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anno_start	anno_end	anno_text	entity_type	sentence	section
6	13	Cryo-EM	experimental_method	Using Cryo-EM to Map Small Ligands on Dynamic Metabolic Enzymes: Studies with Glutamate Dehydrogenase	TITLE
78	101	Glutamate Dehydrogenase	protein_type	Using Cryo-EM to Map Small Ligands on Dynamic Metabolic Enzymes: Studies with Glutamate Dehydrogenase	TITLE
0	24	Cryo-electron microscopy	experimental_method	Cryo-electron microscopy (cryo-EM) methods are now being used to determine structures at near-atomic resolution and have great promise in molecular pharmacology, especially in the context of mapping the binding of small-molecule ligands to protein complexes that display conformational flexibility.	ABSTRACT
26	33	cryo-EM	experimental_method	Cryo-electron microscopy (cryo-EM) methods are now being used to determine structures at near-atomic resolution and have great promise in molecular pharmacology, especially in the context of mapping the binding of small-molecule ligands to protein complexes that display conformational flexibility.	ABSTRACT
75	85	structures	evidence	Cryo-electron microscopy (cryo-EM) methods are now being used to determine structures at near-atomic resolution and have great promise in molecular pharmacology, especially in the context of mapping the binding of small-molecule ligands to protein complexes that display conformational flexibility.	ABSTRACT
30	53	glutamate dehydrogenase	protein_type	We illustrate this here using glutamate dehydrogenase (GDH), a 336-kDa metabolic enzyme that catalyzes the oxidative deamination of glutamate.	ABSTRACT
55	58	GDH	protein_type	We illustrate this here using glutamate dehydrogenase (GDH), a 336-kDa metabolic enzyme that catalyzes the oxidative deamination of glutamate.	ABSTRACT
132	141	glutamate	chemical	We illustrate this here using glutamate dehydrogenase (GDH), a 336-kDa metabolic enzyme that catalyzes the oxidative deamination of glutamate.	ABSTRACT
17	20	GDH	protein_type	Dysregulation of GDH leads to a variety of metabolic and neurologic disorders.	ABSTRACT
39	46	cryo-EM	experimental_method	Here, we report near-atomic resolution cryo-EM structures, at resolutions ranging from 3.2 Å to 3.6 Å for GDH complexes, including complexes for which crystal structures are not available.	ABSTRACT
47	57	structures	evidence	Here, we report near-atomic resolution cryo-EM structures, at resolutions ranging from 3.2 Å to 3.6 Å for GDH complexes, including complexes for which crystal structures are not available.	ABSTRACT
106	109	GDH	protein_type	Here, we report near-atomic resolution cryo-EM structures, at resolutions ranging from 3.2 Å to 3.6 Å for GDH complexes, including complexes for which crystal structures are not available.	ABSTRACT
151	169	crystal structures	evidence	Here, we report near-atomic resolution cryo-EM structures, at resolutions ranging from 3.2 Å to 3.6 Å for GDH complexes, including complexes for which crystal structures are not available.	ABSTRACT
41	45	NADH	chemical	We show that the binding of the coenzyme NADH alone or in concert with GTP results in a binary mixture in which the enzyme is in either anopenor “closed” state.	ABSTRACT
71	74	GTP	chemical	We show that the binding of the coenzyme NADH alone or in concert with GTP results in a binary mixture in which the enzyme is in either anopenor “closed” state.	ABSTRACT
140	144	open	protein_state	We show that the binding of the coenzyme NADH alone or in concert with GTP results in a binary mixture in which the enzyme is in either anopenor “closed” state.	ABSTRACT
150	156	closed	protein_state	We show that the binding of the coenzyme NADH alone or in concert with GTP results in a binary mixture in which the enzyme is in either anopenor “closed” state.	ABSTRACT
12	21	structure	evidence	Whereas the structure of NADH in the active site is similar between the open and closed states, it is unexpectedly different at the regulatory site.	ABSTRACT
25	29	NADH	chemical	Whereas the structure of NADH in the active site is similar between the open and closed states, it is unexpectedly different at the regulatory site.	ABSTRACT
37	48	active site	site	Whereas the structure of NADH in the active site is similar between the open and closed states, it is unexpectedly different at the regulatory site.	ABSTRACT
72	76	open	protein_state	Whereas the structure of NADH in the active site is similar between the open and closed states, it is unexpectedly different at the regulatory site.	ABSTRACT
81	87	closed	protein_state	Whereas the structure of NADH in the active site is similar between the open and closed states, it is unexpectedly different at the regulatory site.	ABSTRACT
132	147	regulatory site	site	Whereas the structure of NADH in the active site is similar between the open and closed states, it is unexpectedly different at the regulatory site.	ABSTRACT
117	138	X-ray crystallography	experimental_method	Our studies thus demonstrate that even in instances when there is considerable structural information available from X-ray crystallography, cryo-EM methods can provide useful complementary insights into regulatory mechanisms for dynamic protein complexes.	ABSTRACT
140	147	cryo-EM	experimental_method	Our studies thus demonstrate that even in instances when there is considerable structural information available from X-ray crystallography, cryo-EM methods can provide useful complementary insights into regulatory mechanisms for dynamic protein complexes.	ABSTRACT
19	43	cryo-electron microscopy	experimental_method	Recent advances in cryo-electron microscopy (cryo-EM) allow determination of structures of small protein complexes and membrane proteins at near-atomic resolution, marking a critical shift in the structural biology field.	INTRO
45	52	cryo-EM	experimental_method	Recent advances in cryo-electron microscopy (cryo-EM) allow determination of structures of small protein complexes and membrane proteins at near-atomic resolution, marking a critical shift in the structural biology field.	INTRO
77	87	structures	evidence	Recent advances in cryo-electron microscopy (cryo-EM) allow determination of structures of small protein complexes and membrane proteins at near-atomic resolution, marking a critical shift in the structural biology field.	INTRO
150	165	crystallization	experimental_method	One specific area of broad general interest in drug discovery is the localization of bound ligands and cofactors under conditions in which efforts at crystallization have not been successful because of structural heterogeneity.	INTRO
7	14	cryo-EM	experimental_method	Recent cryo-EM analyses have already demonstrated that it is now possible to use single-particle cryo-EM methods to localize small bound ligands or inhibitors on target proteins.	INTRO
81	104	single-particle cryo-EM	experimental_method	Recent cryo-EM analyses have already demonstrated that it is now possible to use single-particle cryo-EM methods to localize small bound ligands or inhibitors on target proteins.	INTRO
37	46	mammalian	taxonomy_domain	Here, we address this question using mammalian glutamate dehydrogenase as an example.	INTRO
47	70	glutamate dehydrogenase	protein_type	Here, we address this question using mammalian glutamate dehydrogenase as an example.	INTRO
0	23	Glutamate dehydrogenase	protein_type	Glutamate dehydrogenase (GDH) is a highly conserved enzyme expressed in most organisms.	INTRO
25	28	GDH	protein_type	Glutamate dehydrogenase (GDH) is a highly conserved enzyme expressed in most organisms.	INTRO
35	51	highly conserved	protein_state	Glutamate dehydrogenase (GDH) is a highly conserved enzyme expressed in most organisms.	INTRO
0	3	GDH	protein_type	GDH plays a central role in glutamate metabolism by catalyzing the reversible oxidative deamination of glutamate to generate α-ketoglutarate and ammonia, with the concomitant transfer of a pair of electrons to either NAD+ or NADP+.	INTRO
28	37	glutamate	chemical	GDH plays a central role in glutamate metabolism by catalyzing the reversible oxidative deamination of glutamate to generate α-ketoglutarate and ammonia, with the concomitant transfer of a pair of electrons to either NAD+ or NADP+.	INTRO
103	112	glutamate	chemical	GDH plays a central role in glutamate metabolism by catalyzing the reversible oxidative deamination of glutamate to generate α-ketoglutarate and ammonia, with the concomitant transfer of a pair of electrons to either NAD+ or NADP+.	INTRO
125	140	α-ketoglutarate	chemical	GDH plays a central role in glutamate metabolism by catalyzing the reversible oxidative deamination of glutamate to generate α-ketoglutarate and ammonia, with the concomitant transfer of a pair of electrons to either NAD+ or NADP+.	INTRO
145	152	ammonia	chemical	GDH plays a central role in glutamate metabolism by catalyzing the reversible oxidative deamination of glutamate to generate α-ketoglutarate and ammonia, with the concomitant transfer of a pair of electrons to either NAD+ or NADP+.	INTRO
217	221	NAD+	chemical	GDH plays a central role in glutamate metabolism by catalyzing the reversible oxidative deamination of glutamate to generate α-ketoglutarate and ammonia, with the concomitant transfer of a pair of electrons to either NAD+ or NADP+.	INTRO
225	230	NADP+	chemical	GDH plays a central role in glutamate metabolism by catalyzing the reversible oxidative deamination of glutamate to generate α-ketoglutarate and ammonia, with the concomitant transfer of a pair of electrons to either NAD+ or NADP+.	INTRO
14	17	GDH	protein_type	Regulation of GDH is tightly controlled through multiple allosteric mechanisms.	INTRO
10	50	biochemical and crystallographic studies	experimental_method	Extensive biochemical and crystallographic studies have characterized the enzymatic activity of GDH and its modulation by a chemically diverse group of compounds such as nucleotides, amino acids, steroid hormones, antipsychotic drugs, and natural products.	INTRO
96	99	GDH	protein_type	Extensive biochemical and crystallographic studies have characterized the enzymatic activity of GDH and its modulation by a chemically diverse group of compounds such as nucleotides, amino acids, steroid hormones, antipsychotic drugs, and natural products.	INTRO
0	30	X-ray crystallographic studies	experimental_method	X-ray crystallographic studies have shown that the functional unit of GDH is a homohexamer composed of a trimer of dimers, with a 3-fold axis and an equatorial plane that define its D3 symmetry (Fig. 1A).	INTRO
70	73	GDH	protein_type	X-ray crystallographic studies have shown that the functional unit of GDH is a homohexamer composed of a trimer of dimers, with a 3-fold axis and an equatorial plane that define its D3 symmetry (Fig. 1A).	INTRO
79	90	homohexamer	oligomeric_state	X-ray crystallographic studies have shown that the functional unit of GDH is a homohexamer composed of a trimer of dimers, with a 3-fold axis and an equatorial plane that define its D3 symmetry (Fig. 1A).	INTRO
105	111	trimer	oligomeric_state	X-ray crystallographic studies have shown that the functional unit of GDH is a homohexamer composed of a trimer of dimers, with a 3-fold axis and an equatorial plane that define its D3 symmetry (Fig. 1A).	INTRO
115	121	dimers	oligomeric_state	X-ray crystallographic studies have shown that the functional unit of GDH is a homohexamer composed of a trimer of dimers, with a 3-fold axis and an equatorial plane that define its D3 symmetry (Fig. 1A).	INTRO
12	20	protomer	oligomeric_state	Each 56-kDa protomer consists of three domains.	INTRO
30	45	dimer interface	site	The first is located near the dimer interface and forms the core of the hexamer.	INTRO
72	79	hexamer	oligomeric_state	The first is located near the dimer interface and forms the core of the hexamer.	INTRO
14	39	nucleotide-binding domain	structure_element	The second, a nucleotide-binding domain (NBD) with a Rossmann fold, defines one face of the catalytic cleft bounded by the core domain.	INTRO
41	44	NBD	structure_element	The second, a nucleotide-binding domain (NBD) with a Rossmann fold, defines one face of the catalytic cleft bounded by the core domain.	INTRO
53	66	Rossmann fold	structure_element	The second, a nucleotide-binding domain (NBD) with a Rossmann fold, defines one face of the catalytic cleft bounded by the core domain.	INTRO
92	107	catalytic cleft	site	The second, a nucleotide-binding domain (NBD) with a Rossmann fold, defines one face of the catalytic cleft bounded by the core domain.	INTRO
32	35	NBD	structure_element	During the catalytic cycle, the NBD executes a large movement, hinged around a “pivot” helix, that closes the catalytic cleft, and drives a large conformational change in the hexamer from open to closed states (Fig. 1B).	INTRO
79	92	“pivot” helix	structure_element	During the catalytic cycle, the NBD executes a large movement, hinged around a “pivot” helix, that closes the catalytic cleft, and drives a large conformational change in the hexamer from open to closed states (Fig. 1B).	INTRO
110	125	catalytic cleft	site	During the catalytic cycle, the NBD executes a large movement, hinged around a “pivot” helix, that closes the catalytic cleft, and drives a large conformational change in the hexamer from open to closed states (Fig. 1B).	INTRO
175	182	hexamer	oligomeric_state	During the catalytic cycle, the NBD executes a large movement, hinged around a “pivot” helix, that closes the catalytic cleft, and drives a large conformational change in the hexamer from open to closed states (Fig. 1B).	INTRO
188	192	open	protein_state	During the catalytic cycle, the NBD executes a large movement, hinged around a “pivot” helix, that closes the catalytic cleft, and drives a large conformational change in the hexamer from open to closed states (Fig. 1B).	INTRO
196	202	closed	protein_state	During the catalytic cycle, the NBD executes a large movement, hinged around a “pivot” helix, that closes the catalytic cleft, and drives a large conformational change in the hexamer from open to closed states (Fig. 1B).	INTRO
