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Chapter 3 miRNA and Target Gene Prediction Using Tissue-Selective Motif

3.3.4 Predicted Novel miRNAs

Over 70% of the predicted frequent motifs do not match any seed regions of currently known human miRNAs. As these unmatched motifs may be potential target sites of unknown miRNAs, we check each of them to see whether it is located in any of the stem regions in the secondary structures of the miRNA candidates predicted by Pedersen et al. [104].

Figure 10. Three cases of frequent motifs that match predicted secondary structures of mature miRNA candidates. (A) AAUCUUU and UCCUUGU. (B) AACUUUU and CUGGGCA. (C) UAGGAUG

Because the locations of seed regions in the predicted secondary structures are unknown, we consider three possible binding situations. First, two different motifs match the two primes of the stem part of a miRNA gene candidate (Figure 10A).

Second, two motifs are mapped onto the same stem of a miRNA gene candidate but at two different locations (Figure 10B). The predicted mature sequences are two potential alternative forms of this candidate miRNA. Third, a motif matches a region of a known miRNA gene, but our predicted mature miRNA is different from the known mature sequence in miRBase. For example, in Figure 10C, the motif UAGGAUG matches a region in the mature sequence UUGCAUAUGUAGGAUGUCCCAU of hsa-miR-448, but our predicted mature sequence is ACAUCCUGCAUAGUGCUGCCAG; we call it an alternative mature

form of hsa-miR-448. Tables 2 and 3 show 4 and 11 additional examples, respectively.

When G:U pairing is not allowed, 60 frequent motifs that do not match any known miRNA seed regions give rise to 48 mature miRNA candidates (Table 2).

However, P-9-3p and -5p are two novel alternative mature forms of hsa-miR-652;

that is, these two sequences overlap with that of hsa-miR-652 but have different seed regions. Similarly, P-27-3p and -5p are two novel alternative mature forms of hsa-miR-802 (Table 2). When one G:U pairing is allowed in the seed match, 116 frequent motifs give rise to 93 mature miRNA candidates (Table 3). P-62, P-63-1 and -2, P-64-1 and -2, P-65, P-66, P-67, P-68-1 and -2, and P-69 are novel alternative mature forms of has-miR-544, hsa-miR-1264, has-miR-1298, hsa-miR-873, hsa-miR-376b, hsa-miR-381, hsa-miR-365, and hsa-miR-448, respectively (Table 3).

Table 2. Predicted novel miRNAs with no G:U pairing in the seed match. *P-9 (3p and 5p), P-11 (3p and 5p), and P-27 (3p and 5p) are alternative mature forms of hsa-miR-652, cfa-miR-1839, and hsa-miR-802, respectively; however, no human version of cfa-miR-1839 is found in miRbase. The coordinates and the located gene IDs of these predicted mature miRNAs are given in the last two columns.

Frequent

Predicted miRNA mature sequence Genomic coordinates (NCBI

36.1) Gene located

AUGCAAA artery/aorta,

uterus P-1 AUUUGCAUAAUGGAUGC chr2:205539433-205539449 ENSG00000116117

GACAGAC

AGCUGAA peripheral nerve P-3 UUUCAGCUCAUAAAA chr5:82856771-82856785 ENSG00000038427 UAAUUGG ovary P-4 UCCAAUUAAGUCUUUUAAAU chr8:83086984-83087003

UUCAAAG liver/hepato CCUUUGAAAAUAUAAAAUC chr8:35236713-35236732

UUUCAAA

cerebrum, intestine, liver/hepato, pancreas

P-5 CUUUGAAAAUAUAAAAUC chr8:35236714-35236732 ENSG00000156687

UCAAUUU kidney P-6-3p UAAAUUGAGGUGGAUCCUGU chr15:57977629-57977648 AAAUUGA lymphnode P-6-5p CUCAAUUUAUUCCUAGAAACA chr15:57977582-57977602

UAAAUUG placenta UCAAUUUAUUCCUAGAAACAG chr15:57977583-57977603 AAUUGAG kidney UCUCAAUUUAUUCCUAG chr15:57977581-57977597 GCAAUUU peripheral nerve P-7-3p AAAUUGCCAUAAAGUG chr9:80472606-80472621

