10 20
30 40
50 60
70 80
90
100
110
120
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
structure
Kluyveromyces lactis ------------------------------------------------------------------------------------------------------------------------
Kluyveromyces wickerhamii
------------------------------------------------------------------------------------------------------------------------
Kluyveromyces nonfermentans ------------------------------------------------------------------------------------------------------------------------
Kluyveromyces marxianus AAUUGUAUACUUUUUAAACAACCUAAACAUGUUGUUACGUUCUAUUAUGUUCAGUAACAAUAUAGACAAUUUACCCUGCGUGUUCGAUUCUAUUUGCCAAAGCCUAGAGAGUGAUUGAGG
Kluyveromyces dobzhanskii
------------------------------------------------------------------------------------------------------------------------
Kluyveromyces aestuarii ------------------------------------------------------------------------------------------------------------------------
130
140
150
160
170
180
190
200
210
220
230
240
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
structure
Kluyveromyces lactis --GAAUAUAUGAUGGGUGCGGAAUCACGAUCGGUAUUUGUGUGAUGCAAUUUCUGGGGUGGUAAGGACCAGUGCCACACUUGGGUUUGAUUUUCCUCUCAUCGAGUUUCGUGUGCGCAUG
Kluyveromyces wickerhamii
--AAAAUUUACUGCGAUCUCAACGAACAUAAGCUACUAAAUCUAUUUCAAUUAUUCCAAAACUUAUUUCACGAUGGGAACAAGGAGGCCUUUGGUAGAUCUGGUUGCUUACGAGGAGGUA
Kluyveromyces nonfermentans ---------------------------------------------------------------GUAAAGAAGAUGUGUUAGGUUUACAACGCUCUUCUCAGCAUUGUAUUGAAGUUUGUU
Kluyveromyces marxianus UGGUUAGGACCAGUACCACGUAUGAGACGUGUUUAACUUUUUCCAAAUGCUGACAAGGUAUAUGUCUCGUAUUUCCCUGACGUUCUCUUUACGCCUGGGAAUUGACUGAGUGGACACUC-
Kluyveromyces dobzhanskii
-GAACUCCAUGAGUUCGUAAUCAGCAAGGAUUGUGGUAUUAUUUGCUGAAAGACGAAGUGUGAACGAAGUUUGAGGUGGUAAGGACCAGUGCCAUGCUUGAGUGUUUCUUUCUGAGUUUU
Kluyveromyces aestuarii -------------------------------------------------------AGCAAAGCAUUUGAAAGCUGAUUUGAAUUUCUAUCGUUCUUUUCAAAACUUCAUUGAAGAUAGUU
250
260
270
280
290
300
310 320 330 340 350 360
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
structure
Kluyveromyces lactis UUCGUCUCUCACAUCGAGUUAUACUCUCAUCUUCUAUACAUUAGAAGGAUGCAUCAGCAGUUUGAUCUGCUGAAAGAGUUCAUACACCCAAUUAUUGUUUAAAGGCCUGGUGUUGGGUAG
Kluyveromyces wickerhamii