30	37	antenna	structure_element	The third domain, dubbed the “antenna,” is an evolutionary acquisition in protista and animals.	INTRO
74	82	protista	taxonomy_domain	The third domain, dubbed the “antenna,” is an evolutionary acquisition in protista and animals.	INTRO
87	94	animals	taxonomy_domain	The third domain, dubbed the “antenna,” is an evolutionary acquisition in protista and animals.	INTRO
0	8	Antennae	structure_element	Antennae of adjacent protomers in each trimer intercalate to form a bundle, perpendicular to the pivot helices, that protrudes along the distal extremes of the 3-fold axis.	INTRO
21	30	protomers	oligomeric_state	Antennae of adjacent protomers in each trimer intercalate to form a bundle, perpendicular to the pivot helices, that protrudes along the distal extremes of the 3-fold axis.	INTRO
39	45	trimer	oligomeric_state	Antennae of adjacent protomers in each trimer intercalate to form a bundle, perpendicular to the pivot helices, that protrudes along the distal extremes of the 3-fold axis.	INTRO
97	110	pivot helices	structure_element	Antennae of adjacent protomers in each trimer intercalate to form a bundle, perpendicular to the pivot helices, that protrudes along the distal extremes of the 3-fold axis.	INTRO
7	15	protomer	oligomeric_state	When a protomer undergoes a conformational change, the rotation of its pivot helix is transferred through the antenna to the adjacent subunit.	INTRO
71	82	pivot helix	structure_element	When a protomer undergoes a conformational change, the rotation of its pivot helix is transferred through the antenna to the adjacent subunit.	INTRO
110	117	antenna	structure_element	When a protomer undergoes a conformational change, the rotation of its pivot helix is transferred through the antenna to the adjacent subunit.	INTRO
134	141	subunit	structure_element	When a protomer undergoes a conformational change, the rotation of its pivot helix is transferred through the antenna to the adjacent subunit.	INTRO
21	28	antenna	structure_element	The influence of the antenna, present only in protozoan and metazoan enzymes, has been proposed to explain its cooperative behavior, which is absent in bacterial homologs.	INTRO
46	55	protozoan	taxonomy_domain	The influence of the antenna, present only in protozoan and metazoan enzymes, has been proposed to explain its cooperative behavior, which is absent in bacterial homologs.	INTRO
60	68	metazoan	taxonomy_domain	The influence of the antenna, present only in protozoan and metazoan enzymes, has been proposed to explain its cooperative behavior, which is absent in bacterial homologs.	INTRO
152	161	bacterial	taxonomy_domain	The influence of the antenna, present only in protozoan and metazoan enzymes, has been proposed to explain its cooperative behavior, which is absent in bacterial homologs.	INTRO
51	54	GDH	protein_type	Structure and quaternary conformational changes in GDH. (A) Views of open (PDB ID 1NR7) and closed (PDB 3MW9) states of the GDH hexamer, shown in ribbon representation perpendicular to the 2-fold symmetry axis (side view, top) and 3-fold symmetry axis (top view, bottom).	FIG
69	73	open	protein_state	Structure and quaternary conformational changes in GDH. (A) Views of open (PDB ID 1NR7) and closed (PDB 3MW9) states of the GDH hexamer, shown in ribbon representation perpendicular to the 2-fold symmetry axis (side view, top) and 3-fold symmetry axis (top view, bottom).	FIG
92	98	closed	protein_state	Structure and quaternary conformational changes in GDH. (A) Views of open (PDB ID 1NR7) and closed (PDB 3MW9) states of the GDH hexamer, shown in ribbon representation perpendicular to the 2-fold symmetry axis (side view, top) and 3-fold symmetry axis (top view, bottom).	FIG
124	127	GDH	protein_type	Structure and quaternary conformational changes in GDH. (A) Views of open (PDB ID 1NR7) and closed (PDB 3MW9) states of the GDH hexamer, shown in ribbon representation perpendicular to the 2-fold symmetry axis (side view, top) and 3-fold symmetry axis (top view, bottom).	FIG
128	135	hexamer	oligomeric_state	Structure and quaternary conformational changes in GDH. (A) Views of open (PDB ID 1NR7) and closed (PDB 3MW9) states of the GDH hexamer, shown in ribbon representation perpendicular to the 2-fold symmetry axis (side view, top) and 3-fold symmetry axis (top view, bottom).	FIG
11	20	protomers	oligomeric_state	Only three protomers are shown in the top view for purposes of visual clarity.	FIG
137	141	open	protein_state	The dashed lines and arrows, respectively, highlight the slight extension in length, and twist in shape that occurs with transition from open to the closed state.	FIG
149	155	closed	protein_state	The dashed lines and arrows, respectively, highlight the slight extension in length, and twist in shape that occurs with transition from open to the closed state.	FIG
4	8	open	protein_state	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
28	38	unliganded	protein_state	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
39	42	GDH	protein_type	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
56	62	closed	protein_state	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
73	77	NADH	chemical	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
79	82	GTP	chemical	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
88	97	glutamate	chemical	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
98	103	bound	protein_state	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
109	122	Superposition	experimental_method	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
126	136	structures	evidence	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
141	147	closed	protein_state	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
152	156	open	protein_state	The open state shown is for unliganded GDH, whereas the closed state has NADH, GTP, and glutamate bound. (B) Superposition of structures for closed and open conformations, along with a series of possible intermediate conformations along the trajectory that serve to illustrate the extent of change in structure across different regions of the protein.	FIG
24	30	closed	protein_state	The transition between “closed” andopen” states of GDH is modulated by two allosteric sites in each protomer (Fig. 1A), which are differentially bound by GTP (an inhibitor) and ADP (an activator).	INTRO
37	41	open	protein_state	The transition between “closed” andopen” states of GDH is modulated by two allosteric sites in each protomer (Fig. 1A), which are differentially bound by GTP (an inhibitor) and ADP (an activator).	INTRO
53	56	GDH	protein_type	The transition between “closed” andopen” states of GDH is modulated by two allosteric sites in each protomer (Fig. 1A), which are differentially bound by GTP (an inhibitor) and ADP (an activator).	INTRO
77	93	allosteric sites	site	The transition between “closed” andopen” states of GDH is modulated by two allosteric sites in each protomer (Fig. 1A), which are differentially bound by GTP (an inhibitor) and ADP (an activator).	INTRO
102	110	protomer	oligomeric_state	The transition between “closed” andopen” states of GDH is modulated by two allosteric sites in each protomer (Fig. 1A), which are differentially bound by GTP (an inhibitor) and ADP (an activator).	INTRO
147	155	bound by	protein_state	The transition between “closed” andopen” states of GDH is modulated by two allosteric sites in each protomer (Fig. 1A), which are differentially bound by GTP (an inhibitor) and ADP (an activator).	INTRO
156	159	GTP	chemical	The transition between “closed” andopen” states of GDH is modulated by two allosteric sites in each protomer (Fig. 1A), which are differentially bound by GTP (an inhibitor) and ADP (an activator).	INTRO
179	182	ADP	chemical	The transition between “closed” andopen” states of GDH is modulated by two allosteric sites in each protomer (Fig. 1A), which are differentially bound by GTP (an inhibitor) and ADP (an activator).	INTRO
44	47	GDH	protein_type	These allosteric modulators tightly control GDH function in vivo.	INTRO
42	53	pivot helix	structure_element	In the first site, which sits next to the pivot helix at the base of the antenna (the “GTP binding site”), GTP binding is known to act as an inhibitor, preventing release of the reaction product from the catalytic site by stabilizing the closed conformation of the catalytic cleft.	INTRO
73	80	antenna	structure_element	In the first site, which sits next to the pivot helix at the base of the antenna (the “GTP binding site”), GTP binding is known to act as an inhibitor, preventing release of the reaction product from the catalytic site by stabilizing the closed conformation of the catalytic cleft.	INTRO
87	103	GTP binding site	site	In the first site, which sits next to the pivot helix at the base of the antenna (the “GTP binding site”), GTP binding is known to act as an inhibitor, preventing release of the reaction product from the catalytic site by stabilizing the closed conformation of the catalytic cleft.	INTRO
107	110	GTP	chemical	In the first site, which sits next to the pivot helix at the base of the antenna (the “GTP binding site”), GTP binding is known to act as an inhibitor, preventing release of the reaction product from the catalytic site by stabilizing the closed conformation of the catalytic cleft.	INTRO
204	218	catalytic site	site	In the first site, which sits next to the pivot helix at the base of the antenna (the “GTP binding site”), GTP binding is known to act as an inhibitor, preventing release of the reaction product from the catalytic site by stabilizing the closed conformation of the catalytic cleft.	INTRO
238	244	closed	protein_state	In the first site, which sits next to the pivot helix at the base of the antenna (the “GTP binding site”), GTP binding is known to act as an inhibitor, preventing release of the reaction product from the catalytic site by stabilizing the closed conformation of the catalytic cleft.	INTRO
265	280	catalytic cleft	site	In the first site, which sits next to the pivot helix at the base of the antenna (the “GTP binding site”), GTP binding is known to act as an inhibitor, preventing release of the reaction product from the catalytic site by stabilizing the closed conformation of the catalytic cleft.	INTRO
15	30	regulatory site	site	In the second “regulatory site”, which is situated near the pivot helix between adjacent protomers, ADP acts as an activator of enzymatic activity, presumably by hastening the opening of the catalytic cleft that leads to the release of the reaction product.	INTRO
60	71	pivot helix	structure_element	In the second “regulatory site”, which is situated near the pivot helix between adjacent protomers, ADP acts as an activator of enzymatic activity, presumably by hastening the opening of the catalytic cleft that leads to the release of the reaction product.	INTRO
89	98	protomers	oligomeric_state	In the second “regulatory site”, which is situated near the pivot helix between adjacent protomers, ADP acts as an activator of enzymatic activity, presumably by hastening the opening of the catalytic cleft that leads to the release of the reaction product.	INTRO
100	103	ADP	chemical	In the second “regulatory site”, which is situated near the pivot helix between adjacent protomers, ADP acts as an activator of enzymatic activity, presumably by hastening the opening of the catalytic cleft that leads to the release of the reaction product.	INTRO
191	206	catalytic cleft	site	In the second “regulatory site”, which is situated near the pivot helix between adjacent protomers, ADP acts as an activator of enzymatic activity, presumably by hastening the opening of the catalytic cleft that leads to the release of the reaction product.	INTRO
56	60	NADH	chemical	Interestingly, it has also been shown that the coenzyme NADH can bind to the regulatory site (also bound by the activator ADP), exerting a converse, inhibitory effect on GDH product release, although the role this may play in vivo is not entirely clear.	INTRO
77	92	regulatory site	site	Interestingly, it has also been shown that the coenzyme NADH can bind to the regulatory site (also bound by the activator ADP), exerting a converse, inhibitory effect on GDH product release, although the role this may play in vivo is not entirely clear.	INTRO
99	107	bound by	protein_state	Interestingly, it has also been shown that the coenzyme NADH can bind to the regulatory site (also bound by the activator ADP), exerting a converse, inhibitory effect on GDH product release, although the role this may play in vivo is not entirely clear.	INTRO
122	125	ADP	chemical	Interestingly, it has also been shown that the coenzyme NADH can bind to the regulatory site (also bound by the activator ADP), exerting a converse, inhibitory effect on GDH product release, although the role this may play in vivo is not entirely clear.	INTRO
170	173	GDH	protein_type	Interestingly, it has also been shown that the coenzyme NADH can bind to the regulatory site (also bound by the activator ADP), exerting a converse, inhibitory effect on GDH product release, although the role this may play in vivo is not entirely clear.	INTRO
28	46	crystal structures	evidence	Although there are numerous crystal structures available for GDH in complex with cofactors and nucleotides, they are limited to the combinations that have been amenable to crystallization.	INTRO
61	64	GDH	protein_type	Although there are numerous crystal structures available for GDH in complex with cofactors and nucleotides, they are limited to the combinations that have been amenable to crystallization.	INTRO
65	80	in complex with	protein_state	Although there are numerous crystal structures available for GDH in complex with cofactors and nucleotides, they are limited to the combinations that have been amenable to crystallization.	INTRO
