Summary information

PDB id
6xt7
Class
DNA
Method
X-ray (1.56 Å)
Summary
Tel25 hybrid four-quartet g-quadruplex with k+
Reference
Beseiso D, Chen EV, McCarthy SE, Martin KN, Gallagher EP, Miao J, Yatsunyk LA (2022): "The first crystal structures of hybrid and parallel four-tetrad intramolecular G-quadruplexes." Nucleic Acids Res., 50, 2959-2972. doi: 10.1093/nar/gkac091.
Abstract
G-quadruplexes (GQs) are non-canonical DNA structures composed of stacks of stabilized G-tetrads. GQs play an important role in a variety of biological processes and may form at telomeres and oncogene promoters among other genomic locations. Here, we investigate nine variants of telomeric DNA from Tetrahymena thermophila with the repeat (TTGGGG)n. Biophysical data indicate that the sequences fold into stable four-tetrad GQs which adopt multiple conformations according to native PAGE. Excitingly, we solved the crystal structure of two variants, TET25 and TET26. The two variants differ by the presence of a 3'-T yet adopt different GQ conformations. TET25 forms a hybrid [3 + 1] GQ and exhibits a rare 5'-top snapback feature. Consequently, TET25 contains four loops: three lateral (TT, TT, and GTT) and one propeller (TT). TET26 folds into a parallel GQ with three TT propeller loops. To the best of our knowledge, TET25 and TET26 are the first reported hybrid and parallel four-tetrad unimolecular GQ structures. The results presented here expand the repertoire of available GQ structures and provide insight into the intricacy and plasticity of the 3D architecture adopted by telomeric repeats from T. thermophila and GQs in general.
G4 notes
16 G-tetrads, 4 G4 helices, 4 G4 stems, 3(-P-Lw-Ln), hybrid-1(3+1), UUDU

Base-block schematics in six views

PyMOL session file PDB file View in 3Dmol.js

List of 16 G-tetrads

 1 glyco-bond=s-ss sugar=3--- groove=wn-- planarity=0.202 type=other  nts=4 GGGG A.DG1001,A.DG1019,A.DG1022,A.DG1004
 2 glyco-bond=--s- sugar=---- groove=-wn- planarity=0.172 type=other  nts=4 GGGG A.DG1005,A.DG1010,A.DG1018,A.DG1023
 3 glyco-bond=--s- sugar=---. groove=-wn- planarity=0.182 type=other  nts=4 GGGG A.DG1006,A.DG1011,A.DG1017,A.DG1024
 4 glyco-bond=--s- sugar=---- groove=-wn- planarity=0.353 type=bowl   nts=4 GGGG A.DG1007,A.DG1012,A.DG1016,A.DG1025
 5 glyco-bond=s-ss sugar=3--- groove=wn-- planarity=0.222 type=other  nts=4 GGGG B.DG1001,B.DG1019,B.DG1022,B.DG1004
 6 glyco-bond=--s- sugar=---- groove=-wn- planarity=0.183 type=other  nts=4 GGGG B.DG1005,B.DG1010,B.DG1018,B.DG1023
 7 glyco-bond=--s- sugar=---3 groove=-wn- planarity=0.180 type=other  nts=4 GGGG B.DG1006,B.DG1011,B.DG1017,B.DG1024
 8 glyco-bond=--s- sugar=---- groove=-wn- planarity=0.332 type=bowl   nts=4 GGGG B.DG1007,B.DG1012,B.DG1016,B.DG1025
 9 glyco-bond=s-ss sugar=3.-- groove=wn-- planarity=0.206 type=other  nts=4 GGGG C.DG1001,C.DG1019,C.DG1022,C.DG1004
10 glyco-bond=--s- sugar=---- groove=-wn- planarity=0.188 type=other  nts=4 GGGG C.DG1005,C.DG1010,C.DG1018,C.DG1023
11 glyco-bond=--s- sugar=---- groove=-wn- planarity=0.177 type=other  nts=4 GGGG C.DG1006,C.DG1011,C.DG1017,C.DG1024
12 glyco-bond=--s- sugar=---- groove=-wn- planarity=0.382 type=bowl   nts=4 GGGG C.DG1007,C.DG1012,C.DG1016,C.DG1025
13 glyco-bond=s-ss sugar=3.-- groove=wn-- planarity=0.207 type=other  nts=4 GGGG D.DG1001,D.DG1019,D.DG1022,D.DG1004
14 glyco-bond=--s- sugar=---- groove=-wn- planarity=0.164 type=other  nts=4 GGGG D.DG1005,D.DG1010,D.DG1018,D.DG1023
15 glyco-bond=--s- sugar=---- groove=-wn- planarity=0.161 type=other  nts=4 GGGG D.DG1006,D.DG1011,D.DG1017,D.DG1024
16 glyco-bond=--s- sugar=---- groove=-wn- planarity=0.342 type=bowl   nts=4 GGGG D.DG1007,D.DG1012,D.DG1016,D.DG1025

