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Correlated mutations
The phenomenon of several mutations occurring simultaneously
and dependent on each other
According to the current hypothesis of molecular positive Darwinian
selection, correlated mutations are related to the changes
occurring in their neighborhood, they reflect the protein-to-protein
interaction and they preserve the biological activity
and structural properties of the molecule
Four unrelated protein families have been studied
for correlated mutations occurrence and characteristics:
Eglin-like proteinase inhibitor family (25 sequences)
Bowman-Birk proteinase inhibitor family (52 sequences)
Myoglobins (74 sequences)
Lysozymes (56 sequences)
The multiple alignment of proteins homologous to eglin C from Hirudo medicinalis
(P01051), constructed with the genetic semihomology algorithm (25 sequences)
The conservative consensus residues are shown as white letters on black background. The gray background indicates genetic
semihomology between aligned residues.
P01051|ICIC_HIRME
Q02214
Q40416
Q41361
Q42420
Q43421
Q96465
P01053|ICI2_HORVU
P08626|ICI3_HORVU
P08820|ICIS_VICFA
Q02214|ITR1_NICSY
Q03198|IPIA_TOBAC
P05118|ICI1_LYCES
P16064|ICI1_PHAAN
P19873|ITH5_CUCMA
P24076|BGIA_MOMCH
P20076|IER1_LYCES
Q03199|IPIB_TOBAC
P01052|ICIA_SOLTU
P01054|ICIC_HORVU
P16063|ICIB_HORVU
P16231|ICI1_LYCPE
P16062|ICIA_HORVU
P08454|ICID_SOLTU
Q00783|ICI1_SOLTU
CONSENSUS
8
20
30
40
50
60
70
K-SFPEVVGKTVDQAREYFTLHYPQYD--VYFLPE---GSP-VTLDLRYNRVRVFYNPGTNVVNHVPHVG
KETWPELIGVPAKFAREIIQKENSKLTN-V---PSVLNGSP-VTKDFRCERVRLFVNVLDFVVQI-PRVG
KETWPELIGVPAKLARETIQKENSKLTN-V---PSVLNGSP-VTQDLRCDRVRLFVNLLDIVVQI-PRVG
KNTWPELCGARGEEAAATVETENPSVTA-VIV-PE---GSI-VTTDERCDRVRVWVDENGIVTRV-PVIG
KTSWPEVVGLSVEDAKKVILKDKPDADI-V-VLP-V--GSV-VTADYRPNRVRIFVDI---VAQT-PHIG
KLSWPELVGKDGEEAVKIIQQENPSLD--VILMPR---GQNWATKDYRPNRVRVFNDDSGKVNSI-PRIG
KTEWPELVGCTIKEAKEKIKADRPDLKV-VIV-P-V--GSI-VTQEIDLNRVRVWVDK---VAKV-PKIG
KTEWPELVGKSVEEAKKVILQDKPEAQI--IVLP-V--GTI-VTMEYRIDRVRLFVDKLDNIAQV-PRVG
KTEWPELVEKSVEEAKKVILQDKPEAQI--IVLP-V--GTI-VTMEYRIDRVRLFVDRLDNIAQV-PRVG
RTSWPELVGVSAEEARK-IKEEMPEAEI--QVVPQ---DSF-VTADYKFQRVRLYVDESNKVVRAAP-IG
KETWPELIGVPAKFAREIIQKENSKLTN-V---PSVLNGSP-VTKDFRCERVRLFVNVLDFVVQI-PRVG
KERWPELLGTPAKFAMQIIQKENPKLTN-VQT---ILNGGP-VTEDLRCNRVRLFVNVLDFIVQT-PQIG
KQMWPELIGVPTKLAKEIIEKENPSITN--I--PILLSGSP-ITLDYLCDRVRLFDNILGFVVQM-PVVT
KTSWPELVGVTAEQAETKIKEEMVDVQI--QVSPH---DSF-VTADYNPKRVRLYVDESNKVTRT-PSIG
KSSWPHLVGVGGSVAKAIIERQNPNVKA-VIL--EE--GTP-VTKDFRCNRVRIWVNKRGLVVSP-PRIG
KRSWPQLVGSTGAAAKAVIERENPRVRA-VIV--RV--GSP-VTADFRCDRVRVWVTERGIVARP-PAIG
KESWPELLGTPAKFAKQIIQKENPKLTN-VETL---LNGSA-FTEDLRCNRVRLFVNLLDIVVQT-PKVG
KERWPELLGTPAKFAMQIIQKENPKLTN-VQT---VLNGTP-VTEDLRCNRVRLFVNVLDFVVQT-PQVG
KLQWPELIGVPTKLAKEIIEKQNSLISN-VHI---LLNGSP-VTMDFRCNRVRLFDDILGSVVQI-PRVA
KTSWPEVVGMSAEKAKEIILRDKPNAQIEVI--P-VDAMVP---LNFNPNRVFVLVHKATTVAZVS-RVG
KRSWPEVVGMSAEKAKEIILRDKPDAQIEVI--P-VDAMVP---LDFNPNRIFILVA----VART-PTVG
KQFWPELIGVPALYAKGIIEKENPSITN--I--PILLNGSP-VTKDFRCDRVRLFVNILGDVVQI-PRVT
KTSWPEVVGMSAEKAKEIILRDKPNAQVEVI--P-VDAMVH---LNFDPNRVFVLVA----VART-PTVG
KQRWPELIGVPTKLAKGIIEKENSLITN-VQ---ILLNGSP-VTMDYRCNRVRLFDNILGDVVQI-PRVA
KLRWPELIGVPTKLAKGIIEKENSLISN-VH---ILLNGSP-VTLDIRCDRVRLFDNILGYVVDI-PVVG
KXSWPELVGVPAKXAKXIIXKENPXXXX-VIX-PXVXXGSP-VTXDXRCNRVRLFVNXLXXVVQX-PXVG
The multiple alignment of 52 Bowman-Birk proteinase inhibitors
constructed with the genetic semihomology algorithm
The conservative consensus residues are shown as white letters on black background. The gray background indicates
genetic semihomology between aligned residues.
P01055
P01057
P01056
P01058
P01059
P01063
P17734
P81483
P81484
P16343
P01064
P82469
P01061
P01062
P01060
1BBI:
1D6R:I
1DF9:C
1PI2:
1PBI:A
AAB4719
TISYC2
JC2225
TIZB2
JC2073
JC2072
0506164
0401177
763679A
TISYD2
0907248
1102213
1102213
0404180
TIZB1B
TIMB
TIZB1P
JC1066
Q41066
P80321
Q41065
P81705
P56679
P16346
P01065
P24661
P07679
P19860
P22737
220645
P09864
P09863
CONSENSUS
3
10
20
30
40
50
60
ESSKPCCDQCACTKSNPPQCRCSDMRLNSCHSACKSCICALSYPAQCF-CVDITDFCYEP-CKP
ESSKPCCDECACTKSIPPQCRCTDVRLNSCHSACSSCVCTFSIPAQCV-CVDMKDFCYAP-CKS
QSSKPCCBHCACTKSIPPQCRCTDLRLDSCHSACKSCICTLSIPAQCV-CBBIBDFCYEP-CKS
ESSKPCCDQCSCTKSMPPKCRCSDIRLNSCHSACKSCACTYSIPAKCF-CTDINDFCYEP-CKS
ESSKPCCDLCTCTKSIPPQCHCNDMRLNSCHSACKSCICALSEPAQCF-CVDTTDFCYKS-CHN
ESSKPCCDLCMCTASMPPQCHCADIRLNSCHSACDRCACTRSMPGQCR-CLDTTDFCYKP-CKS
QSSKPCCRQCACTKSIPPQCRCSQVRLNSCHSACKSCACTFSIPAQCF-CGBIBBFCYKP-CKS
-SSKPCCBHCACTKSIPPQCRCSBLRLNSCHSECKGCICTFSIPAQCI-CTDTNNFCYEP-CKS
-SSKPCCBHCACTKSIPPQCRCSBLRLNSCHSECKGCICTFSIPAQCI-CTDTNNFCYEP-CKS
ESSKPCCSSC-CTRSRPPQCQCTDVRLNSCHSACKSCMCTFSDPGMCS-CLDVTDFCYKP-CKS
EYSKPCCDLCMCTRSMPPQCSCEDIRLNSCHSDCKSCMCTRSQPGQCR-CLDTNDFCYKP-CKS
-SSGPCCDRCRCTKSEPPQCQCQDVRLNSCHSACEACVCSHSMPGLCS-CLDITHFCHEP-CKS
ESSHPCCDLCLCTKSIPPQCQCADIRLDSCHSACKSCMCTRSMPGQCR-CLDTHDFCHKP-CKS
ESSEPCCDSCDCTKSIPPECHCANIRLNSCHSACKSCICTRSMPGKCR-CLDTDDFCYKP-CES
QSSPPCCBICVCTASIPPQCVCTBIRLBSCHSACKSCMCTRSMPGKCR-CLBTTBYCYKS-CKS
ESSKPCCDQCACTKSNPPQCRCSDMRLNSCHSACKSCICALSYPAQCF-CVDITDFCYEP-CKP