172	187	crystallization	experimental_method	Although there are numerous crystal structures available for GDH in complex with cofactors and nucleotides, they are limited to the combinations that have been amenable to crystallization.	INTRO
11	16	X-ray	experimental_method	Nearly all X-ray structures of mammalian GDH are in the closed conformation, and the few structures that are in the open conformation are at lower resolution (Table 1).	INTRO
17	27	structures	evidence	Nearly all X-ray structures of mammalian GDH are in the closed conformation, and the few structures that are in the open conformation are at lower resolution (Table 1).	INTRO
31	40	mammalian	taxonomy_domain	Nearly all X-ray structures of mammalian GDH are in the closed conformation, and the few structures that are in the open conformation are at lower resolution (Table 1).	INTRO
41	44	GDH	protein_type	Nearly all X-ray structures of mammalian GDH are in the closed conformation, and the few structures that are in the open conformation are at lower resolution (Table 1).	INTRO
56	62	closed	protein_state	Nearly all X-ray structures of mammalian GDH are in the closed conformation, and the few structures that are in the open conformation are at lower resolution (Table 1).	INTRO
89	99	structures	evidence	Nearly all X-ray structures of mammalian GDH are in the closed conformation, and the few structures that are in the open conformation are at lower resolution (Table 1).	INTRO
116	120	open	protein_state	Nearly all X-ray structures of mammalian GDH are in the closed conformation, and the few structures that are in the open conformation are at lower resolution (Table 1).	INTRO
9	19	structures	evidence	Of those structures in the closed conformation, most include NAD[P]H, GTP, and glutamate (or, alternately, NAD+, GTP, and α-ketoglutarate).	INTRO
27	33	closed	protein_state	Of those structures in the closed conformation, most include NAD[P]H, GTP, and glutamate (or, alternately, NAD+, GTP, and α-ketoglutarate).	INTRO
61	68	NAD[P]H	chemical	Of those structures in the closed conformation, most include NAD[P]H, GTP, and glutamate (or, alternately, NAD+, GTP, and α-ketoglutarate).	INTRO
70	73	GTP	chemical	Of those structures in the closed conformation, most include NAD[P]H, GTP, and glutamate (or, alternately, NAD+, GTP, and α-ketoglutarate).	INTRO
79	88	glutamate	chemical	Of those structures in the closed conformation, most include NAD[P]H, GTP, and glutamate (or, alternately, NAD+, GTP, and α-ketoglutarate).	INTRO
107	111	NAD+	chemical	Of those structures in the closed conformation, most include NAD[P]H, GTP, and glutamate (or, alternately, NAD+, GTP, and α-ketoglutarate).	INTRO
113	116	GTP	chemical	Of those structures in the closed conformation, most include NAD[P]H, GTP, and glutamate (or, alternately, NAD+, GTP, and α-ketoglutarate).	INTRO
122	137	α-ketoglutarate	chemical	Of those structures in the closed conformation, most include NAD[P]H, GTP, and glutamate (or, alternately, NAD+, GTP, and α-ketoglutarate).	INTRO
37	40	GTP	chemical	However, the effects of coenzyme and GTP, bound alone or in concert in the absence of glutamate, have not been analyzed by crystallographic methods.	INTRO
42	53	bound alone	protein_state	However, the effects of coenzyme and GTP, bound alone or in concert in the absence of glutamate, have not been analyzed by crystallographic methods.	INTRO
75	85	absence of	protein_state	However, the effects of coenzyme and GTP, bound alone or in concert in the absence of glutamate, have not been analyzed by crystallographic methods.	INTRO
86	95	glutamate	chemical	However, the effects of coenzyme and GTP, bound alone or in concert in the absence of glutamate, have not been analyzed by crystallographic methods.	INTRO
16	56	single-particle cryo-electron microscopy	experimental_method	Here, we report single-particle cryo-electron microscopy (cryo-EM) studies that show that under these conditions enzyme complexes coexist in both closed and open conformations.	INTRO
58	65	cryo-EM	experimental_method	Here, we report single-particle cryo-electron microscopy (cryo-EM) studies that show that under these conditions enzyme complexes coexist in both closed and open conformations.	INTRO
146	152	closed	protein_state	Here, we report single-particle cryo-electron microscopy (cryo-EM) studies that show that under these conditions enzyme complexes coexist in both closed and open conformations.	INTRO
157	161	open	protein_state	Here, we report single-particle cryo-electron microscopy (cryo-EM) studies that show that under these conditions enzyme complexes coexist in both closed and open conformations.	INTRO
17	27	structures	evidence	We show that the structures in both states can be resolved at near-atomic resolution, suggesting a molecular mechanism for synergistic inhibition of GDH by NADH and GTP (see Table 2 for detailed information on all cryo-EM-derived structures that we report in this work).	INTRO
149	152	GDH	protein_type	We show that the structures in both states can be resolved at near-atomic resolution, suggesting a molecular mechanism for synergistic inhibition of GDH by NADH and GTP (see Table 2 for detailed information on all cryo-EM-derived structures that we report in this work).	INTRO
156	160	NADH	chemical	We show that the structures in both states can be resolved at near-atomic resolution, suggesting a molecular mechanism for synergistic inhibition of GDH by NADH and GTP (see Table 2 for detailed information on all cryo-EM-derived structures that we report in this work).	INTRO
165	168	GTP	chemical	We show that the structures in both states can be resolved at near-atomic resolution, suggesting a molecular mechanism for synergistic inhibition of GDH by NADH and GTP (see Table 2 for detailed information on all cryo-EM-derived structures that we report in this work).	INTRO
214	221	cryo-EM	experimental_method	We show that the structures in both states can be resolved at near-atomic resolution, suggesting a molecular mechanism for synergistic inhibition of GDH by NADH and GTP (see Table 2 for detailed information on all cryo-EM-derived structures that we report in this work).	INTRO
230	240	structures	evidence	We show that the structures in both states can be resolved at near-atomic resolution, suggesting a molecular mechanism for synergistic inhibition of GDH by NADH and GTP (see Table 2 for detailed information on all cryo-EM-derived structures that we report in this work).	INTRO
0	5	X-ray	experimental_method	X-ray structures of mammalian GDH reported in both the open and closed conformations	TABLE
6	16	structures	evidence	X-ray structures of mammalian GDH reported in both the open and closed conformations	TABLE
20	29	mammalian	taxonomy_domain	X-ray structures of mammalian GDH reported in both the open and closed conformations	TABLE
30	33	GDH	protein_type	X-ray structures of mammalian GDH reported in both the open and closed conformations	TABLE
55	59	open	protein_state	X-ray structures of mammalian GDH reported in both the open and closed conformations	TABLE
64	70	closed	protein_state	X-ray structures of mammalian GDH reported in both the open and closed conformations	TABLE
0	3	GDH	protein	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
45	47	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
48	52	NADH	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
55	58	GLU	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
61	64	GTP	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
70	76	Closed	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
83	85	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
86	89	Glu	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
91	94	GTP	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
96	101	NADPH	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
122	128	Closed	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
135	137	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
138	141	Glu	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
143	148	NADPH	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
150	153	GTP	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
168	174	Closed	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
181	183	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
184	187	apo	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
193	197	Open	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
204	206	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
207	212	NADPH	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
214	223	glutamate	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
229	232	GTP	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
238	244	Closed	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
251	253	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
254	259	NADPH	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
262	265	GLU	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
268	271	GTP	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
274	278	Zinc	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
284	290	Closed	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
298	300	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
301	306	NADPH	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
308	311	Glu	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
313	316	GTP	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
339	345	Closed	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
350	352	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
353	356	NAD	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
358	361	PO4	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
367	381	2-oxoglutarate	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
387	393	Closed	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
400	402	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
403	408	NADPH	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
411	414	GLU	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
425	431	Closed	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
445	451	mutant	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
452	455	apo	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
461	465	Open	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
472	474	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
475	478	apo	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
484	488	Open	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
495	497	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
498	501	ADP	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
507	511	Open	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
518	520	WT	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
521	526	NADPH	chemical	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
564	568	Open	protein_state	"GDH	Ligands	PDB ID	Conformation	Resolution	 	WT	NADH + GLU + GTP	3MW9	Closed	2.4	 	WT	Glu, GTP, NADPH, and Bithionol	3ETD	Closed	2.5	 	WT	Glu, NADPH, GTP + GW5074	3ETG	Closed	2.5	 	WT	apo	1L1F	Open	2.7	 	WT	NADPH, glutamate, and GTP	1HWZ	Closed	2.8	 	WT	NADPH + GLU + GTP + Zinc	3MVQ	Closed	2.94	 	WT	NADPH, Glu, GTP, Hexachlorophene	3ETE	Closed	3	 	WT	NAD, PO4, and 2-oxoglutarate	1HWY	Closed	3.2	 	WT	NADPH + GLU + Eu	3MVO	Closed	3.23	 	R463A mutant	apo	1NR1	Open	3.3	 	WT	apo	1NR7	Open	3.3	 	WT	ADP	1NQT	Open	3.5	 	WT	NADPH and Epicatechin-3-gallate (Ecg)	3QMU	Open	3.62	 	"	TABLE
0	7	Cryo-EM	experimental_method	Cryo-EM structures of mammalian GDH determined for this study	TABLE
8	18	structures	evidence	Cryo-EM structures of mammalian GDH determined for this study	TABLE
22	31	mammalian	taxonomy_domain	Cryo-EM structures of mammalian GDH determined for this study	TABLE
32	35	GDH	protein_type	Cryo-EM structures of mammalian GDH determined for this study	TABLE
0	3	GDH	protein_type	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
63	65	WT	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
66	69	apo	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
84	88	Open	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
102	104	WT	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
105	108	GTP	chemical	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
123	127	Open	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
141	143	WT	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
144	148	NADH	chemical	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
163	167	Open	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
181	183	WT	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
184	188	NADH	chemical	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
203	209	Closed	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
223	225	WT	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
226	230	NADH	chemical	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
233	236	GTP	chemical	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
251	255	Open	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
269	271	WT	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
272	276	NADH	chemical	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
279	282	GTP	chemical	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
297	303	Closed	protein_state	"GDH	Ligands	EMDB ID	PDB ID	Conformation	Resolution	Particles	 	WT	apo	EMD-6630	3JCZ	Open	3.26	22462	 	WT	GTP	EMD-6631	3JD0	Open	3.47	39439	 	WT	NADH	EMD-6635	3JD2	Open	3.27	34716	 	WT	NADH	EMD-6634	3JD1	Closed	3.27	34926	 	WT	NADH + GTP	EMD-6632	3JD3	Open	3.55	14793	 	WT	NADH + GTP	EMD-6633	3JD4	Closed	3.40	20429	 	"	TABLE
43	46	apo	protein_state	To explore the conformational landscape of apo-GDH, we first determined its structure in the absence of any added ligands (Supplemental Fig. 1, Fig. 2, A–C).	RESULTS
47	50	GDH	protein	To explore the conformational landscape of apo-GDH, we first determined its structure in the absence of any added ligands (Supplemental Fig. 1, Fig. 2, A–C).	RESULTS