AUGGAGCCAGUACCUUAUCGCCACUUUUCGAUUGACACCAAAGCUCCCACAACGCGAACGUGAAACGCAUGACUGUUCU-----------------------------------------
Kluyveromyces nonfermentans UAGAUAGAGGCGGUUUUGGAACCUCACCUGAUGUACUCUUUCAAAUAUUC-UGUGAUAUU-----ACAAUGUUGCCCUUAGUGAUUG---CAUGUGACCUGUUUUGCGGGAGAAGCGGAC
Kluyveromyces marxianus CAAAGCUUUAGAUAUACCUGGC--------------------------------------------------------------------AUGCUAGAUCGUAUGCAUAUAUUUACGGAU
Kluyveromyces dobzhanskii
AGGUGUGCAUGUUUGUGUGUUUGACGACAUAGUCCCAUUCGUGUAUCUGACAUACCAGAAGUACACGGAAGGAAACUGAGUUUGUUGAAUGAAGUGAGGUGCAGCCACACAGCGAUCAAA
Kluyveromyces aestuarii UAAUUGGGGGCGGUUUUGAACCCUGGCCUGAUAGUUCCUUCAAAAGGUGCAUUUGAUUCUCACGUACAACUUCGCCUCACCCCAUUGAAGUUUGCCAGUAAUUCAGCUAAAUAAGCGGAC
370
380
390
400
410
420
430
440
450
460 470 480
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
structure
Kluyveromyces lactis UAUUUCUAACGGAUUUUAGAAAUGCUAUGACAACAAUACCUUUAGAAUUAUAUUGUGUGGAUAUUUACUUCUUGUUAAACAGACUGUGACAUAUCUCAAUGAGAGGGGAUUAUUCUCGUU
Kluyveromyces wickerhamii
---------------------AUGCACUGUUUAUUUACAAAGCAUUCGGAAAAUACGAUGUAAAUAUGAGGUGUACGCACAAAGUUGUCGAUUGUAAAUGACCUAUCUAACUCAGUUCUA
Kluyveromyces nonfermentans CUUACGACGCUGAAUAGGAAAAUUAGUUUGAACUAAUAUUAUAUCACAUGGAGUUUCAAGGGGUGGAUAAGGUAGCAGUCUAACUGCCUGUCAGGGUAUCAUAUGGAAGUGUAUAUCA-G
Kluyveromyces marxianus UAAAGAUAUGUGUAUGAAUAUGGCGCUAGGAAUUUUGUGCUUAAAAGAGAGUUUGCGUUUAA-UAGGUCUUUCGCAGACUAUACCGGAUAAUACCAAUUUCGGAGGCAUGAGGAUUGGGG
Kluyveromyces dobzhanskii
CCAGUGCUGUUGUUCGGUGCUUCUGACGGAUUUGGAAAACCUGAAUUGAUACGUGCCUUUUGAAAUUUUACUGGGUUGCAGUGUACCUUACUUUACCAGACUGACUCUUGUCUCGAUGGG
Kluyveromyces aestuarii CUUACGAAACUGAACUCAGGCAAACUC-CCUGUAUGAAAAUGAC--UGCUUGGAGCUGUUUGGA-UGAAGAAUGUAAGGGAUAGAUCUUAUGCAAUUGCCUUG---AGAAGUGAACUUCA
490
500
510
520
530
540
550
560
570
580 590 600
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
Template
structure
||||||||||||||||||||||||||||||
Kluyveromyces lactis UUUGGAGUUAUUCUGAUGCCUGUUGCUCUUGCUCCUACGCAUAGAAGCAUAGCAUUGAACUCCAUGGUGUGGUACUUCAAAUCCGUACACCACAUACCUAAUCAAAACCUCCUC-GCAAC
Kluyveromyces wickerhamii
UAGGCCUUACACAUGUUUCAAACCUUGGUCUAGCUUUAAUAUGGUGCAGUGCGAAGAGUUAACAGAUUUGGUACUAU-AAAUCCGUACACCACAUACCUAAUCAAAACCGCCUU-AGAAC
Kluyveromyces nonfermentans GUGUA-----------------------------------------------------------------------UCAAAUCCGUACACCACAUACCUAAUCAAUCCUUUUUUUAGGAC