---KPCCDQCACTKSNPPQCRCSDMRLNSCHSACKSCICALSYPAQCF-CVDITDFCYEP-CKESSEPCCDSCDCTKSIPPQCHCANIRLNSCHSACKSCICTRSMPGKCR-CLDTDDFCYKP-CES
EYSKPCCDLCMCTRSMPPQCSCED-RINSCHSDCKSCMCTRSQPGQCR-CLDTNDFCYKP-CKS
DVKSACCDTCLCTKSNPPTCRCVDVGET-CHSACLSCICAYSNPPKCQ-CFDTQKFCYKQ-CHN
ESSKPCCDQCTCTKSIPPQCRCTDVRLNSCHSACSSCVCTFSIPAQCV-CVDMKDFCYAP-CKS
ESSKPCCDLCMCTASMPPQCHCADIRLNSCHSACDRCACTRSMPGQCR-CLDTTDFCYKP-CKS
ESSKPCCDLCMCTASMPPQCHCADIRLNSCHSACDRCACTRSMPGQCR-CLDTTDFCYKP-CKS
ESSKPCCDQC-CTKSMPPKCRCSDIRLDSCHSACKSCACTYSIPAKCF-CTDINDFCYEP-CKS
ESSKPCCDECKCTKSEPPQCQCVDTRLESCHSACKLCLCALSFPAKCR-CVDTTDFCYKP-CKS
ESSKPCCDECKCTKSEPPQCQCVDTRLESCHSACKLCLCALSFPAKCR-CVDTTDFCYKP-CKS
ESSKPCCDQC-CTKSMPPKCRCSDIRLDSCHSACKSCACTYSIPAKCF-CTDINDFCYEP-CKS
ESSKPCCDLCMCTASMPPQCHCADIRLNSCHSACDRCACTRSMPGQCR-CLDTTDFCYKP-CKS
ESSKPCCDLCMCTASMPPQCHCADIRLNSCHSACDRCACTRSMPGQCR-CLDTTDFCYKP-CKS
EYSKPCCDLCMCTRSMPPQCSCEDIRLNSCHSDCKSCMCTRSQPGQCR-CLDTNDFCYKP-CKS
ESSEPCCDSCRCTKSIPPQCHCADIRLNSCHSACKSCMCTRSMPGKCR-CLDTDDFCYKP-CES
ESSEPCCDLCLCTKSIPPQCQCADIRLNSCHSACKSCMCTRSMPGQCH-CLDTHDFCHKP-CKS
ESSEPCCDLCLCTKSIPPQCQCADIRLNSCHSACKSCMCTRSMPGQCR-CLDTHDFCHKP-CKS
EYSKPCCDLCMCTRSMPPQCSCEDIRLNSCHSDCKSCMCTRSQPGQCR-CLDTNDFCYKP-CKS
ESSHPCCDLCLCTKSIPPQCQCADIRLDSCHSACKSCMCTRSMPGQCH-CLDTHDFCHKP-CKS
ESSEPCCDSCDCTKSKPPQCHCANIRLNSCHSACKSCICTRSMPGKCR-CLDTDDFCYKP-CES
ESSHPCCDLCLCTKSIPPQCQCADIRLNSCHSACKSCMCTRSMPGQCR-CLDTHDFCHKP-CKS
ESSEPCCDSCDCTKSKPPQCHCANIRLNSCHSACKSCICTRSMPGKCR-CLDTDDFCTKP-CES
DVKSACCDTCLCTKSDPPTCRCVDVGET-CHSACDSCICALSYPPQCQ-CFDTHKFCYKA-CHN
STTTACCDFCPCTRSIPPQCQCTDVREK-CHSACKSCLCTLSIPPQCH-CYDITDFCYPS-CRDVKSACCDTCLCTKSNPPTCRCVDVRET-CHSACDSCICAYSNPPKCQ-CFDTHKFCYKA-CHN
--TSACCDKCFCTKSNPPICQCRDVGET-CHSACKFCICALSYPAQCH-CLDQNTFCYDK-CDS
DVKSACCDTCLCTKSNPPTCRCVDVGET-CHSACLSCICAYSNPPKCQ-CFDTQKFCYKA-CHN
--TTACCNFCPCTRSIPPQCRCTDIGET-CHSACKTCLCTKSIPPQCH-CADITNFCYPK-CNDVKSACCDTCLCTRSQPPTCRCVDVGER-CHSACNHCVCNYSNPPQCQ-CFDTHKFCYKA-CHS
DVKSACCDTCLCTKSEPPTCRCVDVGER-CHSACNSCVCRYSNPPKCQ-CFDTHKFCYKS-CHN
KRPWECCDIAMCTRSIPPICRCVDKVDR-CSDACKDCEETEDN--RHV-CFDTYIGDPGPTCHD
ERPWKCCDLQTCTKSIPAFCRCRDLLEQ-CSDACKECGKVRDSDPPRYICQDVYRGIPAPMCHE
ERPWKCCDLQTCTKSIPAFCRCRDLLEQ-CSDACKECGKVRDSDPPRYICQDVYRGIPAPMCHE
ES-EGCCDRCICTKSMPPQCHCHDVRLDSCHSDCETCICTRSYPAQCR-CADTTDFCYKP-C-S
TRPWKCCDRAICTKSFPPMCRCMDMVEQ-CAATCKKCGPATSDSSRRV-CEDXY----------KRPWKCCDQAVCTRSIPPICRCMDQVFE-CPSTCKACGPSVGDPSRRV-CQDQYV---------ESSKPCCDXCXCTKSIPPQCRCXDXRLNSCHSACKSCXCTRSXPXQCX-CXDTXDFCYKP-CKS
The multiple alignment of 74 myoglobins, constructed with the genetic semihomology algorithm
p02144
p02145
p02146
p02147
p02148
p02149
p02150
p02163
p02189
p14396
p02152
p02151
p02153
p02165
p11343
p02164
p02154
p02193
p20856
p02171
p02155
p02181
p02166
p02156
p04249
p04250
p04248
p02161
p02157
p02188
p02176
p02170
p02174
p32428
p02172
p02173
p02177
p02183
p02167
p02178
p02169
p02160
p02185
p02158
p02194
p02159
p02182
p02186
p02179
p02162
p02184
p02187
p02191
p02190
p02180
p02168
p02192
p04247
p02196
p02195
p30562
p02197
p02199
p02203
p02201
p56208
p02202
p02200
p02206
p02205
p14399
p02204
p14397
p14398
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: G L S D G E W Q L V L N V W G K V E A D I P G H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GA T V L T A L G G I L K K K G H H E A E I K P L A Q S H A T K H K I P V K Y L E F I S E C I I Q V L Q S K H P G D F G A D A Q G A M N K A L E L F R K D M A S N Y K E L G F Q G
: G L S D G E W Q L V L N V W G K V E A D I P G H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GA T V L T A L G G I L K K K G H H E A E I K P L A Q S H A T K H K I P V K Y L E F I S E C I I Q V L H S K H P G D F G A D A Q G A M N K A L E L F R K D M A S N Y K E L G F Q G
: G L S D G E W Q L V L N V W G K V E A D I P S H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GA T V L T A L G G I L K K K G H H E A E I K P L A Q S H A T K H K I P V K Y L E F I S E C I I Q V L Q S K H P G D F G A D A Q G A M N K A L E L F R K D M A S N Y K E L G F Q G
: G L S D G E W Q L V L N V W G K V E A D I S G H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GA T V L T A L G G I L K K K G H H E A E I K P L A Q S H A T K H K I P V K Y L E F I S E C I I Q V L Q S K H P G D F G A D A Q G A M N K A L E L F R K D M A S N Y K E L G F Q G
: G L S D G E W Q L V L N V W G K V E A D I P S H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GA T V L T A L G G I L K K K G H H E A E I K P L A Q S H A T K H K I P V K Y L E F I S E S I I Q V L Q S K H P G D F G A D A Q G A M N K A L E L F R K D M A S N Y K E L G F Q G
: G L S D G E W Q L V L N V W G K V E A D I P S H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GA T V L T A L G G I L K K K G H H E A E I K P L A Q S H A T K H K I P V K Y L E L I S E S I I Q V L Q S K H P G D F G A D A Q G A M N K A L E L F R N D M A A K Y K E L G F Q G
: G L S D G E W Q L V L N V W G K V E A DI P S H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GV T V L T A L G G I L K K K G H H E A EI K P L A Q S H A T K H K I P V K Y L EL I S E S I I Q V L Q S K H P G D F G A D A Q G A MN K A L E L F R N D M A A K Y K E L G F Q G
