Selenoprotein evolution: SECIS

Introduction to selenocysteine insertion into selenoproteins

The SECIS element in 3' UTR of selenoproteins allows the insertion of selenocysteine for the appropriate TGA codon at time of translation. Auxillary gene products are needed for this to happen, including a selenocysteine-specific translation elongation factor (eEFSec), Sec-tRNA[Ser]Sec and a SECIS binding protein (SBP2). The SBP2 protein has three domains, including a conserved L7Ae RNA binding motif for SECIS RNA binding, a not completely overlapping ribosome binding site and a redox-sensitive cysteine-rich domain. The kink-turn tertiary structure in the SECIS is at the core of binding recognition. A mutation here leads to abnormal thyroid hormone function, very likely through inadequate production of DIO selenoproteins needed to interconvert thyroid hormones. SBP2 shuttles between the nucleus and the cytoplasm via nuclear localization and nuclear export signal motifs, the latter using the CRM1 pathway.

Surprisingly, despite many studies, a strong paralog of BP2 has never been mentioned. That protein, KIAA0256, also has 17 coding exons with near-identical intron locations and phases and full-length alignability, establishing KIAA0256 and SB2 as resulting from an ancestral gene duplication. The percent identity overall is only 42% but reaches 72% (human to human) in the 3 exons defining the L7Ae kink-turn binding motif. Since both genes can be recovered from chondrichthyes, lamprey, and amphioxus, the duplication event preceded the emergence of chordates. Becaue the roots of the selenoproteome and the insertion mechanism lie much deeper in pre-metazoa, it follows that the parent gene must have assumed the key SECIS recognition roles (as did both descendent genes, at least initially).

We could ask if KIAA0256 still plays a role in SECIS in contemporary organisms. One possibility is that some of the 30-odd selenoproteins use KIAA0256 in place of BP2. Another possibility is differential use during development or in various cell types. It's worth noting that KIAA0256 is considerably more conserved across its length than BP2, which has only a few exons conserved back to fish) suggesting that it is the latter that has the more minor role.

Reference sets of vertebrate SECIS BP2 and KIAA0256 L7Ae motif exons

>SECISBP2_homSap Homo sapiens (human) chr9 uc004aqj exons 14-16
1 YCSQMLSKEVDACVTDLLKELVRFQDRMYQKDPVKAKTKRRLVLGLREVLKHLKLKKLKCVIISPNCEKIQSK 1
2 GGLDDTLHTIIDYACEQNIPFVFALNRKALGRSLNKAVPVSVVGIFSYDGAQDQ 0
0 FHKMVELTVAARQAYKTMLENVQQELVGEPRPQAPPSLPTQGPSCPAEDGPPALKEKEEPHY 1

>KIAA0256_homSap Homo sapiens (human) uc001zxd chr15 exons 14-16 75% 69% 51%
1 YCNQVLCKEIDECVTLLLQELVSFQERIYQKDPVRAKARRRLVMGLREVTKHMKLNKIKCVIISPNCEKIQSK 1
2 GGLDEALYNVIAMAREQEIPFVFALGRKALGRCVNKLVPVSVVGIFNYFGAE 0
0 SLFNKLVELTEEARKAYKDMVAAMEQEQAEEALKNVKKVPHHMGHSRNPSAASAISFCSVISEPISEVNEKEY 1

>SECISBP2_ratNor Rattus norvegicus (rat) exons 14-16
1 YCSQMLSKEVDACVTGLLKELVRFQDRMYQKDPVKAKTKRRLVLGLREVLKHLKLRKLKCIIISPNCEKTQSK 1
2 GGLDDTLHTIIDCACEQNIPFVFALNRKALGRSLNKAVPVSIVGIFSYDGAQDQ 0
0 FHKMVELTMAARQAYKTMLETMRQEQAGEPGPQTPPSPPMQDPIQSTDEGTLASTGEEPHY 1

>SECISBP2_calMil Callorhinchus milii (elephantfish) AAVX01044988
1 YCSQVLSKDVDSCVTDLLKELVRFQDRLYQKDPIKAKKKRRIVMGLREVLKHLKLKRLKCIIISPNCEKIQSR 1
2 GGLDDALHNIISIACEQEIPFVFALNRKALGQCVNKPVPVSVLGIFSYDGAE 0
0 NQFHQMVEITEEARKAYQEMLDALQQELEADEEKGDSEEQPLISSESSTIHFNNVTSQPFSEADEPEY 1

