Details for Accession #: Q8IWI9


Protein Sequence










Peptide Information

ID Peptide Seq Site Residue Position in Peptide Position in Protein
1242 ISPPEPQSFASK S 11 1875
1308 IPGVSTPQTLAGTQK T 6 1558
4015 IPGVSTPQTLAGTQK T 1558
10001103 IPGVSTPQTLAGTQK S 5 1557
10001643 1557
10001688 1872
10002578 1672
10002829 1557
10002842 1872
10003114 1672
10003428 1557
10003429 1561
10003430 1872
10003431 1664
10003432 1866
10005415 ISPPEPQSFASK S 8 1911
10005558 IPGVSTPQTLAGTQK S 5 1557
10006677 IPGVSTPQTLAGTQK S 5 1557
10008363 IPGVSTPQTLAGTQK S 5 1557
10008395 ISPPEPQSFASK S 8 1911
10009449 IPGVSTPQTLAGTQK S 5 1557
10009450 TTGITTPVASVAFPK T 2 1664
10009451 ISPPEPQSFASK S 8 1911
10011693 VMSKVGALQQKIPGVSTPQTLAGTQKFSIRP S 1557
10011694 TLTLRISPPEPQSFASKTGSETKITYSSGGQ S 1875
10011695 SNAPTSPVVYQLPTKSTSYVRTLDSVLKKQS S 861
10013455 VGSFIIELASQRKSR S 1371
10013456 MASSSGTATNRPGK S 1506
10015460 IPGVSTPQTLAGTQK S 5 1557
10015461 S 1872
10016290 IPGVSTPQTLAGTQK S 5 1557
10016624 IPGVSTPQTLAGTQK T 13 1565
10016625 TTGITTPVASVAFPK T 2 1664
10016626 TTGITTPVASVAFPK T 5 1667
10016627 ISPPEPQSFASK S 2 1866
10016776 TTGITTPVASVAFPK T 1 1663
10019488 ISPPEPQSFASK S 1872
10019611 IPGVSTPQTLAGTQK S 1557
10021353 1664
10021406 1561
10022485 TTGITTPVASVAFPK T 2 1663
10022578 ISPPEPQSFASK S 3 1905
10022666 TTGITTPVASVAFPK T 6 1667
10022671 IPGVSTPQTLAGTQK S 6 1557
10022696 IPGVSTPQTLAGTQK T 7 1558
10022740 ISPPEPQSFASK S 9 1911
10022769 TTGITTPVASVAFPK S 11 1672
10023797 IPGVSTPQTLAGTQK S 5 1557
10023798 IPGVSTPQTLAGTQK T 9 1561
10023799 TTGITTPVASVAFPK S 10 1672
10024903 IPGVSTPQTLAGTQK T 6 1558
10024904 ISPPEPQSFASK S 2 1866
10024905 ISPPEPQSFASK S 8 1872
10024906 TTGITTPVASVAFPK T 6 1668
10025281 ISPPEPQSFASK S 8 1872
10025553 IPGVSTPQTLAGTQK S 5 1557
10026022 TTGITTPVASVAFPK T 1 1663
10026023 TTGITTPVASVAFPK S 10 1672
10026203 TTGITTPVASVAFPK T 6 1668
10028274 IPGVSTPQTLAGTQK S 5 1557
10028275 IPGVSTPQTLAGTQK T 9 1561
10028276 TTGITTPVASVAFPK T 1 1663
10028277 TTGITTPVASVAFPK T 2 1664
10028278 TTGITTPVASVAFPK S 10 1672
10028279 ISPPEPQSFASK S 8 1872
10029293 IPGVSTPQTLAGTQK S 5 1557
10029294 IPGVSTPQTLAGTQK T 9 1561
10029295 TTGITTPVASVAFPK T 2 1664
10029296 TTGITTPVASVAFPK S 10 1672
10029297 ISPPEPQSFASK S 2 1866
10029298 ISPPEPQSFASK S 8 1872
10030272 S 1672
10030568 S 1557
10030569 T 1561
10030570 S 1872
10030571 T 1664
10030572 S 1866
10030820 IPGVSTPQTLAGTQK S 5 1557
10031078 ISPPEPQSFASK S 2 1905
10031367 IPGVSTPQTLAGTQK T 13 1565
10031990 TTGITTPVASVAFPK S 10 1672
10031991 TTGITTPVASVAFPK T 1 1663
10031992 TTGITTPVASVAFPK T 2 1664
10031993 ISPPEPQSFASK S 8 1911
10032533 IPGVSTPQTLAGTQK S 1557
10033036 TTGITTPVASVAFPK S 10 1672
10033037 ISPPEPQSFASK S 8 1872
10033110 TTGITTPVASVAFPK T 6 1668
10033601 TTGITTPVASVAFPK T 2 1664
10033943 TTGITTPVASVAFPK S 10 1672
10036208 ISPPEPQSFASK S 2 1866
10037205 IPGVSTPQTLAGTQK 1557
10037266 ISPPEPQSFASK 1872
10037267 TTGITTPVASVAFPK 1672
10037399 IPGVSTPQTLAGTQK 1557
10037426 ISPPEPQSFASK 1872

O-GlcNAcylation Information

ID Method Analytical Throughput Ambiguity Species Sample Type Condition log2Ratio P Value Data Source PMID Comments
1242 MS HTP ambiguous human HEK 293 cells 23301498
1308 MS HTP ambiguous human HEK 293 cells 23301498