: G L S D G E W Q L V L N V W G K V E A D I P G H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GA T V L T A L G G I L K K K G Q H E A Q L K P L A Q S H A T K H K I P V K Y L E F I S E V I I Q V L Q S K H P G D F G A D A Q G A M G K A L E L F R N D I A A K Y K E L G F Q G
: G L S D G E W Q L V L N V W G K V E A D V A G H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H G N T V L T A L G G I L K K K G H H E A E LT P L A Q S H A T K H K I P V K Y L E F I S E A I I Q V L Q S K H P G D F G A D A Q G A M S K A L E L F R N D M A A K Y K E L G F Q G
: GLSDGEWQLVLHVWGKVEADLAGHGQEVLIRLFKGHPETLEKFNKFKHIKSEDEMKASEDLKKHGVTVLTALGGVLKKKGHHEAEIKPLAQSHATKHKIPIKYLEFISEAIIHVLQSKHPGBFGADABGAMNKALELFRKDIAAKYKELGFQG
: G L S D G E W Q L V L N V W G K V E A DI P S H G Q E V L I S L F K G H P E T L E K F D K F K H L K S E D E M K A S EE L K K H G V T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H K I P V K Y L E F I SD A I V H V L Q K K H P G D F G A D A Q G A MK K A L E L F R N D M A A K Y K E L G F Q G
: G L S D G E W Q L V L N V W G K V E A DV P S H G Q E V L I S L F K G H P E T L E K F D K F K H L K S E D E M K A S EE L K K H G V T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H K I P V K Y L E F I SD A I V H V L Q K K H P G D F G A D A Q G A MK K A L E L F R N D M A A K Y K E L G F Q G
: GLSDGEWQLVLNVWGKVEADIPSHGQEVLISLFKGHPETLEKFDKFKHLKSEDEMKASEELKKHGATVLTALGGILKKKGQHEAELKPLAQSHATKHKIPVKYLEFISDAIVHVLQKKHPGDFGADAQGAMKKALELFRNDMAAKYKELGFQG
: GLSDGEWQLVLNVWGKVEADVAGHGQEVLIRLFKGHPETLEKFDKFKHLKTEDEMKASEDLKKHGNTVLSALGGILKKKGQHEAEIKPLAQSHATKHKIPVKYLEFISEAIIQVLQSKHPGDFGADAQAAMSKALELFRNDIAAKYKELGFQG
: G L S D G E W Q L V L N V W G K V E A D L A G H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M KG S E D L K K H G N T V L T A L G G I L K K K G K H E A E L K P L A Q S H A T K H K I P I K Y L E F I S E A I I Q V L Q S K H P G B F G A D A Q G A M K R A L E L F R N D I A A K Y K E L G F Q G
: GLSDAEWQLVLNVWGKVEADIPGHGQDVLIRLFKGHPETLEKFDRFKHLKTEDEMKASEDLKKHGTTVLTALGGILKKKGQHEAEIQPLAQSHATKHKIPVKYLEFISEAIIQVIQSKHSGDFGADAQGAMSKALELFRNDIAAKYKELGFQG
: GLSDGEWQLVLNIWGKVEADIPSHGQEVLISLFKGHPETLEKFDKFKHLKSEDEMKASEELKKHGVTVLTALGGILKKKGQHEAELKPLAQSHATKHKIPVKYLEFISDAIIHALQKKHPGDFGADAQGAMKKALELFRNDMAAKYKELGFQG
: G L S D G E W Q L V L N A W G K V E A D I P G H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GA T V L T A L G N I L K K K G N H E A E L K P L A Q S H A T K H K I S V Q F L E F I S E A I I Q V I Q S K H P G D F G G D A Q A A M G K A L E L F R N D M A A K Y K E L G F Q G
: G L S D G E W Q L V L N A W G K V E T D I G G H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GT T V L T A L G N I L K K K G Q H E A E L A P L A Q S H A T K H K I P V K Y L E F I S E A I I Q V L E S K H P G D F G A D A Q G A M S K A L E L F R N D I A A K Y K E L G F Q G
: GLSDGEWQLVLNVWGKVEADLAGHGQEVLIRLFKNHPETLEKFDKFKNLKSEDEMKGSDDLKKHGNTVLSALGGILKKKGQHEAELKPLAQSHATKHKIPVKYLEFISEAIIQVLQSKHPGDFGADAQGAMSKALELFRNDMAAKYKELGFQG
: GLSDGEWQLVLNIWGKVEADIPSHGQEVLISLFKGHPETLEKFDKFKHLKSEDEMKASEELKKHGTTVLTALGGILKKKGQHEAELKPLAQSHATKHKIPVKYLELISDAIVHVLQKKHPGDFGADAQGAMKKALELFRNDMAAKYKELGFQG
: G L S D G E W Q L V L N I W G K V E A D L A G H G Q D V L I R L F K G H P E T L E K F D K F K H L K T E A E M K A S E D L K K H G N T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H K I PI K Y L E F I S E A I I H V L H S R H P G D F G A D A Q A A M N K A L E L F R K D I A A K Y K E L G F H G
: G L S D G E W Q S V L N V W G K V E A D L A G H G Q E I L I R L F T A H P E T L E K F D K F K N L K T P D E M K A S E D L K K H G V T V L T A L G G I L K K K G H H E A E I K P L A Q S H A T K H K I P V K Y L E F I S E A I I H V L Q S K H P G D F G A D A Q G A MN K A L E L F R N D I A A K Y K E L G F Q G
: GLSDGEWQLVLNVWGKVEADIPGHGQEVLIRLFKDHPETLEKFDKFKHLKSEDEMKSSEDLKKHGTTVLTALGGILKKKGQHEAQLAPLAQSHANKHKIPVKYLEFISEAIIQVLKSKHAGDFGADAQGAMSKALELFRNDIAAKYKELGFQG
: GLSDGEWQLVLNVWGKVEGDLSGHGQEVLIRLFKGHPETLEKFDKFKHLKAEDEMRASEELKKHGTTVLTALGGILKKKGQHAAELAPLAQSHATKHKIPVKYLEFISEAIIQVLQSKHPGDFGADAQGAMSKALELFRNDIAAKYKELGFQG
: GLSDGEWQLVLNVWGKVEADLGGHGQEVLIRLFKGHPETLEKFDKFKHLKAEDEMRASEDLKKHGTTVLTALGGILKKRGQHAAELAPLAQSHATKHKIPVKYLEFISEAIIQVLQSKHPGDFGADAQAAMSKALELFRNDIAAKYKELGFQG
: GLSDGEWQLVLNVWGKVEGDLAGHGQEVLIKLFKNHPETLEKFDKFKHLKSEDEMKGSEDLKKHGNTVLTALGGILKKKGQHAAEIQPLAQSHATKHKIPIKYLEFISEAIIQVLQSKHPGDFGADAQGAMSKALELFRNDIAAKYKELGFQG
: G L S D G E W Q L V L NI W G K V E A D LV G H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M KRS E D L K K H GK T V L T A L G G I L K K K G H HD A E L K P L A Q S H A T K H K I PIK Y L E F I S E A I I H V L Q S K H P G D F G A DT H AA M KK A L E L F R N D I A AK YR E L G F Q G
: G L S D G E W Q L V L N V W G K V E A D L A G H G Q E V L I R L F K G H P E T L E K F D K F K H L K S E D E M KG S E D L K K H G N T V L T A L G G I L K K K G H Q E A E L K P L A Q S H A T K H K I P V K Y L E F I S D A I A Q V L Q S K H P G N F A A E A Q G A M K K A L E L F R N D I A A K Y K E L G F Q G
: GLSDGEWQQVLNVWGKVEADIAGHGQEVLIRLFTGHPETLEKFDKFKHLKTEAEMKASEDLKKHGTVVLTALGGILKKKGHHEAELKPLAQSHATKHKIPIKYLEFISDAIIHVLHSKHPGDFGADAQGAMTKALELFRNDIAAKYKELGFQG
