Histone H2A-IV A2XHJ3 H33_ORYSI
94.6 C Histone H2A-IV Q0JCT1 H33_ORYSJ
94.6 C Histone H2A-IV Q711T2 H33_PINPS
94.6 C Histone H2A-IV Q76N23 H33_TOBAC
94.6 C Histone H2A-IV Q71H73 H33_VITVI
94.6 C Histone H2A-IV Q16695 H31T_HUMAN
94.6 C Histone H2A-IV Q9U281 H332_CAEEL
94.6 C Histone H2A-IV Q8WSF1 H33_TRIPS
93.5 C Histone H2A-IV Q42681 H31_CHLRE
94.6 C Histone H2A-IV Q5RCC9 H33_PONAB
94.6 C Histone H2A-IV Q402E1 H32_LILLO
92.5 C Histone H2A-IV Q9LR02 H3L3_ARATH
92.5 C Histone H2A-IV Q27532 H33L2_CAEEL
92.5 C Histone H2A-IV A5PK61 H3C_BOVIN
92.5 C Histone H2A-IV Q9FKQ3 H3L5_ARATH
92.4 C Histone H2A-IV Q7XYZ0 H3_GRIJA
94.6 C Histone H2A-IV Q6NXT2 H3C_HUMAN
91.4 C Histone H2A-IV Q2Z2F4 H33B_LILLO
93.5 C Histone H2A-IV P50564 H33_CHLRE
91.4 C Histone H2A-IV Q402E2 H33A_LILLO
93.5 C Histone H2A-IV P06902 H3C_CAIMO
91.4 C Histone H2A-IV O15819 H33A_DICDI
91.4 C Histone H2A-IV Q9FX60 H3L1_ARATH
91.4 C Histone H2A-IV P90543 H3_EUPCR
91.4 C Histone H2A-IV Q9FXI7 H3L2_ARATH
90.3 C Histone H2A-IV Q55BN9 H33B_DICDI
92.4 C Histone H2A-IV Q4PB04 H32_USTMA
90.3 C Histone H2A-IV P0CO05 H3_CRYNB
90.3 C Histone H2A-IV P0CO04 H3_CRYNJ
90.3 C Histone H2A-IV A5DWE2 H31_LODEL
92.4 C Histone H2A-IV Q9HDN1 H3_MORAP
90.3 C Histone H2A-IV Q4P7J7 H31_USTMA
90.1 C Histone H2A-IV Q6C0C4 H3_YARLI
95.0 C Histone H2A-IV P06353 H33_HORVU
93.7 C Histone H2A-IV P81197 H33_STYLE
91.1 C Histone H2A-IV P81199 H35_STYLE
92.4 C Histone H2A-IV P81201 H37_STYLE
92.4 C Histone H2A-IV P81196 H39_STYLE
96.7 B Histone H2A-IV Q5RCP8 H2B2E_PONAB
96.7 B Histone H2A-IV Q9PSW9 H2B8_CHICK
95.6 B Histone H2A-IV Q64525 H2B2B_MOUSE
95.6 B Histone H2A-IV Q5QNW6 H2B2F_HUMAN
95.6 B Histone H2A-IV Q5BJA5 H2B1_DANRE
95.6 B Histone H2A-IV Q6PC60 H2B3_DANRE
94.4 B Histone H2A-IV Q2M2T1 H2B1K_BOVIN
95.6 B Histone H2A-IV P0C1H5 H2B7_CHICK
94.4 B Histone H2A-IV P69069 H2B_ONCMY
94.4 B Histone H2A-IV P69070 H2B_SALTR
94.4 B Histone H2A-IV Q9D2U9 H2B3A_MOUSE
94.4 B Histone H2A-IV Q8N257 H2B3B_HUMAN
94.4 B Histone H2A-IV Q8CGP0 H2B3B_MOUSE
93.3 B Histone H2A-IV Q00729 H2B1A_RAT
92.2 B Histone H2A-IV P70696 H2B1A_MOUSE
93.3 B Histone H2A-IV Q00715 H2B1_RAT
96.8 B Histone H2A-IV P02280 H2B_CRONI
100.0 D Histone H2A-IV P84048 H4_ACRAS
100.0 D Histone H2A-IV P35059 H4_ACRFO
100.0 D Histone H2A-IV P84047 H4_ASEAQ
100.0 D Histone H2A-IV P62803 H4_BOVIN
100.0 D Histone H2A-IV P62800 H4_CAIMO
100.0 D Histone H2A-IV Q7KQD1 H4_CHAVR
100.0 D Histone H2A-IV P62801 H4_CHICK
100.0 D Histone H2A-IV P84046 H4_CHITH
100.0 D Histone H2A-IV Q6LAF1 H4_DENKL
100.0 D Histone H2A-IV P84041 H4_DROER
100.0 D Histone H2A-IV P84042 H4_DROHY
100.0 D Histone H2A-IV Q76FE7 H4_DROMA
100.0 D Histone H2A-IV P84040 H4_DROME
100.0 D Histone H2A-IV Q76FF1 H4_DROOR
100.0 D Histone H2A-IV Q76FD9 H4_DROSE
100.0 D Histone H2A-IV P84043 H4_DROSI
100.0 D Histone H2A-IV Q76FF5 H4_DROTE
100.0 D Histone H2A-IV P84044 H4_DROYA
100.0 D Histone H2A-IV P62805 H4_HUMAN
100.0 D Histone H2A-IV Q4R362 H4_MACFA
100.0 D Histone H2A-IV P62806 H4_MOUSE