List of 4 G4-helices

In DSSR, a G4-helix is defined by stacking interactions of G-tetrads, regardless of backbone connectivity, and may contain more than one G4-stem.

Helix#1, 4 G-tetrad layers, INTRA-molecular, with 1 stem

 1  glyco-bond=s-ss sugar=3--- groove=wn-- Major-->WC nts=4 GGGG A.DG1001,A.DG1019,A.DG1022,A.DG1004
 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG A.DG1010,A.DG1018,A.DG1023,A.DG1005
 3  glyco-bond=-s-- sugar=--.- groove=wn-- WC-->Major nts=4 GGGG A.DG1011,A.DG1017,A.DG1024,A.DG1006
 4  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG A.DG1012,A.DG1016,A.DG1025,A.DG1007
  step#1  mm(<>,outward)  area=15.61 rise=3.47 twist=15.5
  step#2  pm(>>,forward)  area=9.63  rise=3.43 twist=31.8
  step#3  pm(>>,forward)  area=14.07 rise=3.47 twist=24.5
  strand#1 DNA glyco-bond=s--- sugar=3--- nts=4 GGGG A.DG1001,A.DG1010,A.DG1011,A.DG1012
  strand#2 DNA glyco-bond=-sss sugar=---- nts=4 GGGG A.DG1019,A.DG1018,A.DG1017,A.DG1016
  strand#3 DNA glyco-bond=s--- sugar=--.- nts=4 GGGG A.DG1022,A.DG1023,A.DG1024,A.DG1025
  strand#4 DNA glyco-bond=s--- sugar=---- nts=4 GGGG A.DG1004,A.DG1005,A.DG1006,A.DG1007

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3 stacking diagrams 
 1  glyco-bond=s-ss sugar=3--- groove=wn-- Major-->WC nts=4 GGGG A.DG1001,A.DG1019,A.DG1022,A.DG1004
 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG A.DG1010,A.DG1018,A.DG1023,A.DG1005
  step#1  mm(<>,outward)  area=15.61 rise=3.47 twist=15.5

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 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG A.DG1010,A.DG1018,A.DG1023,A.DG1005
 3  glyco-bond=-s-- sugar=--.- groove=wn-- WC-->Major nts=4 GGGG A.DG1011,A.DG1017,A.DG1024,A.DG1006
  step#2  pm(>>,forward)  area=9.63  rise=3.43 twist=31.8

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 3  glyco-bond=-s-- sugar=--.- groove=wn-- WC-->Major nts=4 GGGG A.DG1011,A.DG1017,A.DG1024,A.DG1006
 4  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG A.DG1012,A.DG1016,A.DG1025,A.DG1007
  step#3  pm(>>,forward)  area=14.07 rise=3.47 twist=24.5

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Helix#2, 4 G-tetrad layers, INTRA-molecular, with 1 stem

