JBC Papers in Press. Published on October 10, 2012 as Manuscript M112.398768
The latest version is at http://www.jbc.org/cgi/doi/10.1074/jbc.M112.398768
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V0
0.8
1
2
3
0.6
0.4
0.2
0.2
Protein activity (μmol/min)
Protein and Phospholipid
concentration (mg/ml)
1.0
Activity retention
C12E8 solubilized and Tween 20 eluent
2
1
1
(D)
C12E8 solubilized and Tween 20 eluent
6
8
10
12
14
16
18
Stokes radius nm
4
20
Protein concentration (mg/ml)
C12E8 solubilized and C12E8 eluent
1.0
0.4
0.8
0.3
0.6
0.2
0.4
0.1
0.2
6
8
10
12
14
16
18
20
Phospholipid concentration (mg/ml)
Elution Volume (ml)
(B)
0,5
Thyr
8
Fer
6
Cat
4
HSA
CA
2
0.2
(E)
0.4
Kd
0.6
TrpI
0,1
0,0
0.8
1 2 3 4 5 MR 6 7 8 9 10 TV
0.0
,
Elution Volume (ml)
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0.1
0.0
2
Time (Days)
Protein concentration (mg/ml)
(C)
Activity (μmol/mg/min)
C12E8 solubilized and Tween 20 eluent
(A)
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