76	85	structure	evidence	To explore the conformational landscape of apo-GDH, we first determined its structure in the absence of any added ligands (Supplemental Fig. 1, Fig. 2, A–C).	RESULTS
93	103	absence of	protein_state	To explore the conformational landscape of apo-GDH, we first determined its structure in the absence of any added ligands (Supplemental Fig. 1, Fig. 2, A–C).	RESULTS
4	15	density map	evidence	The density map, refined to an average resolution of3.0 Å (Supplemental Fig. 2), is in the open conformation and closely matches the model of unliganded GDH derived by X-ray crystallography at 3.3 Å resolution (PDB ID 1NR7).	RESULTS
93	97	open	protein_state	The density map, refined to an average resolution of3.0 Å (Supplemental Fig. 2), is in the open conformation and closely matches the model of unliganded GDH derived by X-ray crystallography at 3.3 Å resolution (PDB ID 1NR7).	RESULTS
144	154	unliganded	protein_state	The density map, refined to an average resolution of3.0 Å (Supplemental Fig. 2), is in the open conformation and closely matches the model of unliganded GDH derived by X-ray crystallography at 3.3 Å resolution (PDB ID 1NR7).	RESULTS
155	158	GDH	protein	The density map, refined to an average resolution of3.0 Å (Supplemental Fig. 2), is in the open conformation and closely matches the model of unliganded GDH derived by X-ray crystallography at 3.3 Å resolution (PDB ID 1NR7).	RESULTS
170	191	X-ray crystallography	experimental_method	The density map, refined to an average resolution of3.0 Å (Supplemental Fig. 2), is in the open conformation and closely matches the model of unliganded GDH derived by X-ray crystallography at 3.3 Å resolution (PDB ID 1NR7).	RESULTS
81	87	ResMap	experimental_method	The variation in local resolution from the core to the periphery, as reported by ResMap (Supplemental Fig. 3D), is consistent with the B-factor gradient observed in the crystal structure (Supplemental Fig. 3A).	RESULTS
135	152	B-factor gradient	evidence	The variation in local resolution from the core to the periphery, as reported by ResMap (Supplemental Fig. 3D), is consistent with the B-factor gradient observed in the crystal structure (Supplemental Fig. 3A).	RESULTS
169	186	crystal structure	evidence	The variation in local resolution from the core to the periphery, as reported by ResMap (Supplemental Fig. 3D), is consistent with the B-factor gradient observed in the crystal structure (Supplemental Fig. 3A).	RESULTS
64	68	open	protein_state	Extensive classification without imposing symmetry yielded only open structures and failed to detect any closed catalytic cleft in the unliganded enzyme, suggesting that all six protomers are in the open conformation.	RESULTS
69	79	structures	evidence	Extensive classification without imposing symmetry yielded only open structures and failed to detect any closed catalytic cleft in the unliganded enzyme, suggesting that all six protomers are in the open conformation.	RESULTS
105	111	closed	protein_state	Extensive classification without imposing symmetry yielded only open structures and failed to detect any closed catalytic cleft in the unliganded enzyme, suggesting that all six protomers are in the open conformation.	RESULTS
112	127	catalytic cleft	site	Extensive classification without imposing symmetry yielded only open structures and failed to detect any closed catalytic cleft in the unliganded enzyme, suggesting that all six protomers are in the open conformation.	RESULTS
135	145	unliganded	protein_state	Extensive classification without imposing symmetry yielded only open structures and failed to detect any closed catalytic cleft in the unliganded enzyme, suggesting that all six protomers are in the open conformation.	RESULTS
178	187	protomers	oligomeric_state	Extensive classification without imposing symmetry yielded only open structures and failed to detect any closed catalytic cleft in the unliganded enzyme, suggesting that all six protomers are in the open conformation.	RESULTS
199	203	open	protein_state	Extensive classification without imposing symmetry yielded only open structures and failed to detect any closed catalytic cleft in the unliganded enzyme, suggesting that all six protomers are in the open conformation.	RESULTS
37	42	loops	structure_element	Consistent with this conclusion, the loops connecting the β-strands of the Rossmann fold are well-defined (Fig. 2B), implying that there is little movement at the NBD, as the transition between closed and open states is associated with NBD movement (Fig. 1B).	RESULTS
58	67	β-strands	structure_element	Consistent with this conclusion, the loops connecting the β-strands of the Rossmann fold are well-defined (Fig. 2B), implying that there is little movement at the NBD, as the transition between closed and open states is associated with NBD movement (Fig. 1B).	RESULTS
75	88	Rossmann fold	structure_element	Consistent with this conclusion, the loops connecting the β-strands of the Rossmann fold are well-defined (Fig. 2B), implying that there is little movement at the NBD, as the transition between closed and open states is associated with NBD movement (Fig. 1B).	RESULTS
163	166	NBD	structure_element	Consistent with this conclusion, the loops connecting the β-strands of the Rossmann fold are well-defined (Fig. 2B), implying that there is little movement at the NBD, as the transition between closed and open states is associated with NBD movement (Fig. 1B).	RESULTS
194	200	closed	protein_state	Consistent with this conclusion, the loops connecting the β-strands of the Rossmann fold are well-defined (Fig. 2B), implying that there is little movement at the NBD, as the transition between closed and open states is associated with NBD movement (Fig. 1B).	RESULTS
205	209	open	protein_state	Consistent with this conclusion, the loops connecting the β-strands of the Rossmann fold are well-defined (Fig. 2B), implying that there is little movement at the NBD, as the transition between closed and open states is associated with NBD movement (Fig. 1B).	RESULTS
236	239	NBD	structure_element	Consistent with this conclusion, the loops connecting the β-strands of the Rossmann fold are well-defined (Fig. 2B), implying that there is little movement at the NBD, as the transition between closed and open states is associated with NBD movement (Fig. 1B).	RESULTS
0	7	Cryo-EM	experimental_method	Cryo-EM structures of GDH in unliganded and NADH-bound states. (A) Refined cryo-EM map of unliganded GDH at3 Å resolution.	FIG
8	18	structures	evidence	Cryo-EM structures of GDH in unliganded and NADH-bound states. (A) Refined cryo-EM map of unliganded GDH at3 Å resolution.	FIG
22	25	GDH	protein	Cryo-EM structures of GDH in unliganded and NADH-bound states. (A) Refined cryo-EM map of unliganded GDH at3 Å resolution.	FIG
29	39	unliganded	protein_state	Cryo-EM structures of GDH in unliganded and NADH-bound states. (A) Refined cryo-EM map of unliganded GDH at3 Å resolution.	FIG
44	54	NADH-bound	protein_state	Cryo-EM structures of GDH in unliganded and NADH-bound states. (A) Refined cryo-EM map of unliganded GDH at3 Å resolution.	FIG
75	82	cryo-EM	experimental_method	Cryo-EM structures of GDH in unliganded and NADH-bound states. (A) Refined cryo-EM map of unliganded GDH at3 Å resolution.	FIG
83	86	map	evidence	Cryo-EM structures of GDH in unliganded and NADH-bound states. (A) Refined cryo-EM map of unliganded GDH at3 Å resolution.	FIG
90	100	unliganded	protein_state	Cryo-EM structures of GDH in unliganded and NADH-bound states. (A) Refined cryo-EM map of unliganded GDH at3 Å resolution.	FIG
101	104	GDH	protein	Cryo-EM structures of GDH in unliganded and NADH-bound states. (A) Refined cryo-EM map of unliganded GDH at3 Å resolution.	FIG
23	34	density map	evidence	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
67	99	Rossmann nucleotide binding fold	structure_element	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
105	130	pivot and antenna helices	structure_element	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
142	152	unliganded	protein_state	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
153	156	GDH	protein	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
157	160	map	evidence	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
166	173	Cryo-EM	experimental_method	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
182	194	density maps	evidence	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
250	253	GDH	protein	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
257	265	bound to	protein_state	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
279	283	NADH	chemical	(B, C) Illustration of density map in the regions that contain the Rossmann nucleotide binding fold (B), pivot and antenna helices (C) in the unliganded GDH map. (D) Cryo-EM-derived density maps for two coexisting conformations that are present when GDH is bound to the cofactor NADH.	FIG
5	13	protomer	oligomeric_state	Each protomer is shown in a different color and densities for NADH bound in both regulatory (red) and catalytic (purple) sites on one protomer are indicated.	FIG
48	57	densities	evidence	Each protomer is shown in a different color and densities for NADH bound in both regulatory (red) and catalytic (purple) sites on one protomer are indicated.	FIG
62	66	NADH	chemical	Each protomer is shown in a different color and densities for NADH bound in both regulatory (red) and catalytic (purple) sites on one protomer are indicated.	FIG
67	75	bound in	protein_state	Each protomer is shown in a different color and densities for NADH bound in both regulatory (red) and catalytic (purple) sites on one protomer are indicated.	FIG
81	91	regulatory	site	Each protomer is shown in a different color and densities for NADH bound in both regulatory (red) and catalytic (purple) sites on one protomer are indicated.	FIG
102	111	catalytic	site	Each protomer is shown in a different color and densities for NADH bound in both regulatory (red) and catalytic (purple) sites on one protomer are indicated.	FIG
121	126	sites	site	Each protomer is shown in a different color and densities for NADH bound in both regulatory (red) and catalytic (purple) sites on one protomer are indicated.	FIG
134	142	protomer	oligomeric_state	Each protomer is shown in a different color and densities for NADH bound in both regulatory (red) and catalytic (purple) sites on one protomer are indicated.	FIG
99	103	open	protein_state	The overall quaternary structures of the two conformations are essentially the same as that of the open and closed states observed by X-ray crystallography.	FIG
108	114	closed	protein_state	The overall quaternary structures of the two conformations are essentially the same as that of the open and closed states observed by X-ray crystallography.	FIG
134	155	X-ray crystallography	experimental_method	The overall quaternary structures of the two conformations are essentially the same as that of the open and closed states observed by X-ray crystallography.	FIG
5	8	GDH	protein	When GDH is bound to NADH, GTP, and glutamate, the enzyme adopts a closed conformation; this “abortive complex” has been determined to 2.4-Å resolution by X-ray crystallography (PDB 3MW9).	RESULTS
12	20	bound to	protein_state	When GDH is bound to NADH, GTP, and glutamate, the enzyme adopts a closed conformation; this “abortive complex” has been determined to 2.4-Å resolution by X-ray crystallography (PDB 3MW9).	RESULTS
21	25	NADH	chemical	When GDH is bound to NADH, GTP, and glutamate, the enzyme adopts a closed conformation; this “abortive complex” has been determined to 2.4-Å resolution by X-ray crystallography (PDB 3MW9).	RESULTS
27	30	GTP	chemical	When GDH is bound to NADH, GTP, and glutamate, the enzyme adopts a closed conformation; this “abortive complex” has been determined to 2.4-Å resolution by X-ray crystallography (PDB 3MW9).	RESULTS
36	45	glutamate	chemical	When GDH is bound to NADH, GTP, and glutamate, the enzyme adopts a closed conformation; this “abortive complex” has been determined to 2.4-Å resolution by X-ray crystallography (PDB 3MW9).	RESULTS
67	73	closed	protein_state	When GDH is bound to NADH, GTP, and glutamate, the enzyme adopts a closed conformation; this “abortive complex” has been determined to 2.4-Å resolution by X-ray crystallography (PDB 3MW9).	RESULTS
155	176	X-ray crystallography	experimental_method	When GDH is bound to NADH, GTP, and glutamate, the enzyme adopts a closed conformation; this “abortive complex” has been determined to 2.4-Å resolution by X-ray crystallography (PDB 3MW9).	RESULTS
9	27	crystal structures	evidence	However, crystal structures of GDH bound only to NADH or to GTP have not yet been reported.	RESULTS
31	34	GDH	protein	However, crystal structures of GDH bound only to NADH or to GTP have not yet been reported.	RESULTS
35	48	bound only to	protein_state	However, crystal structures of GDH bound only to NADH or to GTP have not yet been reported.	RESULTS
49	53	NADH	chemical	However, crystal structures of GDH bound only to NADH or to GTP have not yet been reported.	RESULTS
57	59	to	protein_state	However, crystal structures of GDH bound only to NADH or to GTP have not yet been reported.	RESULTS
60	63	GTP	chemical	However, crystal structures of GDH bound only to NADH or to GTP have not yet been reported.	RESULTS
22	26	NADH	chemical	To test the effect of NADH binding on GDH conformation in solution, we determined the structure of this binary complex using cryo-EM methods combined with three-dimensional classification.	RESULTS
38	41	GDH	protein	To test the effect of NADH binding on GDH conformation in solution, we determined the structure of this binary complex using cryo-EM methods combined with three-dimensional classification.	RESULTS
86	95	structure	evidence	To test the effect of NADH binding on GDH conformation in solution, we determined the structure of this binary complex using cryo-EM methods combined with three-dimensional classification.	RESULTS