 1  glyco-bond=s-ss sugar=3--- groove=wn-- Major-->WC nts=4 GGGG B.DG1001,B.DG1019,B.DG1022,B.DG1004
 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG B.DG1010,B.DG1018,B.DG1023,B.DG1005
 3  glyco-bond=-s-- sugar=--3- groove=wn-- WC-->Major nts=4 GGGG B.DG1011,B.DG1017,B.DG1024,B.DG1006
 4  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG B.DG1012,B.DG1016,B.DG1025,B.DG1007
  step#1  mm(<>,outward)  area=15.24 rise=3.51 twist=15.5
  step#2  pm(>>,forward)  area=8.90  rise=3.39 twist=31.9
  step#3  pm(>>,forward)  area=13.65 rise=3.49 twist=24.9
  strand#1 DNA glyco-bond=s--- sugar=3--- nts=4 GGGG B.DG1001,B.DG1010,B.DG1011,B.DG1012
  strand#2 DNA glyco-bond=-sss sugar=---- nts=4 GGGG B.DG1019,B.DG1018,B.DG1017,B.DG1016
  strand#3 DNA glyco-bond=s--- sugar=--3- nts=4 GGGG B.DG1022,B.DG1023,B.DG1024,B.DG1025
  strand#4 DNA glyco-bond=s--- sugar=---- nts=4 GGGG B.DG1004,B.DG1005,B.DG1006,B.DG1007

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3 stacking diagrams 
 1  glyco-bond=s-ss sugar=3--- groove=wn-- Major-->WC nts=4 GGGG B.DG1001,B.DG1019,B.DG1022,B.DG1004
 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG B.DG1010,B.DG1018,B.DG1023,B.DG1005
  step#1  mm(<>,outward)  area=15.24 rise=3.51 twist=15.5

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 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG B.DG1010,B.DG1018,B.DG1023,B.DG1005
 3  glyco-bond=-s-- sugar=--3- groove=wn-- WC-->Major nts=4 GGGG B.DG1011,B.DG1017,B.DG1024,B.DG1006
  step#2  pm(>>,forward)  area=8.90  rise=3.39 twist=31.9

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 3  glyco-bond=-s-- sugar=--3- groove=wn-- WC-->Major nts=4 GGGG B.DG1011,B.DG1017,B.DG1024,B.DG1006
 4  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG B.DG1012,B.DG1016,B.DG1025,B.DG1007
  step#3  pm(>>,forward)  area=13.65 rise=3.49 twist=24.9

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Helix#3, 4 G-tetrad layers, INTRA-molecular, with 1 stem

 1  glyco-bond=s-ss sugar=3.-- groove=wn-- Major-->WC nts=4 GGGG C.DG1001,C.DG1019,C.DG1022,C.DG1004
 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG C.DG1010,C.DG1018,C.DG1023,C.DG1005
 3  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG C.DG1011,C.DG1017,C.DG1024,C.DG1006
 4  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG C.DG1012,C.DG1016,C.DG1025,C.DG1007
  step#1  mm(<>,outward)  area=14.87 rise=3.48 twist=16.5
  step#2  pm(>>,forward)  area=9.27  rise=3.41 twist=32.4
  step#3  pm(>>,forward)  area=14.51 rise=3.49 twist=24.3
  strand#1 DNA glyco-bond=s--- sugar=3--- nts=4 GGGG C.DG1001,C.DG1010,C.DG1011,C.DG1012
  strand#2 DNA glyco-bond=-sss sugar=.--- nts=4 GGGG C.DG1019,C.DG1018,C.DG1017,C.DG1016
  strand#3 DNA glyco-bond=s--- sugar=---- nts=4 GGGG C.DG1022,C.DG1023,C.DG1024,C.DG1025
  strand#4 DNA glyco-bond=s--- sugar=---- nts=4 GGGG C.DG1004,C.DG1005,C.DG1006,C.DG1007