Kluyveromyces marxianus AGGCUCGCGCUC-AGUUAGAGAUGGUUAAGC--UCUCAAUG-GGAGUUUA-CAUCCAUUUUAUAAACGUGGUACUUUCAAAUCCGUACACCACAUAACUAAUCAAAACCGCUUC-UAAAC
Kluyveromyces dobzhanskii
GAAAUACUUUAGUAUUGAAGUUCUUUCAACGUUGCACUCUUGUGCACAAUGCCUGGAAUUGUAGCGUUGAUCUUCAUAGCAU-GGCACUUCAAAUCCGUACACCACAUACCUAAUCAAAA
Kluyveromyces aestuarii -GGUGUA---------------------------------------------------------------------UCAAAUCCGUACACCACAUACCUAAUCAAUCCUUUUU--GGAAC
610
620
630
640
650
660
670
680
690
700 710 720
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
|----CS2----|
structure
((((((((( (((
Kluyveromyces lactis GGCUUCCAUAUUCUCGUCUGUAUUCAAUCG-UGGAUAGUGGUUUCAGCUCGACGUGGGUAUGCUAGUGACUCAGAGUUUGGAUGUACUA-GGUUUCAUAAG-------------------
Kluyveromyces wickerhamii
GGUU---AAACUCUGGUGUGUUUC-AUUUG--GCACUGUGGUACGGCCGUUA----GCUACGGCUCUAAUUCAGGGUUUAGA----UCAAGUUUACAAACGGAU----------------
Kluyveromyces nonfermentans AGCCGU--GUUUCUGUUUCAUUGACAACGA----AUAGCGGC-GGAUAAAGCCAGCACUAA--UCGGAAUUGAGGGUUUGGA----UCAAGGUUACAUCAG-------------------
Kluyveromyces marxianus GGCUUUAAA-UUCAUAUCUGUAUCAACUUGAGGGAUAGCGGCGUCGAUUCGA-AAGGGCAA--CAGUGAUUCAGGAUGUGGAAAAGUCAAGGUUGCAUAAG-------------------
Kluyveromyces dobzhanskii
CCGCUUUGAAACGGCUCCCAUUCUCGUCUGUUUCUACCCUGGGAUAGUGUUUUCAAUUCGACGUGGGUAAGCUGCAGAUUCAGGGUUUGGGGCACUAGGUUGCAUAAGAUUUGCAGCAAC
Kluyveromyces aestuarii UGCCAU--AUUUUUUGUUCACUGGCAACCA----AAAGCGGCCAAAGAGGGCUAGCAUCAG--UUGGAAUUAAGGGUUUGGA----UCAAGGUUACACAAA-------------------
730
740
750
760
770
780
790
800
810
820 830 840
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
|------CS2------|
structure
))))))) )))))
Kluyveromyces lactis ---------------------------------------------GAUACCCAACUG-------------GAAAUGUUGC-GGAAAUAUCACAUUCGUCCUUGCUCCAUUGUUAACUCAG
Kluyveromyces wickerhamii
--------------------------------------AACUUGUGUUCUGCAGGU---------------AACUGUGACCAAGUUGCAUCACUUAUGCUUGAUUCUAUGGUUAACCCAG
Kluyveromyces nonfermentans -----------------------------------------------------------------------GAAGGUUUU--GCUUUCAUAUUUUCUUUGCGAAUCCAUGGUUAACCCCG
Kluyveromyces marxianus --------------------------------------GAUGCAUGAGAUGAAAUUC--------------AAAUGUGUCAAAAACUCCUCGUUCCAUCGUUAAUCCGGUGU-CAUUUAG
Kluyveromyces dobzhanskii
AUUCAGUUUCCAUGUAUGGUGCGAAAAGGCAGUAAUACUGUAGGAGAAGGCUGAUCCUUAAUUUUUUAAGUUCAGCUGGAAUGUUGCCAUAAAAAUCACCAUUAUCCUGGCCCCAUUGUU