AUGUUCA artery/aorta P-8 AUGAACAUCUGAUUAUU chr11:132066843-132066859 ENSG00000183715 CCAUUCA retina P-9-3p UUGAAUGGCGCCACUAGGGUU chrX:109185270-109185290

UUGUGCA lymphnode P-9-5p CUGCACAACCCUAGGAGAGGG chrX:109185228-109185248

ENSG00000157600 (hsa-miR-652) ACAAAUC cerebrum P-10 UGAUUUGUUCAAGAUGAUGA chr7:27256727-27256746

GGUCUUG lung P-11-3p UCAAGACCUACUUAUCUACC chr15:81221851-81221870 AUCUACC testis P-11-5p GGUAGAUAGAACAGGUCUUG chr15:81221817-81221836 AUGCAUU lymphnode P-12-3p AAAUGCAUGAAAUAGAU chr1:158032773-158032789 AUAAUGA cerebrum P-13-5p-1 UCAUUAUAAAAUGUGAUAAUGU chr15:51019585-51019606

AAUGUCA placenta P-13-3p GUGACAUUAUGACAUUACAUU chr15:51019643-51019663 CUAAUUA retina UUAAUUAGCAAAAAGGCU chr1:208493640-208493657

GCUAAUU thymus, testis,

UGUUAAU placenta P-15-3p AAUUAACAGAAUAUUAU chr5:146055768-146055784

AUUAACA

P-15-5p UUGUUAAUCAAAAAACUAU chr5:146055739-146055757 ENSG00000156475

UGAAUAU cerebrum, pineal P-16-5p UAUAUUCACAUUUAUUGGAU chr7:146609661-146609680 ENSG00000174469

gland, peripheral blood

UCAAAAC placenta, lung P-17 UGUUUUGAUAACAGUAAUGU chr8:75780484-75780503 UAAUUUC kidney UGAAAUUAUAUUACCAACA chr10:128235007-128235025

UUGUAGU lymphnode P-19-1 GACUACAACUCCCAAGGUA chr1:153246074-153246092

GGAGUUG vein P-19-2 ACAACUCCCAAGGUACAUACA chr1:153246078-153246098 ENSG00000160685

AUUAUCU lymphnode P-20-1 UAGAUAAUUUGCACAUUAU chr14:72223490-72223508 ENSG00000205683

AUUACAG

P-21 CUGUAAUAUAAAUUUAAUUUAUU chr4:126647864-126647886

UUUUAAC cerebrum,

prostate, lung P-23 GUAUAUUGUGACAUACAUGU chr1:1463389-1463408

AAUUAAC placenta P-24-3p GUGUUAAUUAAACCUCUAUUUAC chr8:113724958-113724980 AUUUACA placenta AUGUAAAUACAGAUUUAAUUAAC chr8:113724904-113724926

UAUUUAC cerebrum,

placenta, kidney UGUAAAUACAGAUUUAAUUAACA chr8:113724905-113724927

UUUACAU

UCAUGUU placenta P-26 CCAACAUGAUGCUAAUAAAU chr17:73293262-73293281

AAUCUUU

UCCUUGU ovary, breast P27-3p AACAAGGAGAAUCUUUGUCACU chr21:36014934-36014955

ENSG00000159216 (hsa-miR-802)

CUAAUAA lymphnode,

placenta P-28 UUUAUUAGUGCCAUAUAAUA chr2:219696897-219696916 ENSG00000187736 AUCAAUA cerebrum, testis P-29-3p UUAUUGAUCAGCGUAGCAAACA chr5:3508678-3508699

UCAAUAA prostate, testis,

adrenal gland CUUAUUGAUCAGCGUAGCAA chr5:3508677-3508697 UUGAUCA lymphnode P-29-5p-1 CUGAUCAAUAAUAAGAUUGAU chr5:3508628-3508648

UUAUUGA lymphnode,

kidney P-29-5p-2 AUCAAUAAUAAGAUUGAUAC chr5:3508631-3508650

UAAACUG lymphnode CCAGUUUAUUUUGUAAAUAUA chr1:60826128-60826148 AUAAACU pancreas P-30

CAGUUUAUUUUGUAAAUAUA chr1:60826129-60826148

AAAUAUG

cerebrum, lymphnode, pancreas

P-31 UCAUAUUUUCUAUCUCUUUGCUU chr4:39795271-39795293 ENSG00000078177

UUCCUUA placenta P-32-1 UUAAGGAAAUUAUGCUGAAC chr4:21498378-21498397 ENSG00000185774 AAUUUUC lymphnode P-33 AGAAAAUUAGGUUGAUA chr6:37530191-37530212 ENSG00000137200