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3 stacking diagrams 
 1  glyco-bond=s-ss sugar=3.-- groove=wn-- Major-->WC nts=4 GGGG C.DG1001,C.DG1019,C.DG1022,C.DG1004
 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG C.DG1010,C.DG1018,C.DG1023,C.DG1005
  step#1  mm(<>,outward)  area=14.87 rise=3.48 twist=16.5

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 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG C.DG1010,C.DG1018,C.DG1023,C.DG1005
 3  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG C.DG1011,C.DG1017,C.DG1024,C.DG1006
  step#2  pm(>>,forward)  area=9.27  rise=3.41 twist=32.4

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 3  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG C.DG1011,C.DG1017,C.DG1024,C.DG1006
 4  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG C.DG1012,C.DG1016,C.DG1025,C.DG1007
  step#3  pm(>>,forward)  area=14.51 rise=3.49 twist=24.3

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Helix#4, 4 G-tetrad layers, INTRA-molecular, with 1 stem

 1  glyco-bond=s-ss sugar=3.-- groove=wn-- Major-->WC nts=4 GGGG D.DG1001,D.DG1019,D.DG1022,D.DG1004
 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG D.DG1010,D.DG1018,D.DG1023,D.DG1005
 3  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG D.DG1011,D.DG1017,D.DG1024,D.DG1006
 4  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG D.DG1012,D.DG1016,D.DG1025,D.DG1007
  step#1  mm(<>,outward)  area=15.06 rise=3.46 twist=16.1
  step#2  pm(>>,forward)  area=9.40  rise=3.41 twist=31.7
  step#3  pm(>>,forward)  area=14.51 rise=3.48 twist=24.4
  strand#1 DNA glyco-bond=s--- sugar=3--- nts=4 GGGG D.DG1001,D.DG1010,D.DG1011,D.DG1012
  strand#2 DNA glyco-bond=-sss sugar=.--- nts=4 GGGG D.DG1019,D.DG1018,D.DG1017,D.DG1016
  strand#3 DNA glyco-bond=s--- sugar=---- nts=4 GGGG D.DG1022,D.DG1023,D.DG1024,D.DG1025
  strand#4 DNA glyco-bond=s--- sugar=---- nts=4 GGGG D.DG1004,D.DG1005,D.DG1006,D.DG1007

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3 stacking diagrams 
 1  glyco-bond=s-ss sugar=3.-- groove=wn-- Major-->WC nts=4 GGGG D.DG1001,D.DG1019,D.DG1022,D.DG1004
 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG D.DG1010,D.DG1018,D.DG1023,D.DG1005
  step#1  mm(<>,outward)  area=15.06 rise=3.46 twist=16.1

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 2  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG D.DG1010,D.DG1018,D.DG1023,D.DG1005
 3  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG D.DG1011,D.DG1017,D.DG1024,D.DG1006
  step#2  pm(>>,forward)  area=9.40  rise=3.41 twist=31.7

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 3  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG D.DG1011,D.DG1017,D.DG1024,D.DG1006
 4  glyco-bond=-s-- sugar=---- groove=wn-- WC-->Major nts=4 GGGG D.DG1012,D.DG1016,D.DG1025,D.DG1007
  step#3  pm(>>,forward)  area=14.51 rise=3.48 twist=24.4

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List of 4 G4-stems

In DSSR, a G4-stem is defined as a G4-helix with backbone connectivity. Bulges are also allowed along each of the four strands.