>KIAA0256_calMil Callorhinchus milii (elephantfish) AAVX01105236
1 YCNQVLSKDIDECVTLLLQELVRFQERVYQKDPIKAKMKRRLVMGLREVTKHMKLRKIKCVIISPNCEKIQSK 1
2 GGLDEALYNVIAMAREQEIPFVFALGRKALGRCVNKLVPVSVVGIFNYFGAE 0
0 1

>KIAA0256_petMar Petromyzon marinus (lamprey)
1              LHKLRALIISPNCEKIQAK 0
2 GGLDEALQTVIALASEQSVPFVFALNRKALGHCLNKKVPVSVVGVFHYGGAE 0
0 THFQRLVALTEEARSAYRNMVSSLQRQEAAATSEPTGHTEDPLEASASPPSVPAHDPTALLHLLRPQQGPREDDPAEASGRSPGRNA 1

>SECISBP2_braFlo Branchiostoma floridae (amphioxus) extra exon
1 YCNQVLDKEIDATVTMLLQDLVRFQDRQYHK 00 DPIKAKAKRRIVMGLREVTKHLKLRKLKCIIIAPNLEKIQSK 1
2 GGLDDAIETILNLCMEQDVPFVFALGRKALGRAVNKLVPVSVVGVFNYDGAE 0
0 EHFKTMVELTTQARNAYIDMVTIYRQEWEQMQVGTDCSLV 1

Reference sets of vertebrate SECIS elements

SEPP2: 5 SECIS sequences

>SEPP2_SECIS_monDom Monodelphis domestica (opossum) chr2:9455773-9456962 COVE: 30 55 bp after stop codon
ACGTGCCGCTGCCCCTCCCTCCCTCCAAGAATGACGCCCACAGTGAAACCCAGAGAACTGGTCCCTGTGGGCTGATGCCCCAGAGGGGAGGAGAGGC

>SEPP2_SECIS_macEug Macropus eugenii (wallaby) SECIS COVE score: 26.38 trace ti|975557399 contains terminal exon
CCTCATTCCTCTTCTCCATGGCATCAGCCCACCATGGCCGGTTCCCCGGGTTTCACTGTGGGCATCATTAATTGAGGGAGGGAAGGGCAATGGGCAAG

>SEPP2_ornAna Ornithorhynchus anatinus (platypus)Ultra131:321101-348720 COVE: 34.28 no match to chicken
TCCCTCGACTCCCGCTCCCGCCTCGCACTCATGACGTCCACGGTGTCAACCGGCCCGCCGGGCACCGTGGACTGACGCCGGTCGAGGCGGAGGGGT

>SEPP2_ornAna Ornithorhynchus anatinus (platypus) Ultra131:321101-348720 COVE: 22.82 no match to chicken
GGAATCAGGAACCCAGTAACATGAGGTCATCTTCGGAAGCCTGTGCCTAGAGGACCAAGATAATGGAAAAAGTGACGGACAAGGGTGTGTAGCTGG

>SEPP2_tetNig Tetraodon nigroviridis (pufferfish) COVE: 17
GCTGGACCCAGGCTGCTGGTGGTCCCGTTGATGACGTCTGCGCTGGTAAACCTGCCTGCAGGAGCCTGTGGACCGACGTGTGTGGACCCACCGGCAG

SELI: 17 SECIS sequences

>SELI_SECIS_homSap Homo sapiens (human) COVE score: 21.61
UUUCACUGAA UGAAGUU UGUGCUUGA AUGAA GAGUGUAUCUUA AA CCCCCUUUUUUUGGA CAGGCUGCACUU GGAU AAAAUA GGCACCA CUGUGUUGAU

>SELI_SECIS_panTro Pan troglodytes (chimp) COVE score: 21.61
uuucacugaa ugaaguu ugugcuuga augaa gaguguaucuua aa cccccuuuuuuugga caggcugcacuu ggau aaaaua ggcacca cuguguugau

>SELI_SECIS_macMul Macaca mulatta (rhesus) COVE score: 21.25
uuucacugaa ugaaguu ugugcuuga augaa gaguguaucuua aa ccuccuuuuuuugga caggcugcacuu ggau aacaua ggcacca cuguguugau