UUCAAAA cerebrum P-34 CUUUUUGAGUUUUGAGGAAG chr9:99359944-99359963 ENSG00000136842 UUACAGC placenta P-35 AGCUGUAAACAGCUCUCCA chr17:69079331-69079349

GAAUAAU

cerebrum, eye, lymphnode, ovary, lung, kidney

P-36 AUUAUUCUUUUUAUAAAA chr2:144691036-144691053 ENSG00000121964

UAUAAAC liver/hepato,

pancreas P-37 UGUUUAUAGUAAUGGGAGAUA chr9:127702054-127702074 ENSG00000167081

CAAAUAU peripheral nerve,

placenta, kidney P-38 AAUAUUUGGAAACAUCCA chr9:13932445-13932467

Table 3. Predicted novel miRNAs with 1 G:U pairing in seed match.

Predicted miRNA mature sequence Genomic coordinates (NCBI

36.1) Gene located

UCCAUCU retina P-6-3p GAGGUGGAUCCUGUUCCAAUU chr15:57977635-57977655 AAUUUGU lymphnode P-7-3p UGCAAAUUGCCAUAAAGUG chr9:80472603-80472621 UAAAGUG prostate, testis P-7-5p CAUUUUAUGGCAAUUUGUU chr9:80472560-80472583

UGAAAUA

P-13-5p-2 AAAUGUGAUAAUGUCAUUGC chr15:51019593-51019612

UUAACAG placenta,

pancreas P-15-5p CUUGUUAAUCAAAAAACUAU chr5:146055738-146055757 ENSG00000156475 UGGAUAU cerebrum UAUAUUCAUGAAUAUAU chr7:146609695-146609711

GAUAUAU

UGAUAUA kidney P-18-2 UUAUAUUACCAACAGAAAU chr10:128235012-128235030

AGAUUAU lymphnode P-20-2 GAUAAUUUGCACAUUAU chr14:72223492-72223508 ENSG00000205683

UGAUUUC bone, lung P-32-2 GGAAAUUAUGCUGAACUCAUUU chr4:21498382-21498403 ENSG00000185774

UAAUGUU artery/aorta P-39 UGACAUUAGUUCAUUU chr2:56002041-56002056 ENSG00000115380

UUUUAAG

lymphnode, skin, placenta, lung, pancreas

P-40 GCUUAGAAAAGUGACCUAGA chr2:77350845-77350864 ENSG00000176204

GCUGUUU lung P-41 AAAGCAGCGUGAAGAUGC chr2:105075146-105075163 ENSG00000135972

UGCAUAU

peripheral nerve, placenta,

pancreas

P-42 UAUAUGUAGAUGUAGCUAUAU chr2:192552553-192552573 ENSG00000144339

AGAUUAU lymphnode P-43 AUAAUUUAUUAGAACAAUUAG chr3:60801997-60802017 ENSG00000189283 AGUAAUU lymphnode UAAUUAUUUUUCUCCAUC chr3:62045270-62045287

GUAAUUA lymphnode, kidney, pancreas

P-44

UUAAUUAUUUUUCUCCAUC chr3:62045269-62045287 ENSG00000144724

UUCAAUU lymphnode P-45 UAAUUGGAAAUUUCAUUU chr4:146917406-146917423 ENSG00000151612 GCACAUU lung P-46 UAAUGUGUAAUGCUGUAGUUU chr4:153150954-153150974

GUCUAUA lymphnode, lung P-47 AUGUAGACAAAACAUCCAGAUAA chr5:15476433-15476455

UGUCUAU lung UGUAGACAAAACAUCCAGAU chr5:15476434-15476453

P-48 AUUAUUUUAGUAAUUCAACAG chr5:36750623-36750643

AAUAACU

placenta UAACUUUUAAUGUAAGCCUGG chr6:50426012-50426032

UAAGAGU placenta, pancreas

P-50

AACUUUUAAUGUAAGCC chr6:50426013-50426029

GCAAAAU cerebrum, retina,

placenta, prostate P-51 UGUUUUGCCAGCAUGUGGUUG chr9:72221811-72221831 UAAGUGA cerebrum UUCAUUUAAAAUUAGGC chr10:13522803-13522819