4015 MS HTP ambiguous human T cells 29351928
10001103 MS HTP unambiguous human HeLa cells 20 uM GalNAz/200 uM GalNAz 2.631578947 30059200
10001643 MS HTP unambiguous human HeLa cells Interphase/Mitosis 0.947485278 0.009698 34161081 The p value was -log10p in original paper.
10001688 MS HTP unambiguous human HeLa cells Interphase/Mitosis 2.261658904 1.24E-04 34161081 The p value was -log10p in original paper.
10002578 MS HTP unambiguous human HeLa cells Mitotic exit/Early mitosis 100 34161081 The p value was -log10p in original paper.
10002829 MS HTP unambiguous human HeLa cells Mitotic exit/Early mitosis 1.017723537 0.005176 34161081 The p value was -log10p in original paper.
10002842 MS HTP unambiguous human HeLa cells Mitotic exit/Early mitosis 1.567480811 0.005459 34161081 The p value was -log10p in original paper.
10003114 MS HTP unambiguous human colorectal cancer (CRC) cell lines SW620 cells/SW480 cells 0.777607579 0.004466725 35254053
10003428 MS HTP unambiguous human colorectal cancer (CRC) cell lines SW620 cells/SW480 cells 0.490726431 0.001918758 35254053
10003429 MS HTP unambiguous human colorectal cancer (CRC) cell lines SW620 cells/SW480 cells -0.344828497 0.02681275 35254053
10003430 MS HTP unambiguous human colorectal cancer (CRC) cell lines SW620 cells/SW480 cells -0.240823127 0.027338203 35254053
10003431 MS HTP unambiguous human colorectal cancer (CRC) cell lines SW620 cells/SW480 cells -0.184809239 0.123045237 35254053
10003432 MS HTP unambiguous human colorectal cancer (CRC) cell lines SW620 cells/SW480 cells -0.070389328 0.286825943 35254053
10005415 MS HTP unambiguous human HEK 293T cells glucose-deprived/glucose-sufficient 0.68 37541260
10005558 MS HTP unambiguous human HEK 293T cells glucose-deprived/glucose-sufficient 0.765 37541260
10006677 MS HTP unambiguous human HEK 293T cells 21740066
10008363 MS HTP unambiguous human HEK 293 cells 23301498
10008395 MS HTP unambiguous human HEK 293 cells 23301498 This site was described as S1872 of the protein sequence in the original paper
10009449 MS HTP unambiguous human brain 28657654
10009450 MS HTP unambiguous human brain 28657654
10009451 MS HTP unambiguous human brain 28657654 This site was described as S1872 of the protein sequence in the original paper
10011693 MS HTP unambiguous human HeLa cells 30059200
10011694 MS HTP unambiguous human HeLa cells 30059200
10011695 MS HTP unambiguous human HeLa cells 30059200
10013455 MS HTP unambiguous human HeLa cells 30379171
10013456 MS HTP unambiguous human HeLa cells 30379171
10015460 MS HTP unambiguous human HeLa cells 31492838
10015461 MS HTP unambiguous human HeLa cells 31492838 This position assigment in the protein sequence seems problematic, which could not be curated
10016290 MS HTP unambiguous human MCF-7 cells 32574038
10016624 MS HTP unambiguous human MCF-7 cells 32574038
10016625 MS HTP unambiguous human MCF-7 cells 32574038
10016626 MS HTP unambiguous human MCF-7 cells 32574038
10016627 MS HTP unambiguous human MCF-7 cells 32574038
10016776 MS HTP unambiguous human MCF-7 cells 32574038
10019488 MS HTP unambiguous human Hela cells 29237092
10019611 MS HTP unambiguous human Hela cells 29237092