Kluyveromyces aestuarii -----------------------------------------------------------------------GAACAUUAC--AUUUUCACAUUUGCCUUGCAUUUCCAUGGUUAACCCAG
850
860
870
880
890
900
910
920
930
940
950
960
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
structure
Kluyveromyces lactis UGUC-ACUUUAGGUCCUAUGGGAAACAAUGUUGAGCUGAUGCAUU--UUUCCG-UACAGCCA--UUACCGUUCUUCGGGAUGGCAAGGAUUAGCCCUUCCUAUUACAGGCUUUAUAAUGG
Kluyveromyces wickerhamii
UGUG-AGAGUUGUUACAAUUAAUUUAUAUG-CGAUGAGCUGUUCUUUGAUCUG-UCAUACCAUGUU-UCGUUCUUCGUCAUGGAAAGGAU--AUCCCAGCCGUUACACACUUU-------
Kluyveromyces nonfermentans UGUU----UUGAUCGGUAUGAAAAAGU-UGUCCAACGGGAAAGUACAGAAACC-UUUAUUCAACAUUUGAUAAAAUGAAUUGAUGGAAAUCUGGC---ACCGUUAUGAAAUUC-------
Kluyveromyces marxianus UGCCUAUCGCAAGAGACAAAAAAAACUG-AUCGAGUCGACAUUUUUUAAUUUGCUUGAGUCAUGUUUCUGUCCUUCAGUAUGACGAGAAUUAGCCCUUCCUGUUACAGAUUUUACGGCGG
Kluyveromyces dobzhanskii
AACUCAGCGUCAUUGCAGUCCCAUGGGAAACAAUGUCGAAUAGUGUAUUAAUCUAACAAGCCAUAUUAUCGUUUUCCGAAAUGGUGAGGAUAGACCUUCCUGUUACAGA-UUUACAACGG
Kluyveromyces aestuarii UGUU----CUAACUGAUAUGAAAAAAU-UGCCUAACGGAAAAAUUCAGAUUCC-UUACUUCCUUCUUUGAUUGAAUGGAAUCGUAAAUCUCUGAC---ACCGUUGUGAACAUG-------
970
980
990
1000
1010
1020
1030
1040
1050
1060
1070
1080
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
CS3 CS3
structure
(((((( ))))))
Kluyveromyces lactis AAAACUCU---UAAGAUAUAUUUCUAGGUUUUUUUCCAUGAC-AGAGAA-UUAAGCCUGUGGUUCAUGUUGAUAUUCCAGAAUACGCAACAAUUGCUGCGUAUUCGGUA--GGUUUCUUU
Kluyveromyces wickerhamii
----------------CA-------------------UGUUC-AAAGAG-UUUAGCCUGUGGUUCAGUUUCAUGAUUUCGUCAUUCAAUGAUGUAUUUCUCGUC-------GGUCUCUUU
Kluyveromyces nonfermentans -----------------AUCUUCUCCG--------GAAGAA--GAAGCA---CGUGCUGCGACGUCAUCACGGAGACAAUG-ACAUGUAUUCUAC----------------GACUUGUUU
Kluyveromyces marxianus ACUUUUUUCCUUUUCAUAUCUGUGAGAGGAAACCGCUAAGU--AAAAGAAUAAAGCCUGUGGUUCAUUUCCAUUGUCUCUUUUCUUCUAUUUCCAAAAGAAACAAUGGCAGGCCUCUUUU
Kluyveromyces dobzhanskii
UGAAUU-------AGA-ACGUUUCUCG----UUCACCGCAAC-AGAGGA-UGGAGUCUGUGGUUCGUUUUCAUGA-UUCCAAAUGCAGCAGCCGUUGUUUCGUUUGGCA--GGUUUUUUU
Kluyveromyces aestuarii -----------------AUCUC---------------CUUAAU-GAGGA-AAAAAUAUGUGCCGAAAUCCC-UGA-UCUGGAAUUCCUUUAU-------------------GGCUCCUU-
<<<<<
1090
1100
1110
1120
1130
1140
1150 1160 1170 1180 1190 1200
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
CS4 CS4
structure
) ((((((((((( )))))))))))