100.0 D Histone H2A-IV P84049 H4_MYRRU
100.0 D Histone H2A-IV Q6WV73 H4_MYTCA
100.0 D Histone H2A-IV Q6WV74 H4_MYTCH
100.0 D Histone H2A-IV Q7K8C0 H4_MYTED
100.0 D Histone H2A-IV Q6WV90 H4_MYTGA
100.0 D Histone H2A-IV Q6WV72 H4_MYTTR
100.0 D Histone H2A-IV P62797 H4_ONCMY
100.0 D Histone H2A-IV P62796 H4_ORENI
100.0 D Histone H2A-IV P62802 H4_PIG
100.0 D Histone H2A-IV P62795 H4_PLADU
100.0 D Histone H2A-IV Q5RCS7 H4_PONAB
100.0 D Histone H2A-IV P62804 H4_RAT
100.0 D Histone H2A-IV P84050 H4_RHYAM
100.0 D Histone H2A-IV P84045 H4_TIGCA
100.0 D Histone H2A-IV P62794 H4_URECA
100.0 D Histone H2A-IV P62798 H4_XENBO
100.0 D Histone H2A-IV P62799 H4_XENLA
100.0 D Histone H2A-IV Q28DR4 H4_XENTR
98.7 D Histone H2A-IV Q8I0Y4 H4_OIKDI
98.7 D Histone H2A-IV P82888 H4_OLILU
98.7 D Histone H2A-IV Q27765 H4_STYPL
97.4 D Histone H2A-IV P62785 H41_WHEAT
97.4 D Histone H2A-IV P62786 H42_WHEAT
97.4 D Histone H2A-IV P59259 H4_ARATH
97.4 D Histone H2A-IV Q6PMI5 H4_CHEMJ
97.4 D Histone H2A-IV Q6WZ83 H4_EUCGL
97.4 D Histone H2A-IV Q6LAF3 H4_FLATR
97.4 D Histone H2A-IV P62887 H4_LOLTE
97.4 D Histone H2A-IV P62787 H4_MAIZE
97.4 D Histone H2A-IV P62788 H4_PEA
97.4 D Histone H2A-IV Q76H85 H4_SILLA
97.4 D Histone H2A-IV P0CG89 H4_SOYBN
98.7 D Histone H2A-IV P62784 H4_CAEEL
98.7 D Histone H2A-IV P62776 H4_HOLTU
98.7 D Histone H2A-IV P62782 H4_LYTPI
97.4 D Histone H2A-IV Q9U7D0 H4_MASBA
98.7 D Histone H2A-IV P62780 H4_PARLI
98.7 D Histone H2A-IV P62777 H4_PISBR
98.7 D Histone H2A-IV P62778 H4_PISOC
98.7 D Histone H2A-IV P62781 H4_PSAMI
98.7 D Histone H2A-IV P62779 H4_PYCHE
98.7 D Histone H2A-IV P62783 H4_STRPU
96.1 D Histone H2A-IV Q71V09 H4_CAPAN
96.1 D Histone H2A-IV Q6V9I2 H4_SOLCH
96.1 D Histone H2A-IV P35057 H4_SOLLC
98.7 D Histone H2A-IV P70081 H48_CHICK
98.7 D Histone H2A-IV Q8MTV8 H4_APLCA
97.4 D Histone H2A-IV P91882 H4_DIAPU
97.4 D Histone H2A-IV Q43083 H4_PYRSA
97.4 D Histone H2A-IV P27996 H4_SOLST
96.1 D Histone H2A-IV Q41811 H43_MAIZE
97.4 D Histone H2A-IV P91849 H4_APIME
96.1 D Histone H2A-IV Q9HDF5 H4_MORAP
94.7 D Histone H2A-IV P62793 H4_AGABI
96.1 D Histone H2A-IV Q27443 H4_ASCSU
94.7 D Histone H2A-IV P62792 H4_PHACH
94.7 D Histone H2A-IV P50566 H4_CHLRE
94.7 D Histone H2A-IV P04915 H4_PHYPO
96.1 D Histone H2A-IV P83865 H4_PENVA
90.8 D Histone H2A-IV Q8T7J8 H4_EIMTE
96.1 D Histone H2A-IV P91890 H4_TRICD
94.7 D Histone H2A-IV P08436 H4_VOLCA
92.1 D Histone H2A-IV P23750 H41_EMENI
92.1 D Histone H2A-IV Q711M0 H41_TALFU
92.1 D Histone H2A-IV Q7LKT3 H4_ASPFU
92.1 D Histone H2A-IV Q76MU7 H4_ASPOR
92.1 D Histone H2A-IV P04914 H4_NEUCR
92.1 D Histone H2A-IV P23751 H42_EMENI
92.1 D Histone H2A-IV Q8NIQ8 H42_TALFU
92.0 D Histone H2A-IV Q6ZXX3 H4_USTMA
92.1 D Histone H2A-IV Q8J1L3 H4_BLAAD
92.1 D Histone H2A-IV Q8NIG3 H4_CANGA
90.8 D Histone H2A-IV Q757K0 H41_ASHGO
90.8 D Histone H2A-IV Q75AX1 H42_ASHGO
90.8 D Histone H2A-IV P02309 H4_YEAST
92.0 D Histone H2A-IV P62790 H4_STENO
92.0 D Histone H2A-IV P62791 H4_STYLE