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  • The lower JRFL autologous tier 2 titers may reflect less stability of the trimer, a more inherently neutralization-resistant virus, or exposure of less potent neutralizing determinants. Historically, the JRFL monomer does not elicit tier 2 autologous neutralizing antibodies [], whereas the BG505 monomer does []. In fact, the relative rank order of the trimers by DSC-determined global stability was roughly associated with the ability to elicit tier 2 autologous neutralizing antibodies in vivo. The order from the lowest Tm to the highest, and elicitation of tier 2 autologous neutralization, being JRFL NFL], indicating the consistency of the results generated in different laboratories and in two different small animal models. Although not directly tested in the immunogenicity study presented here, given that X-link JRFL and BG505 NFL were able to more consistently elicit autologous neutralizing antibodies and at higher titers for X-link BG505 NFL compared to Wt, it would be expected then that X-link JRFL and BG505 SOSIP would also be able to more consistently elicit autologous neutralizing antibodies and at higher titers that Wt.

    That the chemically stabilized trimers of both JRFL and BG505 NFL origin became partially resistant to opening following ELISA plate adsorption and elicited enhanced tier 2 autologous neutralization relative to Wt trimers, suggested that some of this enhancement might be linked to quaternary-related neutralizing antibodies. However, even the potent BG505 tier 2 autologous neutralization observed could be inhibited completely by monomeric BG505 gp120, as could virtually all BG505-trimer elicited tier 2 autologous neutralization, whether generated from fixed or unfixed trimers. To note however, the BG505 monomer is somewhat unusual in that it is well-recognized by the usually quaternary-dependent bNAb, PG9, and does consistently elicit tier 2 autologous neutralizing antibodies in small animals as recently reported []. These properties may indicate that the conformation of the BG505 monomer is similar to its conformation on the trimer, relative to most other gp120 monomers. In contrast, for the JRFL X-link trimers, there were indications that some of the neutralizing antibodies may be quaternary-dependent. This apparent difference may be due to inherent relative trimer stabilities (JRFL less than BG505) and more heterogeneity, differential penetration of the fixative into the less stable JRFL trimers, or be a property of the more neutralization resistant JRFL virus. In the future, cloning of quaternary-specific mAbs from the JRFL immunized animals will confirm the indications from the adsorption analysis.

  • The most stable BG505 trimers were very poor at eliciting HXBc2 neutralization, considered an “open” tier 1A virus, which may reflect limited exposure of the occluded CD4bs on the most stable of the spikes assessed here. Alternatively, this might indicate that the cross reactivity at the CD4bs is low between BG505 (clade A) and HXBc2 (clade B). However, some of the BG505 elicited antisera could cross-neutralize HXBc2 and we have seen in another study in progress that more potent HXBc2 neutralization is indeed achievable with BG505 gp160 DNA, presumably due to incomplete cleavage resulting in open spikes akin to foldon trimers [, ]. These data suggest that strategies to better expose this site on the more stable trimers may be needed.

    In terms of trimer stability in vitro, native gel analysis of the JRFL NFL trimer compared to the more stable BG505 NFL indicated that the JRFL trimers displayed a propensity to dissociate to monomers, whereas BG505 trimers appeared to degrade, perhaps due to low levels of protease present in the recombinant trimer preparation. Although chemical fixation does seem to overcome either of these unfolding pathways, the heterogeneity of the oligomeric subtypes detected by the SDS gel analysis following fixation indicated that not all trimers were similarly cross-linked, perhaps due to the disperse locations of the reactive K side chains on the solvent exposed surface of the trimers. In our study, chemical cross-linking of the trimers resulted in a large increase in thermostability for BG505 NFL, although not quite as large of an increase in the Tm as recently reported by the Sattentau group []. The resulting X-link BG505 NFL trimer DSC profile peak did broaden (i.e. greater T1/2), indicating some differences in the cross-linking process, but was more uniform and narrow as a single peak, signifying less sample heterogeneity overall, than the X-link BG505 SOSIP trimers. These differences may be due to slightly different cross-linking conditions or differences in the purification methods for the SOSIP trimers compared to the NFL trimers. That X-link JRFL NFL did not obtain as high as a Tm gain as X-link JRFL SOSIP may indicate that the slightly more stable JRFL SOSIP provided for a few more opportunities for cross-links to form.


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    There was clear intra-clade bias for tier 1B neutralization and is likely due to differences in V3 sequences between clade B (GPGR and flanking residues) and the more similar clades A and C sequences (both with GPGQ at the tip). Not surprisingly, with the exception of the non-V3 matched HXBc2, most tier 1 elicited neutralizing activity mapped to either the clade B V3 (BaL.26) for JRFL or the clade C V3 (ZM109) for BG505 and 16055. Since V3 is slightly exposed on the JRFL SOSIP or NFL trimers [, ], elicitation of V3-directed tier 1 neutralizing activity was not unexpected. Although V3 exposure may be less with the BG505 trimers, some V3-directed neutralization was still elicited by these trimers, indicating that perhaps these trimers partially open in vivo. Of note, although X-link BG505 NFL elicited V3-directed antibodies, it also yielded the highest tier 2 autologous neutralization, indicating that such off target antibodies don’t necessarily preclude the efficient elicitation of tier 2 autologous neutralizing antibodies. Although efforts were made to eliminate or reduce V3 exposure in the BG505 SOSIP context, there was no significant improvement in autologous tier 2 neutralizing serum titers [], consistent with our observation that V3 exposure does not readily affect the elicitation of autologous tier 2 neutralizing antibodies.

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Antibody-binding kinetics were measured for CSL and CSL-SOS trimers using a panel of representative (a) bNAbs and (b) non-NAbs, with additional antibody binding profiles shown in . Sensorgrams were obtained from an Octet RED96 using a trimer titration series of six concentrations (200–12.5nM by twofold dilution). D values calculated from 1:1 global fitting are labelled for V1V2 apex-directed bNAbs (PGDM1400 and PG16) in a. UFO, uncleaved prefusion-optimized.