Stem#1, 3 G-tetrad layers, 3 loops, INTRA-molecular, UUDU, hybrid-(mixed), 3(-P-Lw-Ln), hybrid-1(3+1)

 1  glyco-bond=--s- sugar=---- groove=-wn- WC-->Major nts=4 GGGG A.DG1005,A.DG1010,A.DG1018,A.DG1023
 2  glyco-bond=--s- sugar=---. groove=-wn- WC-->Major nts=4 GGGG A.DG1006,A.DG1011,A.DG1017,A.DG1024
 3  glyco-bond=--s- sugar=---- groove=-wn- WC-->Major nts=4 GGGG A.DG1007,A.DG1012,A.DG1016,A.DG1025
  step#1  pm(>>,forward)  area=9.63  rise=3.43 twist=31.8
  step#2  pm(>>,forward)  area=14.07 rise=3.47 twist=24.5
  strand#1  U DNA glyco-bond=--- sugar=--- nts=3 GGG A.DG1005,A.DG1006,A.DG1007
  strand#2  U DNA glyco-bond=--- sugar=--- nts=3 GGG A.DG1010,A.DG1011,A.DG1012
  strand#3  D DNA glyco-bond=sss sugar=--- nts=3 GGG A.DG1018,A.DG1017,A.DG1016
  strand#4  U DNA glyco-bond=--- sugar=-.- nts=3 GGG A.DG1023,A.DG1024,A.DG1025
  loop#1 type=propeller strands=[#1,#2] nts=2 TT A.DT1008,A.DT1009
  loop#2 type=lateral   strands=[#2,#3] nts=3 GTT A.DG1013,A.DT1014,A.DT1015
  loop#3 type=lateral   strands=[#3,#4] nts=4 GTTG A.DG1019,A.DT1020,A.DT1021,A.DG1022

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Stem#2, 3 G-tetrad layers, 3 loops, INTRA-molecular, UUDU, hybrid-(mixed), 3(-P-Lw-Ln), hybrid-1(3+1)

 1  glyco-bond=--s- sugar=---- groove=-wn- WC-->Major nts=4 GGGG B.DG1005,B.DG1010,B.DG1018,B.DG1023
 2  glyco-bond=--s- sugar=---3 groove=-wn- WC-->Major nts=4 GGGG B.DG1006,B.DG1011,B.DG1017,B.DG1024
 3  glyco-bond=--s- sugar=---- groove=-wn- WC-->Major nts=4 GGGG B.DG1007,B.DG1012,B.DG1016,B.DG1025
  step#1  pm(>>,forward)  area=8.90  rise=3.39 twist=31.9
  step#2  pm(>>,forward)  area=13.65 rise=3.49 twist=24.9
  strand#1  U DNA glyco-bond=--- sugar=--- nts=3 GGG B.DG1005,B.DG1006,B.DG1007
  strand#2  U DNA glyco-bond=--- sugar=--- nts=3 GGG B.DG1010,B.DG1011,B.DG1012
  strand#3  D DNA glyco-bond=sss sugar=--- nts=3 GGG B.DG1018,B.DG1017,B.DG1016
  strand#4  U DNA glyco-bond=--- sugar=-3- nts=3 GGG B.DG1023,B.DG1024,B.DG1025
  loop#1 type=propeller strands=[#1,#2] nts=2 TT B.DT1008,B.DT1009
  loop#2 type=lateral   strands=[#2,#3] nts=3 GTT B.DG1013,B.DT1014,B.DT1015
  loop#3 type=lateral   strands=[#3,#4] nts=4 GTTG B.DG1019,B.DT1020,B.DT1021,B.DG1022

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Stem#3, 3 G-tetrad layers, 3 loops, INTRA-molecular, UUDU, hybrid-(mixed), 3(-P-Lw-Ln), hybrid-1(3+1)