125	132	cryo-EM	experimental_method	To test the effect of NADH binding on GDH conformation in solution, we determined the structure of this binary complex using cryo-EM methods combined with three-dimensional classification.	RESULTS
155	187	three-dimensional classification	experimental_method	To test the effect of NADH binding on GDH conformation in solution, we determined the structure of this binary complex using cryo-EM methods combined with three-dimensional classification.	RESULTS
46	50	open	protein_state	Two dominant conformational states, in an all open or all closed conformation were detected, segregated (Fig. 2D), and further refined to near-atomic resolution (∼3.3 Å; Supplemental Fig. 2).	RESULTS
58	64	closed	protein_state	Two dominant conformational states, in an all open or all closed conformation were detected, segregated (Fig. 2D), and further refined to near-atomic resolution (∼3.3 Å; Supplemental Fig. 2).	RESULTS
0	9	Densities	evidence	Densities for 12 molecules of bound NADH were identified in maps of both open and closed states (Supplemental Fig. 4).	RESULTS
30	35	bound	protein_state	Densities for 12 molecules of bound NADH were identified in maps of both open and closed states (Supplemental Fig. 4).	RESULTS
36	40	NADH	chemical	Densities for 12 molecules of bound NADH were identified in maps of both open and closed states (Supplemental Fig. 4).	RESULTS
60	64	maps	evidence	Densities for 12 molecules of bound NADH were identified in maps of both open and closed states (Supplemental Fig. 4).	RESULTS
73	77	open	protein_state	Densities for 12 molecules of bound NADH were identified in maps of both open and closed states (Supplemental Fig. 4).	RESULTS
82	88	closed	protein_state	Densities for 12 molecules of bound NADH were identified in maps of both open and closed states (Supplemental Fig. 4).	RESULTS
4	14	NADH-bound	protein_state	The NADH-bound closed conformation matches the structure of the quaternary complex observed by X-ray crystallography, with the exception that density corresponding to GTP and glutamate was absent in the cryo-EM-derived map.	RESULTS
15	21	closed	protein_state	The NADH-bound closed conformation matches the structure of the quaternary complex observed by X-ray crystallography, with the exception that density corresponding to GTP and glutamate was absent in the cryo-EM-derived map.	RESULTS
47	56	structure	evidence	The NADH-bound closed conformation matches the structure of the quaternary complex observed by X-ray crystallography, with the exception that density corresponding to GTP and glutamate was absent in the cryo-EM-derived map.	RESULTS
95	116	X-ray crystallography	experimental_method	The NADH-bound closed conformation matches the structure of the quaternary complex observed by X-ray crystallography, with the exception that density corresponding to GTP and glutamate was absent in the cryo-EM-derived map.	RESULTS
142	149	density	evidence	The NADH-bound closed conformation matches the structure of the quaternary complex observed by X-ray crystallography, with the exception that density corresponding to GTP and glutamate was absent in the cryo-EM-derived map.	RESULTS
167	170	GTP	chemical	The NADH-bound closed conformation matches the structure of the quaternary complex observed by X-ray crystallography, with the exception that density corresponding to GTP and glutamate was absent in the cryo-EM-derived map.	RESULTS
175	184	glutamate	chemical	The NADH-bound closed conformation matches the structure of the quaternary complex observed by X-ray crystallography, with the exception that density corresponding to GTP and glutamate was absent in the cryo-EM-derived map.	RESULTS
203	210	cryo-EM	experimental_method	The NADH-bound closed conformation matches the structure of the quaternary complex observed by X-ray crystallography, with the exception that density corresponding to GTP and glutamate was absent in the cryo-EM-derived map.	RESULTS
219	222	map	evidence	The NADH-bound closed conformation matches the structure of the quaternary complex observed by X-ray crystallography, with the exception that density corresponding to GTP and glutamate was absent in the cryo-EM-derived map.	RESULTS
18	28	NADH-bound	protein_state	Comparison of the NADH-bound closed conformation to the NADH-bound open conformation shows that, as expected, the catalytic cleft is closed and the NBDs are displaced toward the equatorial plane, accompanied by a rotation of the pivot helix by7°, concomitant with a large conformational change in the antennae domains (Figs. 1 and 2D).	RESULTS
29	35	closed	protein_state	Comparison of the NADH-bound closed conformation to the NADH-bound open conformation shows that, as expected, the catalytic cleft is closed and the NBDs are displaced toward the equatorial plane, accompanied by a rotation of the pivot helix by7°, concomitant with a large conformational change in the antennae domains (Figs. 1 and 2D).	RESULTS
56	66	NADH-bound	protein_state	Comparison of the NADH-bound closed conformation to the NADH-bound open conformation shows that, as expected, the catalytic cleft is closed and the NBDs are displaced toward the equatorial plane, accompanied by a rotation of the pivot helix by7°, concomitant with a large conformational change in the antennae domains (Figs. 1 and 2D).	RESULTS
67	71	open	protein_state	Comparison of the NADH-bound closed conformation to the NADH-bound open conformation shows that, as expected, the catalytic cleft is closed and the NBDs are displaced toward the equatorial plane, accompanied by a rotation of the pivot helix by7°, concomitant with a large conformational change in the antennae domains (Figs. 1 and 2D).	RESULTS
114	129	catalytic cleft	site	Comparison of the NADH-bound closed conformation to the NADH-bound open conformation shows that, as expected, the catalytic cleft is closed and the NBDs are displaced toward the equatorial plane, accompanied by a rotation of the pivot helix by7°, concomitant with a large conformational change in the antennae domains (Figs. 1 and 2D).	RESULTS
133	139	closed	protein_state	Comparison of the NADH-bound closed conformation to the NADH-bound open conformation shows that, as expected, the catalytic cleft is closed and the NBDs are displaced toward the equatorial plane, accompanied by a rotation of the pivot helix by7°, concomitant with a large conformational change in the antennae domains (Figs. 1 and 2D).	RESULTS
148	152	NBDs	structure_element	Comparison of the NADH-bound closed conformation to the NADH-bound open conformation shows that, as expected, the catalytic cleft is closed and the NBDs are displaced toward the equatorial plane, accompanied by a rotation of the pivot helix by7°, concomitant with a large conformational change in the antennae domains (Figs. 1 and 2D).	RESULTS
229	240	pivot helix	structure_element	Comparison of the NADH-bound closed conformation to the NADH-bound open conformation shows that, as expected, the catalytic cleft is closed and the NBDs are displaced toward the equatorial plane, accompanied by a rotation of the pivot helix by7°, concomitant with a large conformational change in the antennae domains (Figs. 1 and 2D).	RESULTS
303	311	antennae	structure_element	Comparison of the NADH-bound closed conformation to the NADH-bound open conformation shows that, as expected, the catalytic cleft is closed and the NBDs are displaced toward the equatorial plane, accompanied by a rotation of the pivot helix by7°, concomitant with a large conformational change in the antennae domains (Figs. 1 and 2D).	RESULTS
21	31	NADH-bound	protein_state	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
32	36	open	protein_state	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
41	47	closed	protein_state	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
94	101	helix 5	structure_element	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
112	119	171186	residue_range	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
152	160	β-sheets	structure_element	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
218	223	5797	residue_range	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
225	232	122130	residue_range	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
238	247	α-helix 2	structure_element	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
258	263	3654	residue_range	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
298	303	helix	structure_element	A comparison between NADH-bound open and closed conformations also involves a displacement of helix 5 (residues 171186), as well as a tilt of the core β-sheets relative to the equatorial plane of the enzyme (residues 5797, 122130) and α-helix 2 (residues 3654), and a bending of the N-terminal helix.	RESULTS
21	36	catalytic cleft	site	Thus, closure of the catalytic cleft is accompanied by a quaternary structural change that can be described as a global bending of the structure about an axis that runs parallel to the pivot helix, accompanied by an expansion of the core (Figs. 1A and 2D).	RESULTS
135	144	structure	evidence	Thus, closure of the catalytic cleft is accompanied by a quaternary structural change that can be described as a global bending of the structure about an axis that runs parallel to the pivot helix, accompanied by an expansion of the core (Figs. 1A and 2D).	RESULTS
185	196	pivot helix	structure_element	Thus, closure of the catalytic cleft is accompanied by a quaternary structural change that can be described as a global bending of the structure about an axis that runs parallel to the pivot helix, accompanied by an expansion of the core (Figs. 1A and 2D).	RESULTS
25	33	GDH/NADH	complex_assembly	Detailed analysis of the GDH/NADH structures shows that both the adenosine and nicotinamide moieties of NADH bind to the catalytic site within the NBD in nearly the same orientation in both the open and the closed states, and display closely comparable interactions with the Rossmann fold (Fig. 3, A and B).	RESULTS
34	44	structures	evidence	Detailed analysis of the GDH/NADH structures shows that both the adenosine and nicotinamide moieties of NADH bind to the catalytic site within the NBD in nearly the same orientation in both the open and the closed states, and display closely comparable interactions with the Rossmann fold (Fig. 3, A and B).	RESULTS
104	108	NADH	chemical	Detailed analysis of the GDH/NADH structures shows that both the adenosine and nicotinamide moieties of NADH bind to the catalytic site within the NBD in nearly the same orientation in both the open and the closed states, and display closely comparable interactions with the Rossmann fold (Fig. 3, A and B).	RESULTS
121	135	catalytic site	site	Detailed analysis of the GDH/NADH structures shows that both the adenosine and nicotinamide moieties of NADH bind to the catalytic site within the NBD in nearly the same orientation in both the open and the closed states, and display closely comparable interactions with the Rossmann fold (Fig. 3, A and B).	RESULTS
147	150	NBD	structure_element	Detailed analysis of the GDH/NADH structures shows that both the adenosine and nicotinamide moieties of NADH bind to the catalytic site within the NBD in nearly the same orientation in both the open and the closed states, and display closely comparable interactions with the Rossmann fold (Fig. 3, A and B).	RESULTS
194	198	open	protein_state	Detailed analysis of the GDH/NADH structures shows that both the adenosine and nicotinamide moieties of NADH bind to the catalytic site within the NBD in nearly the same orientation in both the open and the closed states, and display closely comparable interactions with the Rossmann fold (Fig. 3, A and B).	RESULTS
207	213	closed	protein_state	Detailed analysis of the GDH/NADH structures shows that both the adenosine and nicotinamide moieties of NADH bind to the catalytic site within the NBD in nearly the same orientation in both the open and the closed states, and display closely comparable interactions with the Rossmann fold (Fig. 3, A and B).	RESULTS
275	288	Rossmann fold	structure_element	Detailed analysis of the GDH/NADH structures shows that both the adenosine and nicotinamide moieties of NADH bind to the catalytic site within the NBD in nearly the same orientation in both the open and the closed states, and display closely comparable interactions with the Rossmann fold (Fig. 3, A and B).	RESULTS
7	22	regulatory site	site	At the regulatory site, where either ADP can bind as an activator or NADH can bind as an inhibitor, the binding of the adenine moiety of NADH is nearly identical between the two conformers.	RESULTS
37	40	ADP	chemical	At the regulatory site, where either ADP can bind as an activator or NADH can bind as an inhibitor, the binding of the adenine moiety of NADH is nearly identical between the two conformers.	RESULTS
69	73	NADH	chemical	At the regulatory site, where either ADP can bind as an activator or NADH can bind as an inhibitor, the binding of the adenine moiety of NADH is nearly identical between the two conformers.	RESULTS
137	141	NADH	chemical	At the regulatory site, where either ADP can bind as an activator or NADH can bind as an inhibitor, the binding of the adenine moiety of NADH is nearly identical between the two conformers.	RESULTS
7	13	closed	protein_state	In the closed state, the nicotinamide group is oriented toward the center of the hexamer, inserted into a narrow cavity between two adjacent subunits in the trimer.	RESULTS
81	88	hexamer	oligomeric_state	In the closed state, the nicotinamide group is oriented toward the center of the hexamer, inserted into a narrow cavity between two adjacent subunits in the trimer.	RESULTS
113	119	cavity	site	In the closed state, the nicotinamide group is oriented toward the center of the hexamer, inserted into a narrow cavity between two adjacent subunits in the trimer.	RESULTS