>SELI_SECIS_otoGar Otolemur garnettii (bushbaby) COVE score: 21.25
uuucacuaag ugaaguu ugugcuuga augaa gaguguaucuu aa acccuuuuuuuuggac agguugcacuu ggau aaaaua ggcacca uuguguugau

>SELI_SECIS_musMus Mus musculus (mouse) COVE score: 24.62
uuccacugaa ugaaguu ugugcuuaa augaa gagugugucuu aa acccuuuuuuuuggac agguugcacuu ggau aaaaua ggcacca cuguguugau

>SELI_SECIS_ratNor Rattus norvegicus (rat) COVE score: 24.62
uuccacugaa ugaaguu ugugcuuaa augaa gagugugucuu aa acccuuuuuuuuggac agguugcacuu ggau aacaua ggcaccg cuguguugau

>SELI_SECIS_cavPor Cavia porcellus (guinea_pig) COVE score: 17.42
uuccagugaa ugaaguu ugugcuuga augaa gaguguaucuua aa cccuuuuuuuuuugga cagguugcacuu ggau aaaaua ggcacca cuguguugau

>SELI_SECIS_oryCun Oryctolagus cuniculus (rabbit) COVE score: 19.47
ucuacugaau gaaguuu gugcuuga augaa gaguguaucuua aa cccuuuuuuuuugga cagguugcacuu ggau aaaau aggcac cacuguugau

>SELI_SECIS_sorAra Sorex araneus (shrew) COVE score: COVE score: 8.13 loose
uuccacugaa ugaaguu ugugcuuga augaa gaguguaucuu aa acccuuuauuuuguuuuuuugga cggauuacacuu ggau auaaua ggcacca uucuguugau

>SELI_SECIS_eriEur Erinaceus europaeus (hedgehog) COVE score: 14.43 loose
cacggaauga aguaugu gcuuga augaa gaguguaucuu aa acccuuuuuuuuuuuugga cagguugcacuu ggau aua auaggc accacucugu

>SELI_SECIS_canFam Canis familiaris (dog) COVE score: 14.04
uuccacugaa ugaaguu ugugcucga augaa gaguguaucuua aa cccuuuuuuuuuugga uggauugcacuu ggau aaaaua ggcacca cuguguugau

>SELI_SECIS_equCab Equus caballus (horse) COVE score: 19.57
uuucacugag ugaaguu ugugcucga augaa gaguguauucuu aa acccuuguuuuugga uggauugcacuu ggau aaaaua agcacca cuguguugau

>SELI_SECIS_dasNov Dasypus novemcinctus (armadillo) COVE score: 17.76 loose
cuacugaauu aaguuug ugcuuga augaa gaguguaucuu aa acccuuuguuuuuuugga cagguugcacuu ggau aaaa uaggca ccacuauguu

>SELI_SECIS_loxAfr Loxodonta africana (elephant) COVE score: 25.81
uuccacugaa ugaaguu ugugcuuga augaa gaguguaucuu aa acccuuuuuuuuggac agguugcacuu ggau aaagua ggcacca cuguguugau

>SELI_SECIS_monDom Monodelphis domestica (opossum) COVE score: 26.46
uucuacugaa ugcaauu ugugcuuga augaa gagugugucuu aa auccuuuauauggac aggcugcacuu ggag aga auaagca caaccauguu

>SELI_SECIS_ornAna Ornithorhynchus anatinus (platypus) COVE score: 19.25
ucccagugaa ugaagcu ugugcuuga augaa gagugcaucuua aa cccauuuuuuuugga aaagcugcacuu ggag agaaag ggcacg acuguguuua

>SELI_SECIS_anoCar Anolis carolinensis (lizard) COVE score: 22.02
uccccuuugu guguguc cuuugugcgugu augaa gagugcggccuc aa cccaggcgucuugga agggccgcaccc ggaa gaa acggagc acagcaaaga

>SELI_SECIS_galGal Gallus gallus (chicken) COVE score: 24.62
uuuuauugag uuuauuu gugcuuaa augaa gagugcgcuuc aa acccagaccaggag agggcgcacuu ggag ugagc gagucaa accuugcucc

SELK: 19 SECIS sequences

>SELK_SECIS_homSap Homo sapiens (human) COVE score: 34.53
acaaggacug cucugug uccucacagauga augag gucaugcuggg aa uucccucugcaggga acuggccugac ugac augcaguuc cauaaa ugcagauguu