Kluyveromyces lactis --UU--AGUGAUUUU--UCCAAACCCAUU--CUUCUCUUCGUUAGUAUAUUUGA-CUCGG----UUCUAUUUCAAUGUGUCGCAUUGAAAUAGAAAUCCUUUGUGCAU--AAAAUCAUU-
Kluyveromyces wickerhamii
--UU--AGUGAUUUU--UCCAGGCUAUUA-------UUGAACUGUUAACAGCAA------------CUAAUUU-------------UAAAUAAG-----UUGGUG-AU--AAAAUCAUU-
Kluyveromyces nonfermentans U--UAU--UGAUUUUUC--AAAGUCCGUC----UUUGUCUUUUUCUGCUUUGGA-----------UCCAUU-----------------AGUAUA------UCCUCC--GAAAAAUCA--U
Kluyveromyces marxianus ---UAU--UGAUUUU--UCCAGGCCUGUAACUUUAACUCUGCUGUUAUUUCCCGCCCCACCGACUUCUAUUAAGCUUUAAGGUUUAUAAA-AGUCACUUUUCGUGCAU--AAAAUCA--U
Kluyveromyces dobzhanskii
--UUAU--UGAUUUU--UCCAAACCUGUA--UUCCGUUAUGCUAGUGUGUUCGAAUUCGU----GUCUAUUUCAC------------AAAUAGAA-UCCUUCGUGCAU--AAAAUCA--U
Kluyveromyces aestuarii -UUUAU--UGAUUUUUC--AAAGCC--------UUUACCAUUUUUUGCUUUGGA-----------UCCCCCUC--------------AAGUAUA------UAGUUC--GAAAAAUCA--U
>>>>>
1210
1220
1230
1240
1250
1260
1270
1280
1290
1300
1310
1320
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
CS5a
structure
((((
Kluyveromyces lactis -UACCUCAAC--UUUCCUCACCAAGUAUCUUUGAAUGUUUUGCUCCCAUCAAGAAAGAAAGAACUUCUACUACAACGAGUCAAUAUGCUAGCAUGGUGGAGGUACAGUUGUCCUAUGGUG
Kluyveromyces wickerhamii
-UACCUCUAU-------UCAAC-----------------UUGGUUGAUCAUCGG-----UUCUUCCUUAUCG-----------UUUGUUAACAGGGGGUGCCUGUAAUG---UGGUAAUG
Kluyveromyces nonfermentans UCACCUCCUU------CUAAACU--------------------CUCUUCAUGCA-----UGUCUGUUUUA---------------UGAUCCAAAGCAGUUGGUACAAAUCUGAAGCAAUG
Kluyveromyces marxianus UUACCUCAAUCAUUUCCCCAACUCUUGGUAAAAUAGAAAAAAAACAAGCACCAGACAAUUUGAUCCAUAGGGAAAGGGAAACUUCUAUAACAACAGAGUAGGUACAGAGCACCUA-UGUG
Kluyveromyces dobzhanskii
UUACCUCAAC--UCCCUUCACCAUUUGUCUUU-------UCGUUGAAAUAUCCACACUCUAAAUCCUUGUUGAAAUGA-----UACACUGGCAUGACCGAGGCACAGUGACUCUAUGGUA
Kluyveromyces aestuarii UCACCUCCACA-----UUAAACU--------------------CUCUUUAUGCA-----CGCAUUCAUGGC--------------UGAUCCAUAGCUACAGGUACAAGUUGAGCACGGUG
1330 1340 1350 1360 1370 1380 1390 1400 1410 1420 1430 1440
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
|-CS5b -| |---CS5b---| |--Sc6--|
structure
(((((( (( )))) )))))) (((( ((
Kluyveromyces lactis GAUCUU-GAAAAGACCCUCGGAAGACAGGGCCACAUCGA-C--GAUCAAUAUUUUGCCUAUCAUCAACAA-AUUGUGACUGAUUUUUCUCUCAUCGUUUAUGAGAGAACCCUCUUGUCUC
Kluyveromyces wickerhamii
GAUCC-CAACAAGCCUUUCGGUU-AUAAGGCUGAAUU---CGGGUC-AAUAUUUUGCCAAUCAUUGGCA---UUUUUGUCAAUUGAA---------------------------------
