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Table 5.

Reported residues of antigenic relevance identified with mouse monoclonal and chicken polyclonal antibodies

Most frequent amino acid residue in naturec
Residuea Conservation %b Antigenic site American Y439 Korean G1-E G1-W BJ94 Published mAb escape Impact on chicken antisera Reference(s)
72 82.2 H9-B G G G G/E G G/E G-E G-E Okamatsu et al. 2008; Peacock et al. 2018
74 73.9 K K K R R R R-K, R-Gd R-G Zhu et al. 2015; Peacock et al. 2018
98 98.9 L L L L L L L-Q Okamatsu et al. 2008; Peacock et al. 2018
109 56.4 K/S/R N R S S R/S S-R, S-I Kaverin et al. 2004; Peacock et al. 2018
115 91.5 H9-B Q Q L Q Q Q/L Q-P, Q-R, Q-L Q-P Peacock et al. 2016, 2018; Jin et al. 2019
120 81.7 H9-B T/R T R T/A T/S T/S T-K Peacock et al. 2016, 2018
121 54.6 I/T I T I I I I-Td I-T Peacock et al. 2018
127 49 Overlap S S T T/D T/S S/D S-Ne T-Ne Kaverin et al. 2004; Okamatsu et al. 2008; Peacock et al. 2018
129 99.6 Site I T T T T T T T-A, T-K T-K Kaverin et al. 2004; Peacock et al. 2018
131 84.8 K/R K K K K K/T K-Ne, K-Id, K-Rd, A-Rd K-I Okamatsu et al. 2008; Peacock et al. 2018
135 62.4 H9-B/site II N/D N G/N/D D/G D D/G D-E, D-N, D-Gd D-G Kaverin et al. 2004; Wan et al. 2014; Peacock et al. 2018
137 99.8 F F F F F F F-L F-L Kaverin et al. 2004; Peacock et al. 2018
139 98.9 H9-B R R R R R R R-G, R-M Peacock et al. 2016, 2018
145 99.8 H9-A T T T T T T T-I T-I, S-N Ping et al. 2008; Zhu et al. 2015; Peacock et al. 2016, 2018
146 70.8 H9-A/site I H H H Q Q Q/R Q-K Q-H Wan et al. 2014; Peacock et al. 2018
147 99.3 Site I K K K K K K K-T Kaverin et al. 2004; Wan et al. 2014; Peacock et al. 2018
148 59.9 Site I S S S S N N/S/D N-D N-D Okamatsu et al. 2008; Zhu et al. 2015; Peacock et al. 2018
149 54.3 H9-A/site I N N N G G N N-D, N-K G-D/K Wan et al. 2014; Peacock et al. 2018
150 33 Site I S S S S/N/F L/N/F/V/G A/N/D/E A-Te, A-D A/N-V/Ld, A-Fd, F-Ld, F-S- Td A-L/F, L-Se/A Wan et al. 2014; Peacock et al. 2018
152 99.5 Site I P P P P P P P-L Kaverin et al. 2004; Peacock et al. 2018
162 50.6 H9-B E/K E W R R/Q Q/R R-W Peacock et al. 2016, 2018
178 95.5 Site II D D D D/E D D/E D-V Wan et al. 2014; Peacock et al. 2018
179 93.6 H9-A/overlap T T T T T T/D T-A T-N Kaverin et al. 2004; Zhu et al. 2015; Peacock et al. 2018
180 46.8 H9-A E E E A/E A/T/V A/V/T A-E-Td, A-Vd, E-Dd A-E Peacock et al. 2018
182 85.7 N/T T M T T T/R T-I T-R Okamatsu et al. 2008; Peacock et al. 2018
183 86.8 H9-A/site II N/D N N N N N/D N-D, N-S, N-T, N-E/Td N-T Kaverin et al. 2004; Okamatsu et al. 2008; Wan et al. 2014; Peacock et al. 2016, 2018
188 75 Overlap T/A A/D A T/N T/N T T-Ne T-Ne Kaverin et al. 2004; Peacock et al. 2018
189 98.7 D D D D D D D-Ne D-Ne Wan et al. 2014; Peacock et al. 2018; Jin et al. 2019
195 59.7 T T T T/A T A/T A-Td NT Peacock et al. 2018
198 46.6 D D D D/N N/D E/D D-N Peacock et al. 2018
200 68.6 N N N N D N/D N-D N-D Wan et al. 2014; Jin et al. 2019
212 96.9 H9-A L L L/I L L L L-H, L-P Okamatsu et al. 2008; Peacock et al. 2016, 2018
216 62.3 Site II Q Q Q Q/L L/Q L/Q L-Q L-Q Kaverin et al. 2004; Peacock et al. 2018
217 55.4 H9-A Q Q Q Q I Q/M I-T I-L/M/Q Peacock et al. 2016, 2018
234 99.8 H9-A R R R R R R R-Q, L-M Peacock et al. 2016, 2018; Jin et al. 2019
249 65.7 V I I I/V V/I I I-V/Ad I-V Peacock et al. 2018
264 51.4 N N N/K N/K N/K/S K/N K-Nd NT Peacock et al. 2018
276 83.7 R/K K K K K K/R K-Rd NT Peacock et al. 2018
288 61.9 I/V V V I I/V V I-Vd NT Peacock et al. 2018
306 99.3 K K K K K K I-Kd NT Peacock et al. 2018

The most frequent residues at those positions for each H9N2 lineage are reported.

(NT) Not tested.

aAll HA positions follow the H9 numbering. Percent of conservation at selected residues is based on the global avian H9N2 HA1 amino acid alignment of 600 sequences from the analysis in Figure 3 with Scop3D entropy calculation (Vermeire et al. 2015).

bPercent of conservation.

cMost frequent amino acid residue at selected positions based on global H9 phylogenetic analysis of HA1.

dIdentified by integrated modeling of HI titers and gene sequence data.

ePotential glycosylation sites.