>SELK_SECIS_panTro Pan troglodytes (chimp) COVE score: 34.53 
acaaggacug cucugug uccucacagauga augag gucaugcuggg aa uucccucugcaggga acuggccugac ugac augcaguuc cauaaa ugcagauguu
>SELK_SECIS_macMul Macaca mulatta (rhesus) COVE score: 36.14 
acaaggacug cucugug uccucacagauga augag gucaugcuagg aa uucccucuacaggga acuggccugac ugac augcaguuc cauaaa ugcagauguu

>SELK_SECIS_otoGar Otolemur garnettii (bushbaby) COVE score: 31.99
caaggauugc ucugugu cuucacagauga augag gucaggcuggg aa uucucucuucaggga acuggccugac ugac augcagu ucuauaa acgcacuuuu
 
>SELK_SECIS_tupBel Tupaia belangeri (tree_shrew) COVE score: 36.65
acaaggauug cucugug uccccacagauga augag guuaugccggg aa uucccuccacaggga ucuggccugac ugau acgcaguuc uauaaa ugcacauguu

>SELK_SECIS_musMus Mus musculus (mouse) COVE score: 31.73
acaaggauug cucugug uccccacagauga augag gucaugcuggg aa uucccucugcagga ucuagccugac ugau acgcaguuc uauaaa uguacauguu
 
>SELK_SECIS_ratNor Rattus norvegicus (rat) COVE score: 33.02
acaaggacug cucugug uccucacagaaga augag gucaugcuggg aa cucccucugcagga ucuggccugac ugau gugcaguuc uauaaa uguacaugug
 
>SELK_SECIS_cavPor Cavia porcellus (guinea_pig) COVE score: 33.72
cucuguguuc ucacaga uaa augag gucacgccagg aa uucucucagcaggga ucuggcuugac ugau acgcagu ucucua aaugcauaug
 
>SELK_SECIS_oryCun Oryctolagus cuniculus (rabbit) COVE score: 38.80 
accaggauug cucugug uccccacagauga augau gucaggcuggg aa uucccuccacaggga ucuggccugau ugag augcaguuc uauaaa ugcguauguu

>SELK_SECIS_sorAra Sorex araneus (shrew) COVE score: 40.91
auaaggacug uucugug uccacacagauga augag gucaugcuggg aa uucccucuacgggga ucuggcaugac ugau augcag uucgaua aaugcacaug

>SELK_SECIS_eriEur Erinaceus europaeus (hedgehog) COVE score: 37.30
acaaggauug cucugug uucucacagguga augag guuaugcuggg aa uucccuccaugggga ucuggcaugac ugau augcaguuc uauaaa ugcacauguu

>SELK_SECIS_canFam Canis familiaris (dog) COVE score: 36.30 
acaaggauug cucugug ccuucacagacgg augag guugugcuagg aa uucccuccccaggga ucuggcaugac ugac augcaguuc uauaaa ugcacauguu

>SELK_SECIS_felCat Felis catus (cat) COVE score: 37.35
acgagaauug cucugug uccucacagacag augag gucgugcuggg aa uucccuccccaggga ucuggcaugac ugac augcaguuc uauaaa ugcacauguu

>SELK_SECIS_bosTau Bos taurus (cow) COVE score: 39.16 
cucugugucc ucacaga cga augag gucaugcuggg aa uucccuccgcaggga ucuggcaugac ugac augcagu ucuaua aaugcacguu

>SELK_SECIS_dasNov Dasypus novemcinctus (armadillo) COVE score: 29.00
acaaggauug cucugug uccucacagauga augag gucauguuuggg aa uucccucugcaggg aucuggcaugac ugac uugcaguuc cauaaa ugcacauguu

>SELK_SECIS_loxAfr Loxodonta africana (elephant) COVE score: 34.85 
acaaagacug cucugug uccccacagacgg augag gcugugcuggg aa uucccucugcaggga ucuggcauggc ugac augcaguuc cauaaa ugcacauguu

>SELK_SECIS_monDom Monodelphis domestica (opossum) COVE score: 21.75
aagaauugcu cugucua cacagauua augau guugugcuggg aa cucccaucuuacagga uccaguguaac ugau ugcaa uuguaua aaugcacaug