10021353 MS HTP unambiguous human HeLa cells 34161081
10021406 MS HTP unambiguous human HeLa cells 34161081
10022485 MS HTP unambiguous human HeLa cells 34846842
10022578 MS HTP unambiguous human HeLa cells 34846842
10022666 MS HTP unambiguous human HeLa cells 34846842
10022671 MS HTP unambiguous human HeLa cells 34846842
10022696 MS HTP unambiguous human HeLa cells 34846842
10022740 MS HTP unambiguous human HeLa cells 34846842
10022769 MS HTP unambiguous human HeLa cells 34846842
10023797 MS HTP unambiguous human THP1 cells 32938750
10023798 MS HTP unambiguous human THP1 cells 32938750
10023799 MS HTP unambiguous human THP1 cells 32938750
10024903 MS HTP unambiguous human Jurkat cells, MCF-7 cells, K562 cells 35132862
10024904 MS HTP unambiguous human Jurkat cells, MCF-7 cells, K562 cells 35132862
10024905 MS HTP unambiguous human Jurkat cells, MCF-7 cells, K562 cells 35132862
10024906 MS HTP unambiguous human Jurkat cells, MCF-7 cells, K562 cells 35132862
10025281 MS HTP unambiguous human HeLa cells 35254053
10025553 MS HTP unambiguous human HeLa cells 35254053
10026022 MS HTP unambiguous human HeLa cells 35254053
10026023 MS HTP unambiguous human HeLa cells 35254053
10026203 MS HTP unambiguous human HeLa cells 35254053
10028274 MS HTP unambiguous human SW480 cells 35254053
10028275 MS HTP unambiguous human SW480 cells 35254053
10028276 MS HTP unambiguous human SW480 cells 35254053
10028277 MS HTP unambiguous human SW480 cells 35254053
10028278 MS HTP unambiguous human SW480 cells 35254053
10028279 MS HTP unambiguous human SW480 cells 35254053
10029293 MS HTP unambiguous human SW620 cells 35254053
10029294 MS HTP unambiguous human SW620 cells 35254053
10029295 MS HTP unambiguous human SW620 cells 35254053
10029296 MS HTP unambiguous human SW620 cells 35254053
10029297 MS HTP unambiguous human SW620 cells 35254053
10029298 MS HTP unambiguous human SW620 cells 35254053
10030272 MS HTP unambiguous human SW480 cells, SW620 cells SW620/SW480 0.777607579 0.004466725 35254053
10030568 MS HTP unambiguous human SW480 cells, SW620 cells SW620/SW480 0.490726431 0.001918758 35254053
10030569 MS HTP unambiguous human SW480 cells, SW620 cells SW620/SW480 -0.344828497 0.02681275 35254053
10030570 MS HTP unambiguous human SW480 cells, SW620 cells SW620/SW480 -0.240823127 0.027338203 35254053
10030571 MS HTP unambiguous human SW480 cells, SW620 cells SW620/SW480 -0.184809239 0.123045237 35254053
10030572 MS HTP unambiguous human SW480 cells, SW620 cells SW620/SW480 -0.070389328 0.286825943 35254053
10030820 MS HTP unambiguous human HeLa cell nucleus 35289036
10031078 MS HTP unambiguous human HeLa cell nucleus 35289036
10031367 MS HTP unambiguous human HeLa cell nucleus 35289036
10031990 MS HTP unambiguous human HeLa cell nucleus 35289036
10031991 MS HTP unambiguous human HeLa cell nucleus 35289036
10031992 MS HTP unambiguous human HeLa cell nucleus 35289036
10031993 MS HTP unambiguous human HeLa cell nucleus 35289036
10032533 MS HTP unambiguous human MCF-7 cells 35513511
10033036 MS HTP unambiguous human Jurkat cells 35705054
10033037 MS HTP unambiguous human Jurkat cells 35705054
10033110 MS HTP unambiguous human Jurkat cells 35705054
10033601 MS HTP unambiguous human HEK 293T cells glucose-deprived/glucose-sufficient 0.83 37541260