: GLSEGEWQLVLNVWGKVEADLAGHGQDVLIRLFKGHPETLEKFDKFKHLKTEAEMKASEDLKKHGNTVLTALGGILKKKGHHDAELKPLAQSHATKHKIPIKYLEFISEAIIHVLHSRHPAEFGADAQGAMNKALELFRKDIATKYKELGFHG
: G L S D A E W Q L V L N V W G K V E A D L A G H G Q E V L I R L FH T H P E T L E K F D K F K H L K S E D E M K A S E D L K K H G N T V L T A L GA I L K K K G H H E A EI K P L A Q S H A T K H K I P V K Y L E F I S E A I I H V LH S K H P G D F G A D A QA A M S K A L E L F R N D I A AQ Y K E L G F Q G
: GLSDGEWQLVLNVWGKVEADLAGHGQDILIRLFKGHPETLEKFDKFKHLKTEADMKASEDLKKHGNTVLTALGAILKKKGHHEAELKPLAQSHATKHKIPIKYLEFISEAIIHVLHSRHPAEFGADAQGAMNKALELFRKDIAAKYKELGFHG
: GLSDGEWQLVLHVWGKVEADLAGHGQDVLIRLFKAHPETLEKFDKFKHIKSEDEMKGSEDLKKHGBTVLTALGGILKKKGHHEAEIKPLAQSHATKHKIPIKYLEFISEAIIHVLZSKHPSBFGADVZGAMKRALELFRNDIAAKYKELGFQG
: GLSDGEWQLVLNVWGKVEADLAGHGQDVLIRLFKGHPETLEKFDKFKHLKTEADMKASEDLKKHGNTVLTALGAILKKKGHHDAELKPLAQSHATKHKIPIKYLEFISEAIIHVLHSRHPAEFGADAQGAMNKALELFRKDIAAKYKELGFHG
: GLSDGEWQLVLNVWGKVEADLAGHGQDILIRLFKGHPETLEKFDKFKHLKTEADMKASEDLKKHGNTVLTALGAILKKKGHHDAELKPLAQSHATKHKIPIKYLEFISEAIIHVLHSRHPAEFGADAQGAMNKALELFRKDIAAKYKELGFHG
: V L S D A E W Q L V L N I W A K V E A D V A G H G Q D I L I R L F K G H P E T L E K F D K F K H L K T E A E M K A S E D L K K H G N T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H K I PI K Y L E F I S D A I I H V L H S R H P G D F G A D A Q A A M N K A L E L F R K D I A A K Y K E L G F Q G
: GLSEAEWQLVLHVWAKVEADLSGHGQEILIRLFKGHPETLEKFDKFKHLKSEAEMKASEDLKKHGHTVLTALGGILKKKGHHEAELKPLAQSHATKHKIPIKYLEFISDAIIHVLHSKHPSDFGADAQGAMTKALELFRKDIAAKYKELGFHG
: GLSDGEWQSVLNVWGKVEADLAGHGQEILIRLFTAHPETLEKFDKFKNLKTPDEMKASEDLKKHGVTVLTALGGILKKKGQHEAEIKPLAQSHATKHKIPVKYLEFISGAIIHVLQSKHPGDFGADAQGAMSKALELFRNDIAAKYKELGFQG
: V L S D A E W Q L V L N I W A K V E A D V A G H G Q D I L I R L F K G H P E T L E K F D K F K H L K T E A E M K A S E D L K K H G N T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H K I PI K Y L E F I S D A I I H V L H S R H P A D F G A D A Q A A M N K A L E L F R K D I A A K Y K E L G F Q G
: GLSDGEWQLVLNVWGKVEADVGGHGQEVLIRLFTGHPETLEKFDKFKHLKTADEMKASEDLKKHGTTVLTALGGILKKKGQHEAELKPLAQSHATKHKIPIKYLEFISDAIVHVLHSKHPAEFGADAQAAMKKALELFRNDIAAKYKELGFQG
: G L S D G E W Q L V L N I W G K V E T D L A G H G Q E V L I R L F K N H P E T L D K F D K F K H L K T E D E M K G S E D L K K H G N T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H K I P V K Y L E F I SD A I I Q V L Q S K H S G D F H A D T E A A M K K A L E L F R N D I A A K Y K E L G F Q G
: VLSEGEWQLVLHVWAKVEADVAGHGQDILIRLFKSHPETLEKFDRFKHLKTEAEMKASEDLKKHGVTVLTALGAILKKKGHHEAELKPLAQSHATKHKIPIKYLEFISEAIIHVLHSRHPGDFGADAQGAMNKALELFRKDIAAKYKELGYQG
: G L S D G E W Q I V L N I W G K V E T D L A G H G Q E V L I R L F K N H P E T L D K F D K F K H L K T E D E M K G S E D L K K H G N T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H K I P V K Y L E F I SD A I I Q V L Q S K H S G D F H A D T E A A M K K A L E L F R N D I A A K Y K E L G F Q G
: G L S D G E W Q L V L NI W G K V E T D E G G H G K D V L I R L F K G H P E T L E K F D K F K H L K S E D E M K A S E D L K K H GI T V L T A L G N I L K K K G H H E A E L K P L A Q S H A T K H K I P VQ F L E F I S D A I I Q V I Q S K H A G N F G A D A Q A A M K K A L E L F R H D M A A K Y K E F G F Q G
: G L S D G E W Q I V L N I W G K V E T D L A G H G Q E V L I R L F K N H P E T L D K F D K F K H L K T E D E M K G S E D L K K H G N T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H K I P V K Y L E F I SD A I I Q V L Q N K H S G D F H A D T E A A M K K A L E L F R N D I A A K Y K E L G F Q G
: GLSEAEWQLVLHVWAKVEADLSGHGQEILIRLFKGHPETLEKFDKFKHLKSEAEMKASEDLKKHGHTVLTALGGILKKKGHHEAELKPLAQSHATKHKIPIKYLEFISDAIIHVLHSRHPSDFGADAQAAMTKALELFRKDIAAKYKELGFHG
: GLSDGEWELVLKTWGKVEADIPGHGETVFVRLFTGHPETLEKFDKFKHLKTEGEMKASEDLKKQGVTVLTALGGILKKKGHHEAEIQPLAQSHATKHKIPIKYLEFISDAIIHVLQSKHPAEFGADAQGAMKKALELFRNDIAAKYKELGFQG
: V L S D A E W H L V L N I W A K V E A D V A G H G Q D I L I R L F K G H P E T L E K F D K F K H L K T E A E M K A S E D L K K H G N T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H K I PI K Y L E F I S D A I I H V L H S R H P A E F G A D A Q A A M N K A L E L F R K D I A A K Y K E L G F Q G
: G L S D G E W HL V L N V W G K V ET D L A G H G Q E V L I R L F KS H P E T L E K F D K F K H L K S E DD M R R S E D LR K H G N T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H KI P IK Y L E F I S E A I I H V LH S K H PA E F G A D A Q A A M K K A L E L F R N D I A A K Y K E L G FH G
: VLSEGEWQLVLHVWAKVEADIAGHGQDILIRLFKHHPETLEKFDRFKHLKSEAEMKASEDLKKHGVTVLTALGAILKKKGHHEAELKPLAQSHATKHKIPIKYLEFISEAIIHVLHSRHPADFGADAQGAMSKALELFRKDIAAKYKELGYQG
: GLSDGEWELVLKTWGKVEADIPGHGEFVLVRLFTGHPETLEKFDKFKHLKTEGEMKASEDLKKQGVTVLTALGGILKKKGHHEAEIQPLAQSHATKHKIPIKYLEFISDAIIHVLQSKHPAEFGADAQAAMKKALELFRNDIAAKYKELGFQG
: GLSDGEWQLVLNAWGKVEADVAGHGQEVLIRLFTGHPETLEKFDKFKHLKTEAEMKASEDLKKHGNTVLTALGGILKKKGHHEAEVKHLAESHANKHKIPVKYLEFISDAIIHVLHAKHPSNFGADAQGAMSKALELFRNDMAAQYKVLGFQG
: GLSDGEWQLVLNAWGKVEADVAGHGQEVLIRLFTGHPETLEKFDKFKHLKTEAEMKASEDLKKHGNTVLTALGGILKKKGHHEAEVKHLAESHANKHKIPVKYLEFISDAIIHVLHAKHPSNFGADAQGAMSKALELFRNDMAAEYKVLGFQG