>SELK_SECIS_ornAna Ornithorhynchus anatinus (platypus) COVE score: 33.68
aauaauugug cugugaa caagcagauua augau guuuugcuggg aa uuccuucaggga uccaguauaac ugau aaagc aauuaua uaaaggcaca

>SELK_SECIS_anoCar Anolis carolinensis (lizard) COVE score: 33.87
aauucccugc ucugcca auuggcgggacc augau guuguccuggg aa uuccuuauucuggga uccagggcaac ugaa aagcaguuc uguuaa auuaaaugca

>SELK_SECIS_galGal Gallus gallus (chicken) COVE score: 30.90
ugagaacugu ucugcaa uauaagcagauga augaa guuguacuggg aa cuccuucaagga uccaguguaac ugaa gugcagug uuauuaa auacauguuu

SEPW: 13 SECIS sequences

>SEPW_SECIS_homSap Homo sapiens (human) COVE score: 36.01
GACCCAGCCC CUCUCAG CAGACGCUUC AUGAU AGGAAGGACUG AA AAGUCUUGUGGACACC UGGUCUUUCCC UGAU GUU CUCGUGG CUGCUGUUGG

>SEPW_SECIS_panTro Pan troglodytes (chimp) COVE score: 36.01
GACCCAGCCC CUCUCAG CAGACGCUUC AUGAU AGGAAGGACUG AA AAGUCUUGUGGACACC UGGUCUUUCCC UGAU GUU CUuGUGG CUGCUGUUGG

>SEPW_SECIS_ponPyg Pongo pygmaeus (orang_sumatran)C OVE score: 30.55
GACCCAGCCC CUCUCAG CAGACGCUUC AUGAU AGGAAGGACUG AA AAAUCUUGUGGACACC UGGUCUUUCCC UGAU GUU CUCGUGG CUGCUGUUGG

>SEPW_SECIS_macMul Macaca mulatta (rhesus) COVE score: 33.55
GACCCAGCCC CUCUCAG CAGACGCUUC AUGAU AGGAAGGACUG AA AAGUCUUGUGGACgCC UGGUCUUUCCC UGAU GUU CUCGUGG CUGCUGUUGG

>SEPW_SECIS_musMus Mus musculus (mouse) COVE score: 34.04
UUGGCCCAGC CCCUCGU GGCAGACGCUUC AUGAU GGGAAGAACUG AA AUGUCUCGUGGACGC CUGGUCUUUCC CUGAU GUCCCU GCGACUG CCACGUAGGG

>SEPW_SECIS_ ratNor Rattus norvegicus (rat) COVE score: 26.55
UGGCCGGCCU UUCUUGG CAGCCGCUUC AUGAC AGGAAGGACUG AA AUGUCUCAAAGACCUG UGGUCUUUCUU CGAU GUU CCUGCGG CCACCAAGUC

>SEPW_SECIS_oryCun Oryctolagus cuniculus (rabbit) no SECIS element found
AGTAACCTTGACCCAGCCCCTTTCATGCCTCAGCCTCGTCTCCATAGGCTAAGACTGGAGAAATGAGTCCCCTGAAGAACTGAAACTGGGGGTAGAGGGTTGGTGTTTTAAGATGTGGATGAGCTGGTCTTTAC

>SEPW_SECIS_canFam Canis familiaris (dog) COVE score: 34.04
UUGGCCCAGC CCCUCGU GGCAGACGCUUC AUGAU GGGAAGAACU GA AAUGUCUCGUGGACGCC UGGUCUUUCC CUGAU GUCCCU GCGACUG CCACGUAGGG

>SEPW_SECIS_susScr Sus scrofa (pig) no SECIS element found
AGTAACCTTGACCCAGCCCCTTTCATGCCTCAGCCTCGTCTCCATAGGCTAAGACTGGAGAAGTCTTGTGGACGCCTGGTCTTTCCCTGATGTTCTCGTGGCTGCTGTTGGGGGCAGAGATGGATGAGCTGGTCTTTAC

>SEPW_SECIS_borAnc Boreoeuthere ancestralis (ancestral) COVE score: 33.54
GGCCCAGCCC CUCUCAG CAGAUGCUUC AUGAC AGGAAGGACUG AA AUGUCUUGUGGACGCC UGGUCUUUCCC UGAU GUU CUUGUGG CUGCUGGUUG

