Complexes formed between BGO-ZP and BGO with TGF-β and TβRII in solution
as assessed using SEC
and SEC–MALS. (A–C) Superdex 200 16/60 SEC chromatograms
for complexes formed by adding 2.5 equiv of TβRII to 1.0 equiv
of TGF-β2TM, 0.75 equiv of BGO-ZP to 1.0 equiv
of 2.5:1 TGF-β2TM/TβRII binary complex, and 3.0 equiv
of BGO to 1.0 equiv of 2.5:1 TGF-β2TM/TβRII
binary complex, respectively. Peaks labeled “el” and
“tv” on the chromatograms correspond to the exclusion
limit and total volume for the column, respectively. Shown in the
inset is a nonreducing SDS–PAGE gel of the major peaks that
eluted. (D) Plot of the SEC partition coefficient, Kav, as a function of the logarithm of the molecular weight
for three proteins studied alone, TβRII, BGO, and
BGO-ZP (black triangles). The red line corresponds
to a fit of the data for the proteins alone (TβRII, BGO, and BGO-ZP), which are of known size, to a straight
line. Green circles shown on the plot correspond to the Kav values for the TGF-β2TM/TβRII, TGF-β2TM/TβRII/BGO, and TGF-β2TM/TβRII/BGO-ZP complexes
plotted as a function of the molecular weights of the complexes assuming
the stoichiometries inferred from the SPR measurements (1:2 TGF-β2TM:TβRII,
1:2:1 TGF-β2TM:TβRII:BGO, and 1:1:1 TGF-β2TM:TβRII:BGO-ZP). (E) Superdex 200 Increase 10/300 GL SEC–MALS
chromatograms obtained for the same three complexes shown in panels
A–C. Complexes are labeled as follows: BG-TC (TGF-β2TM/TβRII/BGO-ZP), BGO-TC (TGF-β2TM/TβRII/BGO), and TβRII-BC (TGF-β2TM/TβRII). Estimated
molecular weights derived from the multiangle light scattering measurements
are shown below the peak for the TGF-β2TM/TβRII binary
complex (blue traces) and above the peaks for the TGF-β2TM/TβRII/BGO and TGF-β2TM/TβRII/BGO-ZP ternary
complexes (green and red traces, respectively). One unexpected observation
is that the peak corresponding to the excess TGF-β2TM/TβRII
binary complex present in the TGF-β2TM/TβRII/BGO-ZP sample eluted at a volume (panel E, red trace, 13.6 mL) slightly
larger than that of the peak for the TGF-β2TM/TβRII binary
complex sample (panel E, blue trace, 12.6 mL). Multiple runs performed
with decreasing amounts of the TGF-β2TM/TβRII complex
loaded show that this is due to a loading effect, with larger amounts
loaded (and thus higher concentrations) eluting earlier (Figure S3). Most likely, the earlier elution
at higher loading concentrations is the result of the preponderance
of 1:2 TGF-β2TM/TβRII binary complexes, while at lower
loading concentrations, there is a preponderance of 1:1 TGF-β2TM/TβRII
binary complexes.