UUAAGUG lymphnode,

placenta, lung UCAUUUAAAAUUAGGC chr10:13522804-13522819

UUUAAGU lymphnode,

placenta, kidney UCAUUUAAAAUUAGGC chr10:13522803-13522819 AAGUGAA cerebrum

P-52

AUUCAUUUAAAAUUAGGC chr10:13522801-13522819

ENSG00000165626

UUAAGAU placenta, lung,

pancreas P-53 CAUCUUGAAAUAAGUCCUCA chr11:122110545-122110564 ENSG00000154127

UUUAAGA

lymphnode, uterus, prostate, testis, lung, kidney, pancreas

AUCUUGAAAUAAGUCCUCAU chr11:122110546-122110565

UUUUAAG

lymphnode, skin, placenta, lung, pancreas

UCUUGAAAUAAGUCCUCAUC chr11:122110547-122110566

UUGAUUA lymphnode, lung,

pancreas P-54 AUAAUCAGAAACACUAAUCA chr13:66389595-66389614 ENSG00000184226

UAAAUGU lymphnode,

placenta P-55 UAUAUUUAACAUACACUUG chr13:104638844-104638862

AUUAGUU lymphnode,

kidney P-56 GAAUUAAUGGUAUUAA chr14:56071430-56071448

UAGUUUA placenta P-57 UAAAUUAAAAUCAAUAUUUU chr16:7601200-7601219 ENSG00000078328 GAAUUUC skin, placenta GGGAAUUCCCACUCUGCAG chr17:9233266-9233289

GGAAUUU kidney P-58

GGAAUUCCCACUCUGCA chr17:9233267-9233288 ENSG00000170310 AUUCUGA kidney P-59 UUUAGAAUUCUAAUUA chr18:26697252-26697267

GCAAAUU lymphnode P-60 UGAUUUGCAUUUUAGU chr18:40000232-40000247 AUGAAGU uterus, lung P-61 UAUUUCAUUUUAAUCUUGA chr19:35550032-35550050 UAGCAAG prostate CUUGUUAAAAAGCAGAUUCU chr14:100584767-100584786

UUUUAGC lung P-62

UGUUAAAAAGCAGAUUCUGA chr14:100584769-100584788 hsa-miR-544

UUGUUGA kidney P-63-1 CUCAAUAAGUAUUUGUUGA chrX:113793391-113793409 ENSG00000147246

UCAAUAA prostate, adrenal

gland P-63-2 UUUGUUGAAAGAAUAAAUAA chrX:113793402-113793421 (hsa-miR-1264)

AACUUUU

cerebrum, lymphnode, prostate, intestine, lung, kidney

P-64-1 AAGAGUUCAUUCGGCUGUCCAG chrX:113855918-113855939

CUGGGCA thymus P-64-2 CUGUCCAGAUGUAUCCAAGU chrX:113855932-113855951

ENSG00000147246 (hsa-miR-1298)

CUCCUGU vein AAUAGGAGACUCACAAGUUCCUG chr9:28878920-28878942 UCCUGUU eye CAAUAGGAGACUCACAAGUU chr9:28878919-28878938 UCUCCUG breast

P-65

AUAGGAGACUCACAAGUUCCUG chr9:28878921-28878942

hsa-miR-873

CAUGUUU lymphnode,

placenta P-66 AAAACGUGGAUAUUCCUUCUAUG chr14:100576545-100576567 hsa-miR-376b GUAUCAA pancreas UUUGGUACUUAAAGCGAGG chr14:100582005-100582023

UAUCAAA testis P-67

GUUUGGUACUUAAAGCGAGG chr14:100582004-100582023 hsa-miR-381 CCUUAUU lymphnode P-68-1 AAAUGAGGGACUUUUGGGGGCA chr16:14310653-14310674

CCUAAAA uterus P-68-2 CUUUUGGGGGCAGAUGUG chr16:14310663-14310680 hsa-miR-365

UAGGAUG bone P-69 ACAUCCUGCAUAGUGCUGCCAG chrX:113964286-113964307 ENSG00000147246 (hsa-miR-448)

3.4 Experimental validation of predicted novel miRNAs and their

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