>SEPW_SECIS_dasNov Dasypus novemcinctus (armadillo) COVE score: 28.71
CCCAGCUGCC CUUGGCA GACGCUUC AUGAG GGGAAGGACCU AA AUGCGUCGUGGAUGCC UGGUCUUUCCC UGAU GCUCCUUCAC CUGCCAG AUGGGGCAGA

>SEPW_SECIS_loxAfr Loxodonta africana (elephant) no SECIS element found
GGGACCTTGGCCCAGCCCCTTTCAGCAGACACTTCATGACAGGAGGACTGAAATGTCTCCCAGACGCCTGGCTCTTTCCCTGAATCTGTCGGCTGCAGGACAGGGCAGCGGTTGACTCTCTCGTTTTTTGCAT

>SEPW_SECIS_echTel Echinops telfairi (tenrec) no SECIS element found
AGGCCAGAGACCTTGGCCCAGTCCCTCCATGACAGGCAGAACTGAAATGTCCTCTGGACAAGTGGTCTTTTCCAGAAACCCCAGGGCTGCTGGGCCGGAGCCGAGGCTGACAACCCTGGTCTTTGCCT

DIO1: 29 SECIS sequences

>DIO1_SECIS_homSap Homo sapiens (human) COVE score: 29
ttttaactctgtgtctttacatatttgtttatgatggccacagcctaaagtacacacggctgtgacttgattcaaaagaaaatgttataag

>DIO1_SECIS_ponPyg Pongo pygmaeus (orang_sumatran) COVE score: 29
ttttaactctgtgtctttacatatttgtttatgatggccacagcctaaagtacacacggctgtgacttgattcaaaagaaaatgttataag

>DIO1_SECIS_macMul Macaca mulatta (rhesus) COVE score: 29
tttcaactctgtgtctttacatatttgtttatgatggccacagcctaaagtacacacggctgtgacttgattcaaaagaaaatgttataag

>DIO1_SECIS_cavPor Cavia porcellus (guinea_pig) COVE score: 24
tgttaactctgcttcttttcatatttgttcatgacggtcacagtctaaagtacacacagctgtgacctgatttgaaagaaaatgttttaag

>DIO1_SECIS_canFam Canis familiaris (dog) COVE score: -
ttttaactctgcttcttttcatgtttgtctatgacggccacagcctaaagcacacacagctgtgacttgatttgaaagaaaatgttttaag

>DIO1_SECIS_felCat Felis catus (cat) COVE score:  -
ttttaactctgcttcttttcacgtttgtctatgacggccacagtctaaagtgcacacagctgtgacttgacttgaacgaaaatgttttaag

>DIO1_SECIS_bosTau Bos taurus (cow) COVE score: 28
ttttaactctgcctcttttcatatttgttcatgacggccacagcctaaagtacacacggctgtgacttgatttgaaagaaaatgttttaag

>DIO1_SECIS_sorAra Sorex araneus (shrew) COVE score: 24
cggaaactcagcttctcttcatatttgtttatgacagccccagctgaaagtacacacagctgtggcttgattggaaagaaaatgttttaag

>DIO1_SECIS_eriEur Erinaceus europaeus (hedgehog) COVE score: 26
tttaactctgctttcttctcatatttgcttatgatggtcacagcttaaagtatacacagctgtgacttgattggaaagaaaatattttaag

>DIO1_SECIS_borAnc Boreoeuthere ancestralis (ancestral) COVE score: 29
ttttaactctgcttcttttcatatttgttcatgatggccacagcctaaagtacacacggctgtgacttgatttgaaagaaaatgttttaag

>DIO1_SECIS_dasNov Dasypus novemcinctus (armadillo) COVE score: -
ttttaactctgcttcttttcatatttgtttatgatggccacagtttaaagtacatacagctgtgacttgatatgaaaaagaaatattttaag

>DIO1_SECIS_loxAfr Loxodonta africana (elephant) COVE score: - 
ttttaactctgcttcttttttcatgtatttatgatgggccacagcctaaagtgcacaacagctgtgacttgatttgaaaaacatctttaag

>DIO1_SECIS_monDom Monodelphis domestica (opossum) COVE score: -
tttccatcctgcttctacaaatatttatttatgacaatcacagcctaaagctcagggcagctgggattcgacgggagaaaaagtttgtaag