141	149	subunits	structure_element	In the closed state, the nicotinamide group is oriented toward the center of the hexamer, inserted into a narrow cavity between two adjacent subunits in the trimer.	RESULTS
157	163	trimer	oligomeric_state	In the closed state, the nicotinamide group is oriented toward the center of the hexamer, inserted into a narrow cavity between two adjacent subunits in the trimer.	RESULTS
41	45	NADH	chemical	There are extensive interactions between NADH and the residues lining this cavity, which may explain the well-defined density of this portion of NADH in the closed state.	RESULTS
75	81	cavity	site	There are extensive interactions between NADH and the residues lining this cavity, which may explain the well-defined density of this portion of NADH in the closed state.	RESULTS
118	125	density	evidence	There are extensive interactions between NADH and the residues lining this cavity, which may explain the well-defined density of this portion of NADH in the closed state.	RESULTS
145	149	NADH	chemical	There are extensive interactions between NADH and the residues lining this cavity, which may explain the well-defined density of this portion of NADH in the closed state.	RESULTS
157	163	closed	protein_state	There are extensive interactions between NADH and the residues lining this cavity, which may explain the well-defined density of this portion of NADH in the closed state.	RESULTS
20	24	open	protein_state	In contrast, in the open conformation, the cavity present in the closed state becomes too narrow for the nicotinamide group; instead, the group is oriented in the opposite direction, parallel to the pivot helix with the amido group extending toward the C-terminal end of the helix.	RESULTS
43	49	cavity	site	In contrast, in the open conformation, the cavity present in the closed state becomes too narrow for the nicotinamide group; instead, the group is oriented in the opposite direction, parallel to the pivot helix with the amido group extending toward the C-terminal end of the helix.	RESULTS
65	71	closed	protein_state	In contrast, in the open conformation, the cavity present in the closed state becomes too narrow for the nicotinamide group; instead, the group is oriented in the opposite direction, parallel to the pivot helix with the amido group extending toward the C-terminal end of the helix.	RESULTS
199	210	pivot helix	structure_element	In contrast, in the open conformation, the cavity present in the closed state becomes too narrow for the nicotinamide group; instead, the group is oriented in the opposite direction, parallel to the pivot helix with the amido group extending toward the C-terminal end of the helix.	RESULTS
275	280	helix	structure_element	In contrast, in the open conformation, the cavity present in the closed state becomes too narrow for the nicotinamide group; instead, the group is oriented in the opposite direction, parallel to the pivot helix with the amido group extending toward the C-terminal end of the helix.	RESULTS
17	21	NADH	chemical	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
38	68	catalytic and regulatory sites	site	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
77	81	NADH	chemical	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
82	89	density	evidence	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
123	138	catalytic sites	site	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
142	148	closed	protein_state	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
157	161	open	protein_state	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
181	185	NADH	chemical	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
186	193	density	evidence	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
224	240	regulatory sites	site	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
244	250	closed	protein_state	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
259	263	open	protein_state	Detailed view of NADH conformation in catalytic and regulatory sites. (A, B) NADH density (purple) and interactions in the catalytic sites of closed (A) and open (B) states. (C, D) NADH density (red) and interactions in the regulatory sites of closed (C) and open (D) states.	FIG
85	91	closed	protein_state	Although there is a difference in orientation of the nicotinamide moiety between the closed and open states in the regulatory site, in both structures the adenine portion of NADH has a similar binding pocket and is located in almost exactly the same position as ADP, a potent activator of GDH function (Supplemental Fig. 5).	RESULTS
96	100	open	protein_state	Although there is a difference in orientation of the nicotinamide moiety between the closed and open states in the regulatory site, in both structures the adenine portion of NADH has a similar binding pocket and is located in almost exactly the same position as ADP, a potent activator of GDH function (Supplemental Fig. 5).	RESULTS
115	130	regulatory site	site	Although there is a difference in orientation of the nicotinamide moiety between the closed and open states in the regulatory site, in both structures the adenine portion of NADH has a similar binding pocket and is located in almost exactly the same position as ADP, a potent activator of GDH function (Supplemental Fig. 5).	RESULTS
140	150	structures	evidence	Although there is a difference in orientation of the nicotinamide moiety between the closed and open states in the regulatory site, in both structures the adenine portion of NADH has a similar binding pocket and is located in almost exactly the same position as ADP, a potent activator of GDH function (Supplemental Fig. 5).	RESULTS
174	178	NADH	chemical	Although there is a difference in orientation of the nicotinamide moiety between the closed and open states in the regulatory site, in both structures the adenine portion of NADH has a similar binding pocket and is located in almost exactly the same position as ADP, a potent activator of GDH function (Supplemental Fig. 5).	RESULTS
193	207	binding pocket	site	Although there is a difference in orientation of the nicotinamide moiety between the closed and open states in the regulatory site, in both structures the adenine portion of NADH has a similar binding pocket and is located in almost exactly the same position as ADP, a potent activator of GDH function (Supplemental Fig. 5).	RESULTS
262	265	ADP	chemical	Although there is a difference in orientation of the nicotinamide moiety between the closed and open states in the regulatory site, in both structures the adenine portion of NADH has a similar binding pocket and is located in almost exactly the same position as ADP, a potent activator of GDH function (Supplemental Fig. 5).	RESULTS
289	292	GDH	protein	Although there is a difference in orientation of the nicotinamide moiety between the closed and open states in the regulatory site, in both structures the adenine portion of NADH has a similar binding pocket and is located in almost exactly the same position as ADP, a potent activator of GDH function (Supplemental Fig. 5).	RESULTS
7	11	open	protein_state	In the open state, the binding of ADP or NADH is further stabilized by His209, a residue that undergoes a large movement during the transition from open to closed conformation (Fig. 3, C and D).	RESULTS
34	37	ADP	chemical	In the open state, the binding of ADP or NADH is further stabilized by His209, a residue that undergoes a large movement during the transition from open to closed conformation (Fig. 3, C and D).	RESULTS
41	45	NADH	chemical	In the open state, the binding of ADP or NADH is further stabilized by His209, a residue that undergoes a large movement during the transition from open to closed conformation (Fig. 3, C and D).	RESULTS
71	77	His209	residue_name_number	In the open state, the binding of ADP or NADH is further stabilized by His209, a residue that undergoes a large movement during the transition from open to closed conformation (Fig. 3, C and D).	RESULTS
148	152	open	protein_state	In the open state, the binding of ADP or NADH is further stabilized by His209, a residue that undergoes a large movement during the transition from open to closed conformation (Fig. 3, C and D).	RESULTS
156	162	closed	protein_state	In the open state, the binding of ADP or NADH is further stabilized by His209, a residue that undergoes a large movement during the transition from open to closed conformation (Fig. 3, C and D).	RESULTS
7	11	open	protein_state	In the open conformation, the distance between His209 and the α-phosphate of NADH is ∼4.4 Å, which is comparable with the corresponding distance in the ADP-bound conformation.	RESULTS
47	53	His209	residue_name_number	In the open conformation, the distance between His209 and the α-phosphate of NADH is ∼4.4 Å, which is comparable with the corresponding distance in the ADP-bound conformation.	RESULTS
77	81	NADH	chemical	In the open conformation, the distance between His209 and the α-phosphate of NADH is ∼4.4 Å, which is comparable with the corresponding distance in the ADP-bound conformation.	RESULTS
152	161	ADP-bound	protein_state	In the open conformation, the distance between His209 and the α-phosphate of NADH is ∼4.4 Å, which is comparable with the corresponding distance in the ADP-bound conformation.	RESULTS
7	13	closed	protein_state	In the closed conformation, however, this key histidine residue is >10.5 Å away from the nearest phosphate group on NADH, altering a critical stabilization point within the regulatory site.	RESULTS
46	55	histidine	residue_name	In the closed conformation, however, this key histidine residue is >10.5 Å away from the nearest phosphate group on NADH, altering a critical stabilization point within the regulatory site.	RESULTS
116	120	NADH	chemical	In the closed conformation, however, this key histidine residue is >10.5 Å away from the nearest phosphate group on NADH, altering a critical stabilization point within the regulatory site.	RESULTS
173	188	regulatory site	site	In the closed conformation, however, this key histidine residue is >10.5 Å away from the nearest phosphate group on NADH, altering a critical stabilization point within the regulatory site.	RESULTS
48	52	NADH	chemical	This suggests that although the conformation of NADH in the open state regulatory site more closely mimics the binding of ADP, the conformation of NADH in the closed state regulatory site is significantly different; these differences may contribute to the opposite effects of NADH and ADP on GDH enzymatic activity.	RESULTS
60	64	open	protein_state	This suggests that although the conformation of NADH in the open state regulatory site more closely mimics the binding of ADP, the conformation of NADH in the closed state regulatory site is significantly different; these differences may contribute to the opposite effects of NADH and ADP on GDH enzymatic activity.	RESULTS
71	86	regulatory site	site	This suggests that although the conformation of NADH in the open state regulatory site more closely mimics the binding of ADP, the conformation of NADH in the closed state regulatory site is significantly different; these differences may contribute to the opposite effects of NADH and ADP on GDH enzymatic activity.	RESULTS
122	125	ADP	chemical	This suggests that although the conformation of NADH in the open state regulatory site more closely mimics the binding of ADP, the conformation of NADH in the closed state regulatory site is significantly different; these differences may contribute to the opposite effects of NADH and ADP on GDH enzymatic activity.	RESULTS
147	151	NADH	chemical	This suggests that although the conformation of NADH in the open state regulatory site more closely mimics the binding of ADP, the conformation of NADH in the closed state regulatory site is significantly different; these differences may contribute to the opposite effects of NADH and ADP on GDH enzymatic activity.	RESULTS
159	165	closed	protein_state	This suggests that although the conformation of NADH in the open state regulatory site more closely mimics the binding of ADP, the conformation of NADH in the closed state regulatory site is significantly different; these differences may contribute to the opposite effects of NADH and ADP on GDH enzymatic activity.	RESULTS
172	187	regulatory site	site	This suggests that although the conformation of NADH in the open state regulatory site more closely mimics the binding of ADP, the conformation of NADH in the closed state regulatory site is significantly different; these differences may contribute to the opposite effects of NADH and ADP on GDH enzymatic activity.	RESULTS
276	280	NADH	chemical	This suggests that although the conformation of NADH in the open state regulatory site more closely mimics the binding of ADP, the conformation of NADH in the closed state regulatory site is significantly different; these differences may contribute to the opposite effects of NADH and ADP on GDH enzymatic activity.	RESULTS
285	288	ADP	chemical	This suggests that although the conformation of NADH in the open state regulatory site more closely mimics the binding of ADP, the conformation of NADH in the closed state regulatory site is significantly different; these differences may contribute to the opposite effects of NADH and ADP on GDH enzymatic activity.	RESULTS
292	295	GDH	protein	This suggests that although the conformation of NADH in the open state regulatory site more closely mimics the binding of ADP, the conformation of NADH in the closed state regulatory site is significantly different; these differences may contribute to the opposite effects of NADH and ADP on GDH enzymatic activity.	RESULTS
7	17	absence of	protein_state	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
18	22	NADH	chemical	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
24	27	GTP	chemical	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