: V L T D A E W H L V L N I W A K V E A D V A G H G Q D I L I S L F K G H P E T L E K F D K F K H L K T E A E M K A S E D L K K H G N T V L T A L G G I L K K K G H H E A E L K P L A Q S H A T K H K I PI K Y L E F I S D A I I H V L H S R H P A D F G A D A Q A A M N K A L E L F R K D I A A K Y K E L G F Q G
: GLSDGEWQLVLKIWGKVEADLAGHGQDVLIRLFTAHPETLEKFDKFKNLKTADEMKASEDLKKHGVTVLTALGGILKKKGQHEAEIKPLAQSHATKHKIPVKYLEFISEAIIHVLQNKHSGDFGTDVQGAMSKALELFRNDIAAKYKELGFQG
: GLSDGEWQLVLNAWGKVEADVAGHGQEVLIRLFTGHPETLEKFDKFKHLKTEAEMKASEDLKKHGNTVLTALGGILKKKGHHEAEVKHLAESHANKHKIPVKYLEFISDAIIHVLHAKHPSDFGADAQAAMSKALELFRNDMAAQYKVLGFHG
: GLSDGEWQLVLNVWGKVEADLAGHGQEVLIGLFKTHPETLDKFDKFKNLKSEEDMKGSEDLKKHGCTVLTALGTILKKKGQHAAEIQPLAQSHATKHKIPVKYLEFISEIIIEVLKKRHSGDFGADAQGAMSKALELFRNDIAAKYKELGFQG
: GLSDGEWQLVLKVWGKVEGDLPGHGQEVLIRLFKTHPETLEKFDKFKGLKTEDEMKASADLKKHGGTVLTALGNILKKKGQHEAELKPLAQSHATKHKISIKFLEYISEAIIHVLQSKHSADFGADAQAAMGKALELFRNDMAAKYKEFGFQG
: GLSDGEWQLVLKVWGKVETDITGHGQDVLIRLFKTHPETLEKFDKFKHLKTEDEMKASADLKKHGGVVLTALGSILKKKGQHEAELKPLAQSHATKHKISIKFLEFISEAIIHVLQSKHSADFGADAQAAMGKALELFRNDMATKYKEFGFQG
: GLSDGEWHLVLNVWGKWETDLAGHGQEVLIRLFKSHPETLEKFDKFKHLKSEDDMRRSFDLRKHGNTVLTALGGILKKKGHHEAELKPLAQSHATKHKIPIKYLEFISEAIIHVLHSKHPAEFGADAQAAMKKALELFRNDIAAKIKELGFHG
: GLSDQEWQQVLTIWGKVEADIAGHGHEVLMRLFHDHPETLDRFDKFKGLKTPDQMKGSEDLKKHGATVLTQLGKILKQKGNHESELKPLAQTHATKHKIPVKYLEFISEVIIKVIAEKHAADFGADSQAAMKKALELFRNDMASKYKEFGFQG
: GLNDQEWQQVLTMWGKVESDLAGHGHAVLMRLFKSHPETMDRFDKFRGLKTPDEMRGSEDMKKHGVTVLT-LGQILKKKGHHEAELKPLSQTHATKHKVPVKYLEFISEAIMKVIAQKHASNFGADAQEAMKKALELFRNDMASKYKEFGFQG
: GLSDEEWKKVVDIWGKVEPDLPSHGQEVIIRMFQNHPETQDRFAKFKNLKTLDEMKNSEDLKKHGTTVLTALGRILKQKGHHEAEIAPLAQTHANTHKIPIKYLEFICEVIVGVIAEKHSADFGADSQEAMRKALELFRNDMASRYKELGFQG
: GLSDDEWHHVLGIWAKVEPDLSAHGQEVIIRLFQVHPETQERFAKFKNLKTIDELRSSEEVKKHGTTVLTALGRILKLKNNHEPELKPLAESHATKHKIPVKYLEFICEIIVKVIAEKHPSDFGADSQAAMRKALELFRNDMASKYKEFGFQG
: GLSDDEWNHVLGIWAKVEPDLSAHGQEVIIRLFQLHPETQERFAKFKNLTTIDALKSSEEVKKHGTTVLTALGRILKQKNNHEQELKPLAESHATKHKIPVKYLEFICEIIVKVIAEKHPSDFGADSQAAMKKALELFRNDMASKYKEFGFQG
: GLSDDEWNHVLGIWAKVEPDLTAHGQEVIIRLFQLHPETQERFAKFKNLTTIDALKSSEEVKKHGTTVLTALGRILKQKNNHEQELKPLAESHATKHKIPVKYLEFICEIIVKVIAEKHPSDFGADSQAAMKKALELFRNDMASKYKEFGFLG
:M E L S D Q E W K H V L D I W T K V E S K L P E H G H E V I I R L L Q E H P E T Q E R F E K F K H M K T A D E M K S S E K M K Q H G N T V F T A L G N I L K Q K G N H A E V L K P L A K S H A L E H K I P V K Y L E F I S E I I V K V I A E K Y P A D F G A D S Q A A M R K A L E L F R N D M A S K Y K E F G Y Q G
:
TEWEHVNKVWAVVEPDIPAVGLAILLRLFKEHKETKDLFPKFKEIPVQQLGNN EDLRKHGVTVLRALGNILKQKGKHSTNVKELADTHINKHKIPPKNFVLITNIAVKVLTEMYPSDMTGPMQESFSKVFTVICSDLETLYKEANFQG
:
ADFDAVLKCWGPVEADYTTMGGLVLTRLFKEHPETQKLFPKFAGIAQADIAGNAAISA HGATVLKKLGELLKAKGSHAAILKPLANSHATKHKIPINNFKLISEVLVKVMHEKAGLDAGGQTALRNVMGIIIA--DLEANYKELGFSG
:
VDWEKVNSVWSAVESDLTAIGQNILLRLFEQYPESQNHFPKFKNKSLGELKDTADIKAQAD TVLSALGNIVKKKGSHSQPVKALAATHITTHKIPPHYFTKITTIAVDVLSEMYPSEMNAQVQAAFSGAFKIICSDIEKEYKAANFQG
:
HDAELVLKCWGGVEADFEGTGGEVLTRLFKQHPETQKLFPKFVGIASNELAGNAAVKA HGATVLKKLGELLKARGDHAAILKPLATTHANTHKIALNNFRLITEVLVKVMAEKAGLDAGGQSALRRVMDVVIGDID--TYYKEIGFAG
:
ADWDKVNSVWSAMEANITAVGQNILLRLFEQYPESQSYFPKLKNKSLGELKDTADIKAQAD TVLKALGNIVKKKGNHSQPVKALAATHITTHKIPPHYFTKITTIAVGVLSEMYPSEMNAQAQEAFSGAFKSICSDIEKEYKAANFQG
:
ABWDKVNSVWSAVEQNITAIGQNILLRLFEQYPESEDYFPKLKNKSLGELKDTADIKAQAD TVLRALGNIVKKKGDHSQPVKALAATHITTHKIPPHYFTKITTIAVGVLSEMYPSEMNAQAQAAFSGAFKNICSDIEKEYKAANFQG
The multiple alignment of 56 lysozymes constructed with the genetic semihomology algorithm
The conservative and frequently occurring consensus residues are shown as white letters on black background
P00695:
P00696:
P00697:
P00698:
P00701:
P00702:
P00703:
P00705:
P04421:
P05105:
P07232:
P79158:
P08905:
P12068:
P12069:
P30200:
P30201:
P50717:
P50718:
P51782:
P79179:
P79239:
P79268:
P79294:
P79687:
P79698:
P79806:
P79811:
P79847:
Q05820:
P49633:
P00699:
P00700:
P00704:
P00706:
P00707:
P00708:
P11375:
P11376:
P12067:
P16973:
P17607:
P19849:
P22910:
P24364:
P24533:
P37156:
P37712:
P37713:
P37714:
P80190:
P81708:
P81709:
P81711:
Q91159:
P11941:
1
1
2
3
4
5
6
7
8
9
0
1
2
3
MKALIVLGLVLLSVT-VQG--KVFE-RCELARTLKRLGMDGYRGISLANWMCLAKWESGYNTRATNY-NAGDRSTDYGIFQINSRYWCNDGKTPGAVNACHLSCSALLQDNIADAVACAKRVVRDPQGIRAWVAWRNRCQNRDVRQYVQGCGV
MKAVIILGLVLLSVT-VQG--KIFE-RCELARTLKRLGLDGYRGISLANWVCLAKWESDYNTQATNY-NPGDQSTDYGIFQINSHYWCNNGKTPGAVNACHISCNALLQDNIADAVTCAKRVVSDPQGIRAWVAWRNHCQNRDVSQYVQGCGV
MKALLVLGFLLLSAS-VQA--KIYE-RCQFARTLKRNGMSGYYGVSLADWVCLAQHESNYNTQARNY-NPGDQSTDYGIFQINSRYWCNDGKTPRAKNACGIPCSALLQDDITQAIQCAKRVVRDPQGIRAWVAWQRHCKNRDLSGYIRNCGV
MRSLLILVLCFLPLA-ALG--KVFG-RCELAAAMKRHGLDNYRGYSLGNWVCAAKFESNFNTQATNR-N-TDGSTDYGILQINSRWWCNDGRTPGSRNLCNIPCSALLSSDITASVNCAKKIVSDGNGMNAWVAWRNRCKGTDVQAWIRGCRL
MRSLLVLVLCFLPLA-ALG--KVYG-RCELAAAMKRHGLDKYQGYSLGNWVCAAKFESNFNTQATNR-N-TDGSTDYGILQINSRWWCNDGRTPGSRNLCNIPCSALLSSDITASVNCAKKIVSDVHGMNAWVAWRNRCKGTDVNAWIRGCRL
MRSLLILVLCFLPLA-APG--KVYG-RCELAAAMKRMGLDNYRGYSLGNWVCAAKFESNFNTGATNR-N-TDGSTDYGILQINSRWWCNDGRTPGSKNLCHIPCSALLSSDITASVNCAKKIVSDGNGMNAWVAWRKHCKGTDVNVWIRGCRL