 1  glyco-bond=--s- sugar=---- groove=-wn- WC-->Major nts=4 GGGG C.DG1005,C.DG1010,C.DG1018,C.DG1023
 2  glyco-bond=--s- sugar=---- groove=-wn- WC-->Major nts=4 GGGG C.DG1006,C.DG1011,C.DG1017,C.DG1024
 3  glyco-bond=--s- sugar=---- groove=-wn- WC-->Major nts=4 GGGG C.DG1007,C.DG1012,C.DG1016,C.DG1025
  step#1  pm(>>,forward)  area=9.27  rise=3.41 twist=32.4
  step#2  pm(>>,forward)  area=14.51 rise=3.49 twist=24.3
  strand#1  U DNA glyco-bond=--- sugar=--- nts=3 GGG C.DG1005,C.DG1006,C.DG1007
  strand#2  U DNA glyco-bond=--- sugar=--- nts=3 GGG C.DG1010,C.DG1011,C.DG1012
  strand#3  D DNA glyco-bond=sss sugar=--- nts=3 GGG C.DG1018,C.DG1017,C.DG1016
  strand#4  U DNA glyco-bond=--- sugar=--- nts=3 GGG C.DG1023,C.DG1024,C.DG1025
  loop#1 type=propeller strands=[#1,#2] nts=2 TT C.DT1008,C.DT1009
  loop#2 type=lateral   strands=[#2,#3] nts=3 GTT C.DG1013,C.DT1014,C.DT1015
  loop#3 type=lateral   strands=[#3,#4] nts=4 GTTG C.DG1019,C.DT1020,C.DT1021,C.DG1022

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Stem#4, 3 G-tetrad layers, 3 loops, INTRA-molecular, UUDU, hybrid-(mixed), 3(-P-Lw-Ln), hybrid-1(3+1)

 1  glyco-bond=--s- sugar=---- groove=-wn- WC-->Major nts=4 GGGG D.DG1005,D.DG1010,D.DG1018,D.DG1023
 2  glyco-bond=--s- sugar=---- groove=-wn- WC-->Major nts=4 GGGG D.DG1006,D.DG1011,D.DG1017,D.DG1024
 3  glyco-bond=--s- sugar=---- groove=-wn- WC-->Major nts=4 GGGG D.DG1007,D.DG1012,D.DG1016,D.DG1025
  step#1  pm(>>,forward)  area=9.40  rise=3.41 twist=31.7
  step#2  pm(>>,forward)  area=14.51 rise=3.48 twist=24.4
  strand#1  U DNA glyco-bond=--- sugar=--- nts=3 GGG D.DG1005,D.DG1006,D.DG1007
  strand#2  U DNA glyco-bond=--- sugar=--- nts=3 GGG D.DG1010,D.DG1011,D.DG1012
  strand#3  D DNA glyco-bond=sss sugar=--- nts=3 GGG D.DG1018,D.DG1017,D.DG1016
  strand#4  U DNA glyco-bond=--- sugar=--- nts=3 GGG D.DG1023,D.DG1024,D.DG1025
  loop#1 type=propeller strands=[#1,#2] nts=2 TT D.DT1008,D.DT1009
  loop#2 type=lateral   strands=[#2,#3] nts=3 GTT D.DG1013,D.DT1014,D.DT1015
  loop#3 type=lateral   strands=[#3,#4] nts=4 GTTG D.DG1019,D.DT1020,D.DT1021,D.DG1022

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List of 8 non-stem G4-loops (including the two closing Gs)

 1 type=lateral   helix=#1 nts=4 GTTG A.DG1001,A.DT1002,A.DT1003,A.DG1004
 2 type=lateral   helix=#1 nts=4 GTTG A.DG1019,A.DT1020,A.DT1021,A.DG1022
 3 type=lateral   helix=#2 nts=4 GTTG B.DG1001,B.DT1002,B.DT1003,B.DG1004
 4 type=lateral   helix=#2 nts=4 GTTG B.DG1019,B.DT1020,B.DT1021,B.DG1022
 5 type=lateral   helix=#3 nts=4 GTTG C.DG1001,C.DT1002,C.DT1003,C.DG1004
 6 type=lateral   helix=#3 nts=4 GTTG C.DG1019,C.DT1020,C.DT1021,C.DG1022
 7 type=lateral   helix=#4 nts=4 GTTG D.DG1001,D.DT1002,D.DT1003,D.DG1004
 8 type=lateral   helix=#4 nts=4 GTTG D.DG1019,D.DT1020,D.DT1021,D.DG1022