Urea, but not guanidinium, destabilizes proteins by forming hydrogen bonds to the peptide group

^aDepartment of Biochemistry and Biophysics, University of Pennsylvania, Philadelphia, PA 19104;
^bDepartment of Molecular Biology, College of Natural Sciences, Pusan National University, Jangjeon-dong, Keumjeong District, Busan 609-735, South Korea; and
^cDepartment of Biochemistry and Molecular Biology, University of Texas Medical Branch, Galveston, TX 77555

Contributed by S. Walter Englander, December 10, 2008 (received for review November 25, 2008)

Footnotes

¹To whom correspondence may be addressed. E-mail: jorosgen{at}utmb.edu or engl{at}mail.med.upenn.edu
Author contributions: W.K.L., J.R., and S.W.E. designed research; W.K.L. performed research; W.K.L. and J.R. analyzed data; and J.R. and S.W.E. wrote the paper.
The authors declare no conflict of interest.
© 2009 by The National Academy of Sciences of the USA

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	Abstract	Full	Pdf
Feb 2009	4306	106	640
Mar 2009	1670	122	332
Apr 2009	281	63	109
May 2009	166	38	57
Jun 2009	104	30	31
Jul 2009	123	39	47
Aug 2009	86	40	55
Sep 2009	60	66	65
Oct 2009	82	56	52
Nov 2009	60	48	59
Dec 2009	62	48	53
Total 2009	7000	656	1500
Jan 2010	62	52	47
Feb 2010	68	66	48
Mar 2010	61	57	47
Apr 2010	63	65	46
May 2010	32	56	30
Jun 2010	53	39	25
Jul 2010	34	52	37
Aug 2010	44	45	22
Sep 2010	33	93	55
Oct 2010	62	95	47
Nov 2010	44	100	39
Dec 2010	67	59	40
Total 2010	623	779	483
Jan 2011	45	71	68
Feb 2011	85	117	48
Mar 2011	65	97	41
Apr 2011	39	82	43
May 2011	29	63	34
Jun 2011	29	111	32
Jul 2011	32	73	27
Aug 2011	33	67	37
Sep 2011	30	138	40
Oct 2011	39	115	35
Nov 2011	34	112	26
Dec 2011	41	81	35
Total 2011	501	1127	466
Jan 2012	33	151	53
Feb 2012	47	186	71
Mar 2012	70	144	59
Apr 2012	35	167	49
May 2012	71	193	76
Jun 2012	51	155	63
Jul 2012	44	187	65
Aug 2012	38	177	48
Sep 2012	28	226	62
Oct 2012	37	194	36
Nov 2012	37	212	41
Dec 2012	45	171	36
Total 2012	536	2163	659
Jan 2013	34	232	56
Feb 2013	50	243	51
Mar 2013	49	198	41
Apr 2013	49	164	63
May 2013	50	132	53
Jun 2013	32	113	34
Jul 2013	37	119	37
Aug 2013	43	134	35
Sep 2013	47	169	58
Oct 2013	41	203	47
Nov 2013	36	172	54
Dec 2013	44	115	44
Total 2013	512	1994	573
Jan 2014	45	170	51
Feb 2014	50	143	56
Mar 2014	28	121	30
Apr 2014	44	109	38
May 2014	85	130	49
Jun 2014	52	113	36
Jul 2014	52	147	47
Aug 2014	26	115	30
Sep 2014	44	172	39
Oct 2014	38	138	35
Nov 2014	37	117	44
Dec 2014	25	92	31
Total 2014	526	1567	486
Jan 2015	33	193	54
Feb 2015	37	217	53
Mar 2015	50	192	44
Apr 2015	53	208	43
May 2015	31	202	48
Jun 2015	37	117	45
Jul 2015	33	118	38
Aug 2015	37	109	22
Sep 2015	47	237	41
Oct 2015	28	217	61
Nov 2015	33	168	66
Dec 2015	31	156	50
Total 2015	450	2134	565
Jan 2016	51	149	103
Feb 2016	35	169	83
Mar 2016	25	133	79
Apr 2016	40	136	103
May 2016	37	145	67
Jun 2016	31	153	52
Jul 2016	42	124	47
Aug 2016	41	108	63
Sep 2016	23	147	104
Oct 2016	40	205	85
Nov 2016	52	190	119
Dec 2016	43	131	70
Total 2016	460	1790	975
Jan 2017	26	146	112
Feb 2017	69	242	177
Mar 2017	65	227	82
Apr 2017	28	33	41
May 2017	30	221	124
Jun 2017	18	174	52
Jul 2017	19	129	50
Aug 2017	28	187	70
Sep 2017	48	275	99
Oct 2017	28	296	82
Nov 2017	21	176	53
Dec 2017	22	125	47
Total 2017	402	2231	989
Jan 2018	24	169	47
Feb 2018	188	230	66
Mar 2018	172	181	66
Apr 2018	187	199	93
May 2018	217	222	65
Jun 2018	181	172	50
Jul 2018	158	160	77
Aug 2018	173	171	60
Sep 2018	232	233	56
Oct 2018	49	254	82
Nov 2018	6	237	60
Dec 2018	5	141	36
Total 2018	1592	2369	758
Jan 2019	9	131	28
Feb 2019	15	159	50
Mar 2019	21	179	64
Apr 2019	16	248	71
May 2019	21	189	47
Jun 2019	11	128	42
Jul 2019	6	143	42
Aug 2019	11	110	45
Sep 2019	12	182	27
Oct 2019	18	216	34
Nov 2019	3	194	23
Dec 2019	4	284	26
Total 2019	147	2163	499
Jan 2020	11	261	36
Feb 2020	16	333	45
Mar 2020	3	251	43
Apr 2020	14	289	64
May 2020	8	404	43
Jun 2020	9	367	53
Jul 2020	11	329	54
Aug 2020	7	330	39
Sep 2020	9	748	72
Oct 2020	14	541	70
Nov 2020	13	441	77
Dec 2020	12	320	47
Total 2020	127	4614	643
Jan 2021	9	306	49
Feb 2021	7	400	43
Total 2021	16	706	92
Total	12892	24293	8688