MRSLLILVLCFLPLA-ALG--KVYG-RCELAAAMKRLGLDNYRGYSLGNWVCAAKFESNFNTHATNR-N-TDGSTDYGILQINSRWWCNDGRTPGSKNLCNIPCSALLSSDITASVNCAKKIASGGNGMNAWVAWRNRCKGTDVHAWIRGCRL
MKALLTLVFCLLPLA-AQG--KVYS-RCELAAAMKRLGLDNYRGYSLGNWVCAANYESGFNTQATNR-N-TDGSTDYGILQINSRWWCDNGKTPRSKNACGIPCSVLLRSDITEAVRCAKRIVSDGDGMNAWVAWRNRCRGTDVSKWIRGCRL
MKALVILGFLFLSVA-VQG--KVFE-RCELARTLKKLGLDGYKGVSLANWLCLTKWESSYNTKATNY-NPSSESTDYGIFQINSKWWCNDGKTPNAVDGCHVSCRELMENDIAKAVACAKHIVSE-QGITAWVAWKSHCRDHDVSSYVEGCTL
MTKYVILLAVLAFALHCDA--KRF-TRCGLVQELRRLGFDETL-MS--NWVCLVENESGRFTDKIGKVNKN-GSRDYGLFQINDKYWCSKGTTPG--KDCNVTCNQLLTDDISVAATCAKKIYKRHK-FDAWYGWKNHCQ-HGLP-DISDC
MKALTILGLVLLSVT-VQG--KIFE-RCELARTLKKLGLDGYKGVSLANWVCLAKWESGYNTEATNY-NPGDESTDYGIFQINSRYWCNNGKTPGAVDACHISCSALLQNNIADAVACAKRVVSDPQGIRAWVAWRNHCQNKDVSQYVKGCGV
MKVLIILGLVLLSVM-VQG--KVFE-RCELARTLKRFGLDGYRGISLANWMCLAKWESDYNTRATNY-NPGDQSTDYGIFQINSHYWCNNGRTPGAVNACHISCNALLQDDITEAVACAKRVVRDPQGIRAWVAWKAHCQNRDVSQYVQGCGV
MKTLLTLGLLLLSVT-AQA--KVYE-RCEFARTLKRNGMAGYYGVSLADWVCLAQHESNYNTRATNY-NRGDQSTDYGIFQINSRYWCNDGKTPRAVNACGINCSALLQDDITAAIQCAKRVVRDPQGIRAWVAWRAHCQNRDLSQYIRNCGV
MKTLLVLALLLLSVS-VQA--KVYD-RCEFARILKKSGMDGYRGVSLANWVCLAKWESDFNTKATNH-N--VGSTDYGIFQINSRYWCNDGKTPKAVNACHISCKVLLDDDLSQDIECAKRVVRDPLGVKAWVAWRAHCQNKDVSQYIRGCKL
MKTLLVLALLLLSVS-VQA--KVYD-RCEFARILKKSGMDGYRGVSLANWVCLAKWESNFNTKATNY-NPGSQSTDYGIFQINSRYWCNDGKTPKAVNACHISCKVLLDDDLSQDIECAKRVVRDPQGIKAWVAWKAHCQNKDVSQYIRGCKL
MKAVIILGLVLLSVT-VQG--KIFE-RCELARTLKRLGLDGYRGISLANWVCLAKWESGYNTQATNY-NPGDQSTDYGIFQINSHYWCNNGKTPGAVNACHISCNALLQDNIADAVTCAKRVVRDPQGIRAWVAWRNHCQNRDVSQYVQGCGV
MKAVIILGLVLLSVT-VQG--KIFE-RCELARTLKRLGLDGYRGISLANWVCLAKWESNYNTQATNY-NPGDQSTDYGIFQINSHYWCNNGKTPGAVNACHISCNALLQDNIADAVTCAKRVVSDPQGIRAWVAWRNHCQNRDVSQYVQGCGV
MQKLAVFLFAIAAVC-IHCEAKYYSTRCDLVRELRKQGFPENQ---MGDWVCLVENESGRKTDKVGPVNKN-GSKDYGLFQINDKYWCSNTRTPG--KDCNVTCADLLLDDITKASTCAKKIFKRHN-FRAWYGWRNHCDGKTLP-DTSNC
MQKLRVFLLALAALC-ISCEAKYFATNCELVHELRRQGFPEDK---MRDWVCLIQNESGRNTSKMGTINKN-GSRDYGLFQINDKYWCSKTSTPG--KDCNVTCAEMLLDDITKASKCAKKIYKRHK-FQAWYGWRNHCQG-TLP-DISKC
MKVLLLLGFIFCSMA-AHG--KRME-RCEFARRIKQLHLDGYHQISLANWVCLAQWESGFDTKATNY-NPGDQSTDYGILQINSHYWCDDGKTPHAANECKVRCSELQEDDLVKAVNCAKKIV-DQQGIRAWVAWRNKCEGKDLSKYLEGCHL
MKALIVLGLVLLSVM-VQG--KVFE-RCELARTLKRLGMDGYRGISLANWMCLAKWESGYNTRATNY-NAGDRSTDYGIFQINSRYWCNDGKTPGAVNACHLSCSALLQDNIADAVACAKRVVRDPQGIRAWVAWRNRCQNRDVRQYVQGCGV
MKALIILGLVLLSVT-VQS--KVFE-RCELARTLKRLGMDGYRGISLANWMCLAKWESGYNTRATNY-NPGDRSTDYGIFQINSRYWCNDGKTPGAVNACHLSCSALLQDNIADAVACAKRVVRDPQGIRAWVAWRNRCQNRDVRQYVQGCGV
MKVLILLGLVLLSVM-VQG--KVFE-RCELARTLKRLGLDGYRGISLANWMCLAKWESDYNTRATNY-NPGDQSTDYGIFQINSHYWCNNGRTPGAVNACHISCNALLQDDITEAVACAKRVVRDPQGIRAWVAWKAHCQNRDVSQYIQGCGV
MKVLVILGLVLLSVM-VQG--KVFE-RCELARTLKRLGMDGYRGISLANWMCLAKWESDYNTRATNY-NPGDQSTDYGIFQINSHYWCNNGRTPGAVNACHISCNALLQDDITQAVACAKRVVRDPQGIRAWVAWKAHCQNRDVSQYVQGCGV
MKAVIILGLVLLSVT-VQG--KIFE-RCELARTLKRLGLDGYRGISLANWVCLAKWESDYNTQATNY-NPGDQSTDYGIFQINSHYWCNNGKTPGAVNACRISCNALLQDNIADAVTCAKRVVRDPQGIRAWVAWRNHCQNRDVSQYVQGCGV
MKALIILGLVLLSVT-VQG--KIFE-RCELARTLKKLGLDGYKGVSLANWVCLAKWESGYNTDATNY-NPGDESTDYGIFQINSRYWCNNGKTPGAVNACHISCNALLQNNIADAVACAKRVVSDPQGIRAWVAWKKHCQNRDVSQYVEGCGV
MKAVIILGLVLLSVT-VQG--KIFE-RCELARTLKRLGLDGYRGISLANWVCLAKWESDYNTQATNY-NPGDQSTDYGIFQINSHYWCNNGKTPGAVNACHISCNALLQDNIADAVTCAKRVVRDPQGIRAWVAWRNHCHNRDVSQYVQGCGV
MKALIILGLVLLSVT-VQG--KIFE-RCELARTLKKLGLDGYKGVSLANWVCLAKWESGYNTEATNY-NPGDESTDYGIFQINSRYWCNNGKTPGAVDACHISCSALLQNNIADAVACAKRVVSDPQGIRAWVAWRNHCQNRDVSQYVKGCGV
MKALIILGLVLLSVT-VQG--KIFE-RCELARTLKKLGLDGYKGVSLANWVCLAKWESGYNTEATNY-NPGDESTDYGIFQINSRYWCNNGKTPGAVDACHISCSALLQNNIADAVACAKRVVSDPQGVRAWVAWRNHCQNKDVSQYVKGCGV
MKALLVLGFLLLSAS-VQA--KVFK-HCELARILRSSALAGYRGVSLENWMCMAQHESNFDTEATNY-NSTDQSTDYGIFQINSRYWCNDGKTPRAVNACGIPCSALLQDDITQAIQCAKRVVRDPQGIRAWVAWQRHCQNRDLSGYIRNCGV
------------------G--KVYG-RCELAAAMKRMGLDNYRGYSLGNWVCAAKFESNFNTGATNR-N-TDGSTDYGILQINSRWWCNDGRTPGSKNLCHIPCSALLSSDITASVNCAKKIVSDGDGMNAWVAWRKHCKGTDVNVWIRGCRL
---------------------KVFG-RCELAAAMKRHGLDNYRGYSLGNWVCAAKFESNFNSQATNR-N-TDGSTDYGVLQINSRWWCNDGRTPGSRNLCNIPCSALLSSDITATVNCAKKIVSDGNGMNAWVAWRNRCKGTDVHAWIRGCRL
---------------------KVFG-RCELAAAMKRHGLDNYRGYSLGNWVCAAKFESNFNSQATNR-N-TDGSTDYGVLQINSRWWCNDGKTPGSRNLCNIPCSALLSSDITATVNCAKKIVSDGNGMNAWVAWRNRCKGTDVQAWIRGCRL
---------------------KVFG-RCELAAAMKRHGLDNYRGYSLGNWVCAAKFESNFNSQATNR-N-TDGSTDYGVLQINSRWWCNDGRTPGSRNLCNIPCSALQSSDITATANCAKKIVSDGNGMNAWVAWRKHCKGTDVRVWIKGCRL
---------------------KVYE-RCELAAAMKRLGLDNYRGYSLGNWVCAANYESSFNTQATNR-N-TDGSTDYGILEINSRWWCDNGKTPRAKNACGIPCSVLLRSDITEAVKCAKRIVSDGDGMNAWVAWRNRCKGTDVSRWIRGCRL
---------------------KIYK-RCELAAAMKRYGLDNYRGYSLGNWVCAARYESNYNTQATNR-NS-NGSTDYGILQINSRWWCNDGRTPGTKNLCHISCSALMGADIAPSVRCAKRIVSDGDGMNAWVAWRKHCKGTDVSTWIKDCKL
---------------------KDI-PRCELVKILRRHGFEGFVGKTVANWVCLVKHESGYRTTAFNN-NGPN-SRDYGIFQINSKYWCNDGKTRGSKNACNINCSKLRDDNIADDIQCAKKIAREARGLTPWVAWKKYCQGKDLSSYVRGC
---------------------KVFS-KCELAHKLKAQEMDGFGGYSLANWVCMAEYESNFNTRAFNGKNA-NGSYDYGLFQLNSKWWCKDNKRSSS-NACNIMCSKLLDDNIDDDISCAKRVVRDPKGMSAWKAWVKHCKDKDLSEYLASCNL
---------------------KVFS-KCELAHKLKAQEMDGFGGYSLANWVCMAEYESNFNTRAFNGKNA-NGSSDYGLFQLNNKWWCKDNKRSSS-NACNIMCSKLLDENIDDDISCAKRVVRDPKGMSAWKAWVKHCKDKDLSEYLASCNL