44	47	GDH	protein	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
55	76	dissociation constant	evidence	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
88	95	Cryo-EM	experimental_method	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
108	111	GDH	protein	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
112	126	incubated with	protein_state	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
127	130	GTP	chemical	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
145	154	structure	evidence	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
215	219	open	protein_state	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
265	269	NBDs	structure_element	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
277	281	open	protein_state	In the absence of NADH, GTP binds weakly to GDH with a dissociation constant of20 μM. Cryo-EM analysis of GDH incubated with GTP resulted in a structure at an overall resolution of 3.5 Å, showing that it is in an open conformation (Supplemental Fig. 6), with all NBDs in the open state.	RESULTS
4	11	density	evidence	The density for GTP is not very well defined, suggesting considerable wobble in the binding site.	RESULTS
16	19	GTP	chemical	The density for GTP is not very well defined, suggesting considerable wobble in the binding site.	RESULTS
84	96	binding site	site	The density for GTP is not very well defined, suggesting considerable wobble in the binding site.	RESULTS
0	11	Subtraction	experimental_method	Subtraction of the GTP-bound map with that of the apo state shows that GTP binding can nevertheless be visualized specifically in the GTP binding site (Supplemental Fig. 6).	RESULTS
19	28	GTP-bound	protein_state	Subtraction of the GTP-bound map with that of the apo state shows that GTP binding can nevertheless be visualized specifically in the GTP binding site (Supplemental Fig. 6).	RESULTS
29	32	map	evidence	Subtraction of the GTP-bound map with that of the apo state shows that GTP binding can nevertheless be visualized specifically in the GTP binding site (Supplemental Fig. 6).	RESULTS
50	53	apo	protein_state	Subtraction of the GTP-bound map with that of the apo state shows that GTP binding can nevertheless be visualized specifically in the GTP binding site (Supplemental Fig. 6).	RESULTS
71	74	GTP	chemical	Subtraction of the GTP-bound map with that of the apo state shows that GTP binding can nevertheless be visualized specifically in the GTP binding site (Supplemental Fig. 6).	RESULTS
134	150	GTP binding site	site	Subtraction of the GTP-bound map with that of the apo state shows that GTP binding can nevertheless be visualized specifically in the GTP binding site (Supplemental Fig. 6).	RESULTS
28	31	GTP	chemical	Importantly, the binding of GTP alone does not appear to drive the transition from the open to the closed state of GDH.	RESULTS
87	91	open	protein_state	Importantly, the binding of GTP alone does not appear to drive the transition from the open to the closed state of GDH.	RESULTS
99	105	closed	protein_state	Importantly, the binding of GTP alone does not appear to drive the transition from the open to the closed state of GDH.	RESULTS
115	118	GDH	protein	Importantly, the binding of GTP alone does not appear to drive the transition from the open to the closed state of GDH.	RESULTS
32	36	NADH	chemical	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
41	44	GTP	chemical	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
72	76	open	protein_state	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
80	86	closed	protein_state	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
110	120	determined	experimental_method	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
121	131	structures	evidence	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
135	138	GDH	protein	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
139	154	in complex with	protein_state	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
160	164	NADH	chemical	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
169	172	GTP	chemical	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
178	185	without	protein_state	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
186	195	glutamate	chemical	To further dissect the roles of NADH and GTP in the transition from the open to closed conformations, we next determined structures of GDH in complex with both NADH and GTP, but without glutamate.	RESULTS
5	9	NADH	chemical	When NADH and GTP are both present, classification reveals the presence of both closed and open GDH conformations, similar to the condition when only NADH is present (Fig. 4, A and B).	RESULTS
14	17	GTP	chemical	When NADH and GTP are both present, classification reveals the presence of both closed and open GDH conformations, similar to the condition when only NADH is present (Fig. 4, A and B).	RESULTS
36	50	classification	experimental_method	When NADH and GTP are both present, classification reveals the presence of both closed and open GDH conformations, similar to the condition when only NADH is present (Fig. 4, A and B).	RESULTS
63	74	presence of	protein_state	When NADH and GTP are both present, classification reveals the presence of both closed and open GDH conformations, similar to the condition when only NADH is present (Fig. 4, A and B).	RESULTS
80	86	closed	protein_state	When NADH and GTP are both present, classification reveals the presence of both closed and open GDH conformations, similar to the condition when only NADH is present (Fig. 4, A and B).	RESULTS
91	95	open	protein_state	When NADH and GTP are both present, classification reveals the presence of both closed and open GDH conformations, similar to the condition when only NADH is present (Fig. 4, A and B).	RESULTS
96	99	GDH	protein	When NADH and GTP are both present, classification reveals the presence of both closed and open GDH conformations, similar to the condition when only NADH is present (Fig. 4, A and B).	RESULTS
150	154	NADH	chemical	When NADH and GTP are both present, classification reveals the presence of both closed and open GDH conformations, similar to the condition when only NADH is present (Fig. 4, A and B).	RESULTS
0	37	Reconstruction without classification	experimental_method	Reconstruction without classification, however, yields a structure clearly in the closed conformation, suggesting that, in coordination with NADH, GTP may further stabilize the closed conformation.	RESULTS
57	66	structure	evidence	Reconstruction without classification, however, yields a structure clearly in the closed conformation, suggesting that, in coordination with NADH, GTP may further stabilize the closed conformation.	RESULTS
82	88	closed	protein_state	Reconstruction without classification, however, yields a structure clearly in the closed conformation, suggesting that, in coordination with NADH, GTP may further stabilize the closed conformation.	RESULTS
141	145	NADH	chemical	Reconstruction without classification, however, yields a structure clearly in the closed conformation, suggesting that, in coordination with NADH, GTP may further stabilize the closed conformation.	RESULTS
147	150	GTP	chemical	Reconstruction without classification, however, yields a structure clearly in the closed conformation, suggesting that, in coordination with NADH, GTP may further stabilize the closed conformation.	RESULTS
177	183	closed	protein_state	Reconstruction without classification, however, yields a structure clearly in the closed conformation, suggesting that, in coordination with NADH, GTP may further stabilize the closed conformation.	RESULTS
16	19	GTP	chemical	The location of GTP in the open and closed states of the GDH/NADH/GTP complex is similar to that in the crystal structure observed in the presence of NADH, GTP, and glutamate.	RESULTS
27	31	open	protein_state	The location of GTP in the open and closed states of the GDH/NADH/GTP complex is similar to that in the crystal structure observed in the presence of NADH, GTP, and glutamate.	RESULTS
36	42	closed	protein_state	The location of GTP in the open and closed states of the GDH/NADH/GTP complex is similar to that in the crystal structure observed in the presence of NADH, GTP, and glutamate.	RESULTS
57	69	GDH/NADH/GTP	complex_assembly	The location of GTP in the open and closed states of the GDH/NADH/GTP complex is similar to that in the crystal structure observed in the presence of NADH, GTP, and glutamate.	RESULTS
104	121	crystal structure	evidence	The location of GTP in the open and closed states of the GDH/NADH/GTP complex is similar to that in the crystal structure observed in the presence of NADH, GTP, and glutamate.	RESULTS
138	149	presence of	protein_state	The location of GTP in the open and closed states of the GDH/NADH/GTP complex is similar to that in the crystal structure observed in the presence of NADH, GTP, and glutamate.	RESULTS
150	154	NADH	chemical	The location of GTP in the open and closed states of the GDH/NADH/GTP complex is similar to that in the crystal structure observed in the presence of NADH, GTP, and glutamate.	RESULTS
156	159	GTP	chemical	The location of GTP in the open and closed states of the GDH/NADH/GTP complex is similar to that in the crystal structure observed in the presence of NADH, GTP, and glutamate.	RESULTS
165	174	glutamate	chemical	The location of GTP in the open and closed states of the GDH/NADH/GTP complex is similar to that in the crystal structure observed in the presence of NADH, GTP, and glutamate.	RESULTS
26	30	NADH	chemical	Likewise, the position of NADH in the open and closed states closely resembles the position of NADH in the GDH/NADH open and closed structures.	RESULTS
38	42	open	protein_state	Likewise, the position of NADH in the open and closed states closely resembles the position of NADH in the GDH/NADH open and closed structures.	RESULTS
47	53	closed	protein_state	Likewise, the position of NADH in the open and closed states closely resembles the position of NADH in the GDH/NADH open and closed structures.	RESULTS
95	99	NADH	chemical	Likewise, the position of NADH in the open and closed states closely resembles the position of NADH in the GDH/NADH open and closed structures.	RESULTS
107	115	GDH/NADH	complex_assembly	Likewise, the position of NADH in the open and closed states closely resembles the position of NADH in the GDH/NADH open and closed structures.	RESULTS
116	120	open	protein_state	Likewise, the position of NADH in the open and closed states closely resembles the position of NADH in the GDH/NADH open and closed structures.	RESULTS
125	131	closed	protein_state	Likewise, the position of NADH in the open and closed states closely resembles the position of NADH in the GDH/NADH open and closed structures.	RESULTS
132	142	structures	evidence	Likewise, the position of NADH in the open and closed states closely resembles the position of NADH in the GDH/NADH open and closed structures.	RESULTS
31	35	open	protein_state	One key difference between the open and closed states of these structures is the position of the His209 residue: As mentioned above, His209 swings away from the adenine moiety of NADH in the closed state.	RESULTS
40	46	closed	protein_state	One key difference between the open and closed states of these structures is the position of the His209 residue: As mentioned above, His209 swings away from the adenine moiety of NADH in the closed state.	RESULTS
63	73	structures	evidence	One key difference between the open and closed states of these structures is the position of the His209 residue: As mentioned above, His209 swings away from the adenine moiety of NADH in the closed state.	RESULTS
97	103	His209	residue_name_number	One key difference between the open and closed states of these structures is the position of the His209 residue: As mentioned above, His209 swings away from the adenine moiety of NADH in the closed state.	RESULTS
133	139	His209	residue_name_number	One key difference between the open and closed states of these structures is the position of the His209 residue: As mentioned above, His209 swings away from the adenine moiety of NADH in the closed state.	RESULTS
179	183	NADH	chemical	One key difference between the open and closed states of these structures is the position of the His209 residue: As mentioned above, His209 swings away from the adenine moiety of NADH in the closed state.	RESULTS
191	197	closed	protein_state	One key difference between the open and closed states of these structures is the position of the His209 residue: As mentioned above, His209 swings away from the adenine moiety of NADH in the closed state.	RESULTS
5	8	GTP	chemical	When GTP is present in the GTP binding site, His209 instead interacts with GTP, probably stabilizing the closed conformation (Fig. 4, C and D).	RESULTS
27	43	GTP binding site	site	When GTP is present in the GTP binding site, His209 instead interacts with GTP, probably stabilizing the closed conformation (Fig. 4, C and D).	RESULTS
45	51	His209	residue_name_number	When GTP is present in the GTP binding site, His209 instead interacts with GTP, probably stabilizing the closed conformation (Fig. 4, C and D).	RESULTS
75	78	GTP	chemical	When GTP is present in the GTP binding site, His209 instead interacts with GTP, probably stabilizing the closed conformation (Fig. 4, C and D).	RESULTS
105	111	closed	protein_state	When GTP is present in the GTP binding site, His209 instead interacts with GTP, probably stabilizing the closed conformation (Fig. 4, C and D).	RESULTS
6	9	GTP	chemical	Thus, GTP binding to GDH appears synergistic with NADH and displaces the conformational landscape toward the closed state.	RESULTS
21	24	GDH	protein	Thus, GTP binding to GDH appears synergistic with NADH and displaces the conformational landscape toward the closed state.	RESULTS
50	54	NADH	chemical	Thus, GTP binding to GDH appears synergistic with NADH and displaces the conformational landscape toward the closed state.	RESULTS