Kluyveromyces nonfermentans GAUCUC-GCAAAGAU--UCGGAU--UCAAAUUCU-UUG--CAGAUU-AAUAUUUCGCUAUACACAGACAACAUGAU-GUGUGUUGAU---------------------------------
Kluyveromyces marxianus GGUCCUG-AAAAGACUUUCGAAAGAUUGAGUCG--U---AAAGGACUAAUAUUUUGCCUACCAUUAACAA-AAGACAAAAAAUCAAAGCAG-AUAGUU-----AAAUCUCUGUCAGAUUU
Kluyveromyces dobzhanskii
AAUCUC-GAAAAGAUCCUCGGAAGAUAGGUUCACAUCGA-CGAUU--AAUAUUUUGCCUAUCAUCACCAA-AUCGCUACUGGUGUCUCCAUCGAUGU-------AGAACCUUUUAGCC-C
Kluyveromyces aestuarii AAUCUC-GCAAAAAAA-UCGGAU--UCAAAUUUU-UU---AGAGAUUAAUAUUUCGCAAUCGAUGGACAAUAAAAAAGUCUGUCGUU---------------------------------
1450
1460
1470
1480
1490
1500
1510 1520 1530 1540 1550 1560
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
|-CS6--| CS5a Sm site
structure
)) )))) )))) |||||||||||||
Kluyveromyces lactis UUGUGAUUGAC--------GCUUAUAUGUCACCAUGGCAUGCAGCUGCUCCUUAUAACCACUUUUUGGUACAUCUCCCUACUAAGACUCGGGAACUGUUGCACUUAUCAAUUUAUUCUUG
Kluyveromyces wickerhamii
-------------------GCUUAUAUGUCAUUGCCAAUCAUUACGUUCCUUAU-AACUA-UUUUUGGACGUCUCU------------------------------------------
Kluyveromyces nonfermentans -------------------CCUUAUAUGUCAUUGUGUCUGGUUUGUUCCAUGU--AACCAUUUUUUGGAGUAUUGUUU----------------------------------------
Kluyveromyces marxianus UUUUCGUUUAUUAAUUGAUGCUUAUAUGUCAUUAUAGGCUUGCACUGUCCCUUAUAACCAUUUUUUGGCACCUAAACACACUUGCUUCAAGCAUCUUGAAUGUUUGAACAACCUUUUUUU
Kluyveromyces dobzhanskii
UUGUGAUUGAC--------GCUUAUAUGCUACGAUAGAAUGUUACUGCACCGAA-AACCAAUUUUUGGAUAUUCUCCUUUG-------------------------------------
Kluyveromyces aestuarii -------------------GCUUAUAUGUCAGCGUGCUUGAUUUGUUCCAUAU--AACCAUUUUUUGGAAAAUUAUU-----------------------------------------
|----CS7----|
1570
1580
1590
1600
1610
1620
1630
1640
1650
1660
1670
1680
....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|....|
structure
Kluyveromyces lactis AGAUAAGGGCAACA
Kluyveromyces wickerhamii
Kluyveromyces nonfermentans
Kluyveromyces marxianus CUUUUUAUUUUUCCUCUCUUGUUUGUAUAUUGUGUUUAACGUAUCCUUAGAGCCAUUGUGGCGCUUUGUCGGCUCGCCGUAGACACAUCUUAUCAAAUCACUUUUAUCAGCUCUUGAUGU
Kluyveromyces dobzhanskii
Kluyveromyces aestuarii
1690
....|....|....|..
structure
Kluyveromyces lactis
Kluyveromyces wickerhamii
Kluyveromyces nonfermentans
Kluyveromyces marxianus AGUCUGCUGUUAUCGAU
Kluyveromyces dobzhanskii
Kluyveromyces aestuarii