10033943 MS HTP unambiguous human HEK 293T cells glucose-deprived/glucose-sufficient 1.16 37541260
10036208 MS HTP unambiguous human Jurkat cells nucleus/cytoplasm 0.92 35705054
10037205 MS HTP unambiguous human HEK 293T cells TMG/control 1.137503524 30620550 Peptide quantification value was applied to each modification site
10037266 MS HTP unambiguous human HEK 293T cells TMG/control 0.263034406 30620550 Peptide quantification value was applied to each modification site
10037267 MS HTP unambiguous human HEK 293T cells TMG/control 0.263034406 30620550 Peptide quantification value was applied to each modification site
10037399 MS HTP unambiguous human HepG2 cells orafenib sensitive/resistant 0.584962501 30620550 Peptide quantification value was applied to each modification site
10037426 MS HTP unambiguous human HepG2 cells orafenib sensitive/resistant 0.137503524 30620550 Peptide quantification value was applied to each modification site

ID Peptide Seq Site Residue Position in Peptide Position in Protein Method Analytical Throughput Ambiguity Species Sample Type Data Source PMID Comments
1242 ISPPEPQSFASK S 11 1875 MS HTP ambiguous human HEK 293 cells 23301498
1308 IPGVSTPQTLAGTQK T 6 1558 MS HTP ambiguous human HEK 293 cells 23301498
4015 IPGVSTPQTLAGTQK T 1558 MS HTP ambiguous human T cells 29351928
10001103 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human HeLa cells 30059200
10001643 1557 MS HTP unambiguous human HeLa cells 34161081 The p value was -log10p in original paper.
10001688 1872 MS HTP unambiguous human HeLa cells 34161081 The p value was -log10p in original paper.
10002578 1672 MS HTP unambiguous human HeLa cells 34161081 The p value was -log10p in original paper.
10002829 1557 MS HTP unambiguous human HeLa cells 34161081 The p value was -log10p in original paper.
10002842 1872 MS HTP unambiguous human HeLa cells 34161081 The p value was -log10p in original paper.
10003114 1672 MS HTP unambiguous human colorectal cancer (CRC) cell lines 35254053
10003428 1557 MS HTP unambiguous human colorectal cancer (CRC) cell lines 35254053
10003429 1561 MS HTP unambiguous human colorectal cancer (CRC) cell lines 35254053
10003430 1872 MS HTP unambiguous human colorectal cancer (CRC) cell lines 35254053
10003431 1664 MS HTP unambiguous human colorectal cancer (CRC) cell lines 35254053
10003432 1866 MS HTP unambiguous human colorectal cancer (CRC) cell lines 35254053
10005415 ISPPEPQSFASK S 8 1911 MS HTP unambiguous human HEK 293T cells 37541260
10005558 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human HEK 293T cells 37541260
10006677 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human HEK 293T cells 21740066
10008363 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human HEK 293 cells 23301498
10008395 ISPPEPQSFASK S 8 1911 MS HTP unambiguous human HEK 293 cells 23301498 This site was described as S1872 of the protein sequence in the original paper
10009449 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human brain 28657654
10009450 TTGITTPVASVAFPK T 2 1664 MS HTP unambiguous human brain 28657654