---------------------KVYD-RCEFARILKKSGMDGYRGVSLANWVCLAKWESDFNTKAINR-N-V-GSTDYGIFQINSRYWCNDGKTPKAVNACHISCKVLLDDDLSQDIECAKRVVRDPQGIKAWVAWRTHCQNKDVSQYIRGCKL
---------------------KIYE-RCELARTLKKLGLDGYKGVSLANWMCLAKWESSYNTRATNY-NPGDKSTDYGIFQINSRYWCNDGKTPRAVNACHIPCSDLLKDDITQAVACAKRVVSDPQGIRAWVAWRNHCQNQDLTPYIRGCGV
---------------------KVFE-RCELARTLKELGLDGYKGVSLANWLCLTKWESSYNTKATNY-NPGSESTDYGIFQINSKWWCNDGKTPNAVDGCHVSCSELMENNIAKAVACAKHIVSE-QGITAWVAWKSHCRDHDVSSYVEGCSL
---------------------KVYG-RCELAAAMKRLGLDNYRGYSLGNWVCAAKFESNFNTHATNR-N-TDGSTDYGILQINSRWWCNDGRTPGSRNLCNIPCSALLSSDITASVNCAKKIVSDRNGMNAWVAWRNRCKGTDVHAWIRGCRL
---------------------KVYG-RCELAAAMKRLGLDNYRGYSLGNWVCAAKFESNFNTHATNR-N-TDGSTDYGILQINSRWWCNDGRTPGSRNLCHIPCSALLSSDITASVNCAKKIVSDGNGMNAWVAWRNRCKGTDVNAWIRGCRL
---------------------KVYG-RCELAAAMKRLGLDNYRGYSLGNWVCAAKYESNFNTHATNR-N-TDGSTDYGILQINSRWWCNDGKTPGSRNLCHIPCSALLSSDITASVNCAKKIVSDGNGMNAWVAWRNRCKGTDVSVWTRGCRL
---------------------KVYG-RCELAAAMKRLGLDNYRGYSLGNWVCAAKFESNFNTHATNR-N-TDGSTDYGILQINSRWWCNDGRTPGSRNLCHISCSALLSSDITASVNCAKKIVSDRNGMNAWVAWRNRCKGTDVNAWIRGCRL
----------------KIL--KKQE-LCKNLVA---QGMNGYQHITLPNWVCTAFHESSYNTRATNH-N-TDGSTDYGILQINSRYWCHDGKTPGSKNACNISCSKLLDDDITDDLKCAKKIAGEAKGLTPWVAWKSKCRGHDLS--KFKC
---------------------KVWE-RCALARKLKELGMDGYRGVSLANWMCLTKWESDYNTDATNY-NPSSESTDYGIFQINSRYWCNNGKTPHAVNGCGINCNVLLEDDITKAVQCAKRVVRDPQGVRAWVAWKNHCEGHDVEQYVEGCDL
---------------------KVFE-RCELARTLKKLGLDDYKGVSLANWLCLTKWESGYNTKATNY-NPGSESTDYGIFQINSKFWCNDGKTPDAVDGCHVSCSELMENDIEKAVACAKHIVSE-QGITAWVAWKSHCRDHDVSSYVEGCTL
---------------------KVFE-RCELARTLKELGLDGYKGVSLANWLCLTKWESSYNTKATNY-NPGSESTDYGIFQINSKFWCNDGKTPNAVDGCHVSCSELMENNIAKAVACAKQIVSE-QGITAWVAWKSHCRDHDVSSYVEGCTL
---------------------KVFE-RCELARTLKRFGMDGFRGISLANWMCLARWESSYNTQATNY-NSGDRSTDYGIFQINSHWWCNDGKTPGAVNACHIPCSALLQDDITQAVACAKRVVSDPQGIRAWVAWRSHCQNQDLTSYIQGCGV
---------------------KIFS-KCELARKLKSMGMDGFHGYSLANWVCMAEYESNFNTQAFNGRNSNGSS-DYGIFQLNSKWWCKSNSHSSA-NACNIMCSKFLDDNIDDDIACAKRVVKDPNGMSAWVAWVKHCKGKDLSKYLASCNL
---------------------KIFE-RCELARTLKNLGLAGYKGVSLANWVCLAKWESNYNTRATNY-NPGSKSTDYGIFQINSRYWCNDGKTPRAVNACHISCSALLQDDITQAVACAKRVVSDPNGIRAWVAWRAHCENRDVSQYVRNCGV
---------------------KVYG-RCELAAAMKRLGLDNFRGYSLGNWVCAAKFESNFNTHATNR-NTDGS-TDYGILQINSRWWCNDGRTPGSRNLCNIPCSALLSSDTIASVNCAKKIVSDGNGMNAWVAWRKRCKGTDVNAWTRGCRL
MLFFGFLLAFLSAVPGTEGEII---PRCELVKILREHGFEGFEGTTIADWICLVQHESDYNTEAYNN-NGPSR--DYGIFQINSKYWCNDGKTSGAVDGCHISCSELMTNDLEDDIKCAKKIARDAHGLTPWYGWKNHCEGRDLSSYVKGC
MRAVVVLLLVAVASA------KVYD-RCELARALKASGMDGYAGNSLPNWVCLSKWESSYNTQATNR-NTDGS-TDYGIFQINSRYWCDDGRTPGAKNVCGIRCSQLLTADLTVAIRCAKRVVLDPNGIGAWVAWRLHCQNQDLRSYVAGCGV
Relative amino acid composition of eglin-like
and Bowman-Birk inhibitors
Amino acid contents [%]
25
20
15
10
5
0
A C D E F G H
I
K
L M N P Q R S
Amino acid
T V W Y
Eglins
Bowman-Birk inhibitors
The amino acids occurring at variable positions of eglin
and Bowman-Birk family
EGLIN-LIKE PROTEINS
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
KR
-ELNQRST
EFMQRST
FW
P
EHQ
LV
CILV
EG
ACKLMSTV
DGPRST
AGITV
ADEKLS
ADEFKLQVY
A
AEKMRV
AEGKQT
-U1IKTVY
FIV
EKLQT
AEKLQRT
DEHQ
KMNRY
PSV
DEKLNQRS
AILVY
DEKQRST
-AINV
-E
-V
-EHIQY
-FILTV
-LMSV
-P
-EHIQRS
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
-EILV
-DL
-ANS
DGM
GQSTV
AFHINPV
-W
-AFIV
-T
AEKLMQT
DEN
EFILY
DKLNR
CFILPY
DEKNQ
R
IV
FR
ILV
FLWY
DNVY
ADHNT
-DEIKLPRV
-AGLNRS
-DGNT
-DFIKLNSTY
IV
ANTV
DKNQRSZ
AHIMPTV
-ASV
-P
-AHKQRSTV
IV
AGT
BOWMAN-BIRK INHIBITORS
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
-DEKQST
-RSTVY
-KPST
EGHKPSTW
AEGKP
C
C
DNRS
EFHIKLQRST
ACQ
-ADFIKLMPRSTV
C
C
AKR
S
DEFIKMNQR
P
AP
EFIKMQT
C
HQRSTV
C
AEHMNQRSTV
DNQ
-IKLMQTV
GLRV
DEFIL
DEKNQRT
-S
C
AHPS
ADS
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
ADET
C
DEKLNS
ADEFGHKLRST
C
AEGILMV
CEKP
ANRSTV
EFHKLRTVY
DGS
DEFIMNQSY
-DPS
-AGPS
KLMPQR
CHR
FHIQRSVY
-I
C
ABEFGLQTVY
DN
-IMQTV
DHKNQTY
-DHIKNRTV
-FGY
-CDI
-HPTY
-ADEGKP
-AKPQS
-MT
-C
-DEHKNR
-DENPS
The position variability patterns of myoglobins and lysozymes
N u m b e r o f a m in o a cid s
M Y O G L O B IN S
12
10
8
6
4
2
0
1
11
21
31
41
51
61
71
81
91
101
111
121
131
141
151
121
131
P o s itio n n u m b e r
N u m b er of am in o acid s
LYS O ZYME S
12
10
8
6
4
2
0
1
11
21
31
41
51
61
71
P o s itio n n u m b e r
81
91
101
111
The observed number and contribution of three correlation types in four
different protein families
The correlation sets consist of 2 to over 20 residues
The correlation statistics
The protein
family (number of
correlated
positions/set)
Total
number of
correlation
sets
observed
Number of
dispersed
sets
Number of
narrow
clusters
Number of
undirected
clusters
Number of
sets
related to
active
center
20
7
7
6
1
23
4
13
6
9
Myoglobins (229)
41
23
9
9
n.a.
Lysozymes (2-15)
41
25
9
7
9
All families
125 (100%)
59 (47.2%)
38 (30.4%)
28 (22.4%)
-
Eglin-like
proteins (2-13)
Bowman-Birk
proteinase
inhibitors (2-28)
Program FEEDBACK – what does it do?
The program FEEDBACK is designed to analyze the multiple
aligned protein sequences for correlated mutations occurrence.