>DIO1_SECIS_ornAna Ornithorhynchus anatinus (platypus) COVE score: 24
ccccggatccggttccgtgaatattggtttatgagggtcacagtgtaaagcgcatgcagctgtgacttgatctgagaaaatatttctgcggc
 
 
>DIO2_SECIS_homSap Homo sapiens (human) 5,630 bp utr COVE score: 30
cagagatgtgcagagttgaccagtgtgcggatgataactactgacgaaagagtcatcgactcagttagtggttggatgtagtcacattagtttgcctctc

>DIO2_SECIS_macMul Macaca mulatta (rhesus) COVE score: 30
cagagatgtgcagagttgaccagtgtgcggatgataactactgacgaaagagtcatcgactcagttagtggttggatgtagtcacattagtttgcctctc

>DIO2_SECIS_musMus Mus musculus (mouse) COVE score: 29
cggagatgttcagagctcactggtgtgcgaatgataactactgacgaaagagctgtctgctcagtctgtggttggatgtagtcacacgagtctgcctttctgca

>DIO2_SECIS_ratNor Rattus norvegicus (rat) COVE score: 28
ccgagatgttcggagctcactggtgtgcgaatgataactactgacgaaagagtcatctgctcagtctgtggttggatgtagtcacacgagtctgcctctccatc

>DIO2_SECIS_canFam Canis familiaris (dog) COVE score: 27
ctgggatgtgcagaggtgaccagtgtgcgaatgataactactgatgaaagagtcactgactcagttagtggttggatacagtcacattagttttcctct 

>DIO2_SECIS_borAnc Boreoeuthere ancestralis (ancestral) COVE score: 30
ctgggatgtgcagaggtgaccagtgtgcaaatgataactactgatgaaagagtcattgactcagttagtggttggatgtagtcacattagtttgcctctc

>DIO2_SECIS_dasNov Dasypus novemcinctus (armadillo) iodothyronine deiodinase type II
ctgggaagttcagaggctaccagtgtgccaatgataactactgacgaaagaggcatcgactcagttagtggttggatgtagccacattagtttgcctctc
 
 
>DIO3_SECIS_homSap Homo sapiens (human) COVE score: 31
ttgggtgcacaggagccccactgctgatgacgaactatctctaactggtcttgaccacgagctagttctgaattgcaggggcctcaaagcagca

>DIO3_SECIS_macMul Macaca mulatta (rhesus) COVE score: 30 
ttgggtgcacaggagccccactgctgatgacgaactgtctctaactggtcttgaccacgagctagttctgaattgcaggggcctcaaaacagca

>DIO3_SECIS_musMus Mus musculus (mouse) COVE score: 26  
ttgggtgcgctggagccctggctgctgatgacgaaccgcctctaactgggcttgaccacgggtcggctctgaattgcagagaggctcgaaacagc

>DIO3_SECIS_ratNor Rattus norvegicus (rat) COVE score: 26 
ttgggtgcgctggagccctggctgctgatgacgaaccgcctctaactgggcttgaccacgggtcggctctgaattgcagagaggctcgaaacagc

>DIO3_SECIS_canFam Canis familiaris (dog) COVE score: 26
ttgggtgctggcgagccccactgctgatgacgagccgcctctaactggtcttgaccacgagctggttctgagttgcaggggggcttgcagcggc

>DIO3_SECIS_bosTau Bos taurus (cow) COVE score: 30
ttgggtgctcacgagccccactgctgatgaagagctgtctctaactggcctcgaccacgagctggttctgatttgcaggaggctcgcagcagc

>DIO3_SECIS_borAnc Boreoeuthere ancestralis (ancestral) COVE score: 27
ttgggtgctcaggagccccactgctgatgacgaactgtctctaactggtcttgaccacgagctggttctgaattgcagggggctcgcagcagca

>DIO3_SECIS_loxAfr Loxodonta africana (elephant) COVE score: 22
ttcggtgcgctagagccccactgctgatgacgaactgtctctaactggtcttgaccacgagctgattccgaattgcagggaactcgcagcagc

>DIO3_SECIS_echTel Echinops telfairi (tenrec) COVE score: -
ttcggtgctctgcagccccactgctgatgacgaactgcctctcactggtcttgaccacgagctgcttctgaaatgcaggggactcgcagccgca