10009451 ISPPEPQSFASK S 8 1911 MS HTP unambiguous human brain 28657654 This site was described as S1872 of the protein sequence in the original paper
10011693 VMSKVGALQQKIPGVSTPQTLAGTQKFSIRP S 1557 MS HTP unambiguous human HeLa cells 30059200
10011694 TLTLRISPPEPQSFASKTGSETKITYSSGGQ S 1875 MS HTP unambiguous human HeLa cells 30059200
10011695 SNAPTSPVVYQLPTKSTSYVRTLDSVLKKQS S 861 MS HTP unambiguous human HeLa cells 30059200
10013455 VGSFIIELASQRKSR S 1371 MS HTP unambiguous human HeLa cells 30379171
10013456 MASSSGTATNRPGK S 1506 MS HTP unambiguous human HeLa cells 30379171
10015460 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human HeLa cells 31492838
10015461 S 1872 MS HTP unambiguous human HeLa cells 31492838 This position assigment in the protein sequence seems problematic, which could not be curated
10016290 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human MCF-7 cells 32574038
10016624 IPGVSTPQTLAGTQK T 13 1565 MS HTP unambiguous human MCF-7 cells 32574038
10016625 TTGITTPVASVAFPK T 2 1664 MS HTP unambiguous human MCF-7 cells 32574038
10016626 TTGITTPVASVAFPK T 5 1667 MS HTP unambiguous human MCF-7 cells 32574038
10016627 ISPPEPQSFASK S 2 1866 MS HTP unambiguous human MCF-7 cells 32574038
10016776 TTGITTPVASVAFPK T 1 1663 MS HTP unambiguous human MCF-7 cells 32574038
10019488 ISPPEPQSFASK S 1872 MS HTP unambiguous human Hela cells 29237092
10019611 IPGVSTPQTLAGTQK S 1557 MS HTP unambiguous human Hela cells 29237092
10021353 1664 MS HTP unambiguous human HeLa cells 34161081
10021406 1561 MS HTP unambiguous human HeLa cells 34161081
10022485 TTGITTPVASVAFPK T 2 1663 MS HTP unambiguous human HeLa cells 34846842
10022578 ISPPEPQSFASK S 3 1905 MS HTP unambiguous human HeLa cells 34846842
10022666 TTGITTPVASVAFPK T 6 1667 MS HTP unambiguous human HeLa cells 34846842
10022671 IPGVSTPQTLAGTQK S 6 1557 MS HTP unambiguous human HeLa cells 34846842
10022696 IPGVSTPQTLAGTQK T 7 1558 MS HTP unambiguous human HeLa cells 34846842
10022740 ISPPEPQSFASK S 9 1911 MS HTP unambiguous human HeLa cells 34846842
10022769 TTGITTPVASVAFPK S 11 1672 MS HTP unambiguous human HeLa cells 34846842
10023797 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human THP1 cells 32938750
10023798 IPGVSTPQTLAGTQK T 9 1561 MS HTP unambiguous human THP1 cells 32938750
10023799 TTGITTPVASVAFPK S 10 1672 MS HTP unambiguous human THP1 cells 32938750
10024903 IPGVSTPQTLAGTQK T 6 1558 MS HTP unambiguous human Jurkat cells, MCF-7 cells, K562 cells 35132862
10024904 ISPPEPQSFASK S 2 1866 MS HTP unambiguous human Jurkat cells, MCF-7 cells, K562 cells 35132862
10024905 ISPPEPQSFASK S 8 1872 MS HTP unambiguous human Jurkat cells, MCF-7 cells, K562 cells 35132862
10024906 TTGITTPVASVAFPK T 6 1668 MS HTP unambiguous human Jurkat cells, MCF-7 cells, K562 cells 35132862
10025281 ISPPEPQSFASK S 8 1872 MS HTP unambiguous human HeLa cells 35254053
10025553 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human HeLa cells 35254053
10026022 TTGITTPVASVAFPK T 1 1663 MS HTP unambiguous human HeLa cells 35254053
10026023 TTGITTPVASVAFPK S 10 1672 MS HTP unambiguous human HeLa cells 35254053
10026203 TTGITTPVASVAFPK T 6 1668 MS HTP unambiguous human HeLa cells 35254053
10028274 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human SW480 cells 35254053