It returns in result all possible residues occurring at all
sequence positions of aligned proteins for each residue
occurring at each position.
The result visualization is assisted by MS EXCEL.
This application is available as freeware upon request.
KR
-E L N Q R S T E F M Q R S T F W
P
EH Q
LV
C IL V
EG
ACKLM S TV
DGP RS T
A G IT V
(1 )(K )2 4
K
-E L N Q R S T E F M Q R S T F W
P
EH Q
LV
C IL V
EG
ACKLM S TV
DGP RS T
A G IT V
(1 )(R ) 1
R
T
S
W
P
E
L
V
G
V
S
A
(2 )(-) 1
K
-
S
F
P
E
V
V
G
K
T
V
(2 )(E ) 6
K
E
RS T
W
P
E
L
IL
G
TV
P
A
(2 )(L ) 3
K
L
QRS
W
P
E
L
IV
G
KV
DP
GT
(2 )(N ) 1
K
N
T
W
P
E
L
C
G
A
R
G
(2 )(Q ) 3
K
Q
FM R
W
P
E
L
I
G
V
P
AT
(2 )(R ) 2
K
R
S
W
P
EQ
LV
V
G
MS
ST
AG
(2 )(S ) 1
K
S
S
W
P
H
L
V
G
V
G
G
(2 )(T ) 8
KR
T
ES
W
P
E
LV
V
EG
CKLM V
ST
A IV
(3 )(E ) 3
K
T
E
W
P
E
L
V
EG
CK
ST
IV
(3 )(F ) 1
K
Q
F
W
P
E
L
I
G
V
P
A
(3 )(M ) 1
K
Q
M
W
P
E
L
I
G
V
P
T
(3 )(Q ) 1
K
L
Q
W
P
E
L
I
G
V
P
T
(3 )(R ) 4
K
EL Q
R
W
P
E
L
IL
G
TV
P
AT
(3 )(S )1 1
KR
-E L R S T
S
FW
P
EH Q
LV
LV
G
KLM S TV
DGP S T
AGV
The three types of distribution of correlated positions present in eglin-like
proteins.
The residue location and relative distribution is shown on tertiary structure of eglin C
(P01051)
The dispersed correlation
Position no. and
occurring residues
67 [–DGNT]
10 [–ELNQRST]
Correlation
versus position 67
D (8)
G (9)
ET
LNQRS
The three types of distribution of correlated positions present in eglin-like
proteins.
The residue location and relative distribution is shown on tertiary structure of eglin C
(P01051)
The narrow correlation cluster
Position no. and
occurring residues
34 [DEKQRST]
Correlation
versus position 34
T (10)
Q (6)
15 [CILV]
CIL
V
17 [DGPRST]
PR
ST
27 [EKLQT]
EQ
KL
28 [AEKLQRT]
KT
EQR
30 [KMNRY]
N
KM
KLS
DEN
C
IP
DFI
–KNT
32 [DEKLNQRS]
56 [CFILPY]
68 [–DFIKLNSTY]
The three types of distribution of correlated positions present in eglin-like
proteins.
The residue location and relative distribution is shown on tertiary structure of eglin C
(P01051)
The spot correlation cluster
Position no. and
occurring residues
Correlation
versus position 30
30 [KMNRY]
K (6)
N (15)
18 [DGPRST]
S
DGPRT
27 [EKLQT]
L
EQ
29 [DEHQ]
D
EQ
33 [AILVY]
A
ILV
35 [–AINV]
IV
–AN
–NS
DFIKLS
Y
68 [–DFIKLNSTY]
The three types of distribution of correlated positions present
in Bowman-Birk inhibitor family
The residue location and relative distribution is shown on tertiary structure of BowmanBirk inhibitor from soybean (P01055)
The dispersed correlation
Position no. and
occurring residues
29 [DEFIL]
Correlation
versus position 29
L (37)
E (12)
EGKST
W
–AFILPRT
M
40 [AEGILMV]
AILMV
E
48 [KLMPQR]
KLMQ
R
6
[EGHKPSTW]
13 [–ADFIKLMPRSTV]
The three types of distribution of correlated positions present
in Bowman-Birk inhibitor family
The residue location and relative distribution is shown on tertiary structure of BowmanBirk inhibitor from soybean (P01055)
The narrow correlation cluster
Position no. and
occurring residues
13 [–ADFIKLMPRSTV]
Correlation
versus position 13
L (11)
M
(10)
A (8)
4 [–RSTVY]
V
–S
S
5 [–KPST]
K
–S
S
7 [AEGKP]
A
P
P
11 [EFHIKLQRST]
T
EHQ
S
21 [EFIKMQT]
T
Q
EQ
The three types of distribution of correlated positions present
in Bowman-Birk inhibitor family
The residue location and relative distribution is shown on tertiary structure of BowmanBirk inhibitor from soybean (P01055)
The spot correlation cluster
Position no. and
occurring residues
25 [AEHMNQRSTV]
11 [EFHIKLQRST]
23 [HQRSV]
50 [FHIQRSVY]
Correlation
versus position 25
A (15)
V (9)
EFKLRS
HQ
Q
R
HRS
FI
The three types of distribution of correlated positions present in
myoglobins.
The residue location and relative distribution is shown on tertiary structure of human
myoglobin (P0244, pdb1bzp)
The dispersed correlation
Position no. and
occurring residues
74
[AGPQST]
Correlation
versus position 74
A (6)
G (49)
N (9)
128 [ABEHQ]
Q
BEHQ
Q
137 [ILNSV]
L
L
ILNSV
The three types of distribution of correlated positions present in
myoglobins
The residue location and relative distribution is shown on tertiary structure of human
myoglobin (P0244, pdb1bzp)
The narrow correlation cluster
Position no. and
occurring residues
Correlation
versus position 23
23 [AEGST]
A (7)
G (55)
S (10)
22 [AEGPSTV]
PST
AEGP
STV
P
26 [EGHKLQ]
LQ
EGH
QK
Q
27 [ADEFLNT]
AEN
ADEF
T
E
IL
IMTV
I
53 [ADEGQ]
DEQ
ADEG
D
54 [ADEILQ]
AEL
DELQ
E
59 [ADEF]
DE
ADEF
E
Q
ABEH
Q
Q
30 [ILMTV]
128 [ABEHQ]
The three types of distribution of correlated positions present in
myoglobins
The residue location and relative distribution is shown on tertiary structure of human
myoglobin (P0244, pdb1bzp)
The spot correlation cluster
Position no. and occurring
residues
127 [AMSTV]
27
[ADEFLNT]
31
78
Correlation
versus position 127
A (58)
S (7)
ADEFNT
E
[GKRS]
GKRS
R
[AKLQ]
K
ALQ
DEGT
E
AEHKQS
A
AEKQS
E
BDEN
D
109 [DEGNT]
116 [AEHKQST]
117 [AEKNQS]
122 [BDEN]
The three types of distribution of correlated positions present in
lysozymes
The residue location and relative distribution is shown on tertiary structure of lysozyme
from rat (P00697, pdb5lyz)
The dispersed correlation
Position no. and occurring
residues
80 [GHKNR]
Correlation
versus position 80
G (7)
H (31)
N (16)
30 [ILMV]
MV
ILMV
V
40 [DFKNR]
DN
N
FKNR
The three types of distribution of correlated positions present in
lysozymes
The residue location and relative distribution is shown on tertiary structure of lysozyme
from rat (P00697, pdb5lyz)
The narrow correlation cluster
Position no. and occurring
residues
Correlation
versus position 59
59 [FL]
F (38)
L (18)
26 [–ILMV]
–ILMV
L
33 [AISTV]
AISTV
A
44 [FIMRTVY]
FIMRTVY
T
54 [–KRSTY]
–KRSTY
T
84 [AKNRS]
AKNRS
S
The three types of distribution of correlated positions present in
lysozymes
The residue location and relative distribution is shown on tertiary structure of lysozyme
from rat (P00697, pdb5lyz)
The spot correlation cluster
Position no. and
occurring residues
126 [–DWY]
Correlation
versus position 126
W (16)
Y (36)
13 [–ILM]
M
IL
15 [–AEKNQRS]
R
AEKNQ
RS
20 [DEGKN]
KN
DG
27 [–AEGPR]
G
AEP
44 [FIMRTVY]
T
FIRTY
46 [GHKNPRTY]
R
GHNRY
105 [–AGHPQRV]
GRV
–APQ
109 [DGKNQRST]
N
GKRST
121 [–HKQRT]
T
HKQR
CONCLUSIONS
Almost 50% of the observed correlated mutations refer to residues
that are not in contact nor interact with each other
The dispersed correlations are present in various protein families
and they occur independently on the mechanism of structure stabilization
The phenomenon of correlated mutations is not limited to interacting
residues and/or known biological activity determination
The current hypothesis of positive Darwinian selection does not fully
explain the mechanism and occurrence of correlated mutations
Correlated mutations team
Similarity estimation team
Łukasz Becella1
Monika Sobczyk1
Monika Grabiec1
Jacek Leluk1,2
1 Institute
of Biochemistry and Molecular Biology, Univeristy of Wrocław
2 Interdisciplinary Centre for Mathematical and Computational Modelling,
University of Warsaw