109	115	closed	protein_state	Thus, GTP binding to GDH appears synergistic with NADH and displaces the conformational landscape toward the closed state.	RESULTS
0	7	Cryo-EM	experimental_method	Cryo-EM structure of GDH bound to both NADH and GTP.	FIG
8	17	structure	evidence	Cryo-EM structure of GDH bound to both NADH and GTP.	FIG
21	24	GDH	protein	Cryo-EM structure of GDH bound to both NADH and GTP.	FIG
25	33	bound to	protein_state	Cryo-EM structure of GDH bound to both NADH and GTP.	FIG
39	43	NADH	chemical	Cryo-EM structure of GDH bound to both NADH and GTP.	FIG
48	51	GTP	chemical	Cryo-EM structure of GDH bound to both NADH and GTP.	FIG
34	38	open	protein_state	(A, B) Observation of co-existing open (A) and closed (B) conformations in the GDH-NADH-GTP ternary complex.	FIG
47	53	closed	protein_state	(A, B) Observation of co-existing open (A) and closed (B) conformations in the GDH-NADH-GTP ternary complex.	FIG
79	91	GDH-NADH-GTP	complex_assembly	(A, B) Observation of co-existing open (A) and closed (B) conformations in the GDH-NADH-GTP ternary complex.	FIG
0	9	Densities	evidence	Densities for GTP (yellow) as well as NADH bound to both catalytic (purple) and regulatory (red) sites in each protomer are shown.	FIG
14	17	GTP	chemical	Densities for GTP (yellow) as well as NADH bound to both catalytic (purple) and regulatory (red) sites in each protomer are shown.	FIG
38	42	NADH	chemical	Densities for GTP (yellow) as well as NADH bound to both catalytic (purple) and regulatory (red) sites in each protomer are shown.	FIG
43	51	bound to	protein_state	Densities for GTP (yellow) as well as NADH bound to both catalytic (purple) and regulatory (red) sites in each protomer are shown.	FIG
57	66	catalytic	site	Densities for GTP (yellow) as well as NADH bound to both catalytic (purple) and regulatory (red) sites in each protomer are shown.	FIG
80	90	regulatory	site	Densities for GTP (yellow) as well as NADH bound to both catalytic (purple) and regulatory (red) sites in each protomer are shown.	FIG
97	102	sites	site	Densities for GTP (yellow) as well as NADH bound to both catalytic (purple) and regulatory (red) sites in each protomer are shown.	FIG
111	119	protomer	oligomeric_state	Densities for GTP (yellow) as well as NADH bound to both catalytic (purple) and regulatory (red) sites in each protomer are shown.	FIG
56	71	regulatory site	site	(C, D) Detailed inspection of the interactions near the regulatory site show that the orientation of His209 switches between the two states, which may allow interactions with bound GTP in the closed (D), but not open (C) conformation.	FIG
101	107	His209	residue_name_number	(C, D) Detailed inspection of the interactions near the regulatory site show that the orientation of His209 switches between the two states, which may allow interactions with bound GTP in the closed (D), but not open (C) conformation.	FIG
175	180	bound	protein_state	(C, D) Detailed inspection of the interactions near the regulatory site show that the orientation of His209 switches between the two states, which may allow interactions with bound GTP in the closed (D), but not open (C) conformation.	FIG
181	184	GTP	chemical	(C, D) Detailed inspection of the interactions near the regulatory site show that the orientation of His209 switches between the two states, which may allow interactions with bound GTP in the closed (D), but not open (C) conformation.	FIG
192	198	closed	protein_state	(C, D) Detailed inspection of the interactions near the regulatory site show that the orientation of His209 switches between the two states, which may allow interactions with bound GTP in the closed (D), but not open (C) conformation.	FIG
212	216	open	protein_state	(C, D) Detailed inspection of the interactions near the regulatory site show that the orientation of His209 switches between the two states, which may allow interactions with bound GTP in the closed (D), but not open (C) conformation.	FIG
4	22	structural studies	experimental_method	Our structural studies thus establish that whether or not GTP is bound, NADH binding is detectable at catalytic and regulatory sites, in both the open and closed conformational states.	RESULTS
58	61	GTP	chemical	Our structural studies thus establish that whether or not GTP is bound, NADH binding is detectable at catalytic and regulatory sites, in both the open and closed conformational states.	RESULTS
65	70	bound	protein_state	Our structural studies thus establish that whether or not GTP is bound, NADH binding is detectable at catalytic and regulatory sites, in both the open and closed conformational states.	RESULTS
72	76	NADH	chemical	Our structural studies thus establish that whether or not GTP is bound, NADH binding is detectable at catalytic and regulatory sites, in both the open and closed conformational states.	RESULTS
102	132	catalytic and regulatory sites	site	Our structural studies thus establish that whether or not GTP is bound, NADH binding is detectable at catalytic and regulatory sites, in both the open and closed conformational states.	RESULTS
146	150	open	protein_state	Our structural studies thus establish that whether or not GTP is bound, NADH binding is detectable at catalytic and regulatory sites, in both the open and closed conformational states.	RESULTS
155	161	closed	protein_state	Our structural studies thus establish that whether or not GTP is bound, NADH binding is detectable at catalytic and regulatory sites, in both the open and closed conformational states.	RESULTS
33	37	NADH	chemical	Whereas the orientation in which NADH binds at the catalytic site is similar for both conformations, the orientation of the nicotinamide portion of NADH in the regulatory site is different between the open and closed conformations (Figs. 3 and 4).	RESULTS
51	65	catalytic site	site	Whereas the orientation in which NADH binds at the catalytic site is similar for both conformations, the orientation of the nicotinamide portion of NADH in the regulatory site is different between the open and closed conformations (Figs. 3 and 4).	RESULTS
148	152	NADH	chemical	Whereas the orientation in which NADH binds at the catalytic site is similar for both conformations, the orientation of the nicotinamide portion of NADH in the regulatory site is different between the open and closed conformations (Figs. 3 and 4).	RESULTS
160	175	regulatory site	site	Whereas the orientation in which NADH binds at the catalytic site is similar for both conformations, the orientation of the nicotinamide portion of NADH in the regulatory site is different between the open and closed conformations (Figs. 3 and 4).	RESULTS
201	205	open	protein_state	Whereas the orientation in which NADH binds at the catalytic site is similar for both conformations, the orientation of the nicotinamide portion of NADH in the regulatory site is different between the open and closed conformations (Figs. 3 and 4).	RESULTS
210	216	closed	protein_state	Whereas the orientation in which NADH binds at the catalytic site is similar for both conformations, the orientation of the nicotinamide portion of NADH in the regulatory site is different between the open and closed conformations (Figs. 3 and 4).	RESULTS
7	13	closed	protein_state	In the closed state, the nicotinamide moiety is inserted into a well-defined cavity at the interface between two adjacent protomers in the trimer.	RESULTS
77	83	cavity	site	In the closed state, the nicotinamide moiety is inserted into a well-defined cavity at the interface between two adjacent protomers in the trimer.	RESULTS
91	100	interface	site	In the closed state, the nicotinamide moiety is inserted into a well-defined cavity at the interface between two adjacent protomers in the trimer.	RESULTS
122	131	protomers	oligomeric_state	In the closed state, the nicotinamide moiety is inserted into a well-defined cavity at the interface between two adjacent protomers in the trimer.	RESULTS
139	145	trimer	oligomeric_state	In the closed state, the nicotinamide moiety is inserted into a well-defined cavity at the interface between two adjacent protomers in the trimer.	RESULTS
25	31	cavity	site	As mentioned above, this cavity is much narrower in the open state, suggesting that this cavity may be unavailable to the NADH nicotinamide moiety when the enzyme is in the open conformation.	RESULTS
56	60	open	protein_state	As mentioned above, this cavity is much narrower in the open state, suggesting that this cavity may be unavailable to the NADH nicotinamide moiety when the enzyme is in the open conformation.	RESULTS
89	95	cavity	site	As mentioned above, this cavity is much narrower in the open state, suggesting that this cavity may be unavailable to the NADH nicotinamide moiety when the enzyme is in the open conformation.	RESULTS
122	126	NADH	chemical	As mentioned above, this cavity is much narrower in the open state, suggesting that this cavity may be unavailable to the NADH nicotinamide moiety when the enzyme is in the open conformation.	RESULTS
173	177	open	protein_state	As mentioned above, this cavity is much narrower in the open state, suggesting that this cavity may be unavailable to the NADH nicotinamide moiety when the enzyme is in the open conformation.	RESULTS
72	79	density	evidence	These structural features provide a potential explanation of the weaker density for the nicotinamide moiety of NADH in the open state, and may account for the higher reported affinity of NADH for the closed state.	RESULTS
111	115	NADH	chemical	These structural features provide a potential explanation of the weaker density for the nicotinamide moiety of NADH in the open state, and may account for the higher reported affinity of NADH for the closed state.	RESULTS
123	127	open	protein_state	These structural features provide a potential explanation of the weaker density for the nicotinamide moiety of NADH in the open state, and may account for the higher reported affinity of NADH for the closed state.	RESULTS
187	191	NADH	chemical	These structural features provide a potential explanation of the weaker density for the nicotinamide moiety of NADH in the open state, and may account for the higher reported affinity of NADH for the closed state.	RESULTS
200	206	closed	protein_state	These structural features provide a potential explanation of the weaker density for the nicotinamide moiety of NADH in the open state, and may account for the higher reported affinity of NADH for the closed state.	RESULTS
93	97	open	protein_state	The role of the nicotinamide moiety in acting as a wedge that prevents the transition to the open conformation also suggests a structural explanation of the mechanism by which NADH binding inhibits the activity of the enzyme by stabilizing the closed conformation state.	RESULTS
176	180	NADH	chemical	The role of the nicotinamide moiety in acting as a wedge that prevents the transition to the open conformation also suggests a structural explanation of the mechanism by which NADH binding inhibits the activity of the enzyme by stabilizing the closed conformation state.	RESULTS
244	250	closed	protein_state	The role of the nicotinamide moiety in acting as a wedge that prevents the transition to the open conformation also suggests a structural explanation of the mechanism by which NADH binding inhibits the activity of the enzyme by stabilizing the closed conformation state.	RESULTS
23	30	cryo-EM	experimental_method	The rapid emergence of cryo-EM as a tool for near-atomic resolution structure determination provides new opportunities for complementing atomic resolution information from X-ray crystallography, as illustrated here with GDH.	RESULTS
68	91	structure determination	experimental_method	The rapid emergence of cryo-EM as a tool for near-atomic resolution structure determination provides new opportunities for complementing atomic resolution information from X-ray crystallography, as illustrated here with GDH.	RESULTS
172	193	X-ray crystallography	experimental_method	The rapid emergence of cryo-EM as a tool for near-atomic resolution structure determination provides new opportunities for complementing atomic resolution information from X-ray crystallography, as illustrated here with GDH.	RESULTS
220	223	GDH	protein	The rapid emergence of cryo-EM as a tool for near-atomic resolution structure determination provides new opportunities for complementing atomic resolution information from X-ray crystallography, as illustrated here with GDH.	RESULTS
130	139	structure	evidence	Perhaps the most important contribution of these methods is the prospect that when there are discrete subpopulations present, the structure of each state can be determined at near-atomic resolution. What we demonstrate here with GDH is that by employing three-dimensional image classification approaches, we not only can isolate distinct, coexisting conformations, but we can also localize small molecule ligands in each of these conformations.	RESULTS
229	232	GDH	protein	Perhaps the most important contribution of these methods is the prospect that when there are discrete subpopulations present, the structure of each state can be determined at near-atomic resolution. What we demonstrate here with GDH is that by employing three-dimensional image classification approaches, we not only can isolate distinct, coexisting conformations, but we can also localize small molecule ligands in each of these conformations.	RESULTS
254	303	three-dimensional image classification approaches	experimental_method	Perhaps the most important contribution of these methods is the prospect that when there are discrete subpopulations present, the structure of each state can be determined at near-atomic resolution. What we demonstrate here with GDH is that by employing three-dimensional image classification approaches, we not only can isolate distinct, coexisting conformations, but we can also localize small molecule ligands in each of these conformations.	RESULTS