10028275 IPGVSTPQTLAGTQK T 9 1561 MS HTP unambiguous human SW480 cells 35254053
10028276 TTGITTPVASVAFPK T 1 1663 MS HTP unambiguous human SW480 cells 35254053
10028277 TTGITTPVASVAFPK T 2 1664 MS HTP unambiguous human SW480 cells 35254053
10028278 TTGITTPVASVAFPK S 10 1672 MS HTP unambiguous human SW480 cells 35254053
10028279 ISPPEPQSFASK S 8 1872 MS HTP unambiguous human SW480 cells 35254053
10029293 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human SW620 cells 35254053
10029294 IPGVSTPQTLAGTQK T 9 1561 MS HTP unambiguous human SW620 cells 35254053
10029295 TTGITTPVASVAFPK T 2 1664 MS HTP unambiguous human SW620 cells 35254053
10029296 TTGITTPVASVAFPK S 10 1672 MS HTP unambiguous human SW620 cells 35254053
10029297 ISPPEPQSFASK S 2 1866 MS HTP unambiguous human SW620 cells 35254053
10029298 ISPPEPQSFASK S 8 1872 MS HTP unambiguous human SW620 cells 35254053
10030272 S 1672 MS HTP unambiguous human SW480 cells, SW620 cells 35254053
10030568 S 1557 MS HTP unambiguous human SW480 cells, SW620 cells 35254053
10030569 T 1561 MS HTP unambiguous human SW480 cells, SW620 cells 35254053
10030570 S 1872 MS HTP unambiguous human SW480 cells, SW620 cells 35254053
10030571 T 1664 MS HTP unambiguous human SW480 cells, SW620 cells 35254053
10030572 S 1866 MS HTP unambiguous human SW480 cells, SW620 cells 35254053
10030820 IPGVSTPQTLAGTQK S 5 1557 MS HTP unambiguous human HeLa cell nucleus 35289036
10031078 ISPPEPQSFASK S 2 1905 MS HTP unambiguous human HeLa cell nucleus 35289036
10031367 IPGVSTPQTLAGTQK T 13 1565 MS HTP unambiguous human HeLa cell nucleus 35289036
10031990 TTGITTPVASVAFPK S 10 1672 MS HTP unambiguous human HeLa cell nucleus 35289036
10031991 TTGITTPVASVAFPK T 1 1663 MS HTP unambiguous human HeLa cell nucleus 35289036
10031992 TTGITTPVASVAFPK T 2 1664 MS HTP unambiguous human HeLa cell nucleus 35289036
10031993 ISPPEPQSFASK S 8 1911 MS HTP unambiguous human HeLa cell nucleus 35289036
10032533 IPGVSTPQTLAGTQK S 1557 MS HTP unambiguous human MCF-7 cells 35513511
10033036 TTGITTPVASVAFPK S 10 1672 MS HTP unambiguous human Jurkat cells 35705054
10033037 ISPPEPQSFASK S 8 1872 MS HTP unambiguous human Jurkat cells 35705054
10033110 TTGITTPVASVAFPK T 6 1668 MS HTP unambiguous human Jurkat cells 35705054
10033601 TTGITTPVASVAFPK T 2 1664 MS HTP unambiguous human HEK 293T cells 37541260
10033943 TTGITTPVASVAFPK S 10 1672 MS HTP unambiguous human HEK 293T cells 37541260
10036208 ISPPEPQSFASK S 2 1866 MS HTP unambiguous human Jurkat cells 35705054
10037205 IPGVSTPQTLAGTQK 1557 MS HTP unambiguous human HEK 293T cells 30620550 Peptide quantification value was applied to each modification site
10037266 ISPPEPQSFASK 1872 MS HTP unambiguous human HEK 293T cells 30620550 Peptide quantification value was applied to each modification site
10037267 TTGITTPVASVAFPK 1672 MS HTP unambiguous human HEK 293T cells 30620550 Peptide quantification value was applied to each modification site
10037399 IPGVSTPQTLAGTQK 1557 MS HTP unambiguous human HepG2 cells 30620550 Peptide quantification value was applied to each modification site
10037426 ISPPEPQSFASK 1872 MS HTP unambiguous human HepG2 cells 30620550 Peptide quantification value was applied to each modification site