Gene Details:

Functional Descriptions:

  • Pb1 encodes a coiled-coil-nucleotide-binding site-leucine-rich repeat (CC-NB-LRR) protein and confers durable, broad-spectrum resistance to Magnaporthe oryzae races.
  • Panicle blast 1 (Pb1) is a blast-resistance gene derived from the indica cultivar ‘Modan’.
  • Pb1-mediated resistance, which is characterized by durability of resistance and adult/panicle blast resistance, has been introduced into elite varieties for commercial cultivation.
  • We discuss possible determinants for the durability of Pb1-mediated blast resistance.
  • Durable panicle blast-resistance gene Pb1 encodes an atypical CC-NBS-LRR protein and was generated by acquiring a promoter through local genome duplication.
  • Panicle blast 1 (Pb1) is a panicle blast resistance gene derived from the indica rice cultivar Modan.
  • Here, we investigated the molecular mechanisms underlying Pb1-mediated blast resistance.
  • Pb1-mediated panicle blast resistance was largely compromised when WRKY45 was knocked down in a Pb1-containing rice cultivar.
  • Leaf-blast resistance by Pb1 overexpression (Pb1-ox) was also compromised in WRKY45 knockdown/Pb1-ox rice.
  • Blast infection induced higher accumulation of WRKY45 in Pb1-ox than in control Nipponbare rice.
  • Overexpression of Pb1-Quad, a coiled-coil domain mutant that had weak interaction with WRKY45, resulted in significantly weaker blast resistance than that of wild-type Pb1.
  • Overexpression of Pb1 with a nuclear export sequence failed to confer blast resistance to rice.
  • These results suggest that the blast resistance of Pb1 depends on its interaction with WRKY45 in the nucleus.
  • In a transient system using rice protoplasts, coexpression of Pb1 enhanced WRKY45 accumulation and increased WRKY45-dependent transactivation activity, suggesting that protection of WRKY45 from ubiquitin proteasome system degradation is possibly involved in Pb1-dependent blast resistance.
  • Blast resistance of CC-NB-LRR protein Pb1 is mediated by WRKY45 through protein-protein interaction.
  • Based on the cosegregation of the resistant genotypes of Pb1 and Stvb-i, a rice stripe virus (RSV) resistance gene derived from an Indica cultivar Modan, we examined the linkage relationships between Pb1 and 13 Stvb-i-linked RFLP markers located on the long arm of chromosome 11.
  • Pb1 transcript levels increased during the development of Pb1+ cultivars; this expression pattern accounts for their adult/panicle resistance.
  • The Pb1 protein interacted with WRKY45, a transcription factor involved in induced resistance via the salicylic acid signaling pathway that is regulated by the ubiquitin proteasome system.
  • We precisely mapped the Pb1 locus for the panicle blast resistence of a quantitative nature on rice chromosome 11 using RFLP markers.
  • Identification of a RFLP Marker Tightly Linked to the Panicle Blast Resistance Gene, Pb1, in Rice.
  • Pb1 is classified as a quantitative resistance gene and it does not have fungal specificity.
  • Mapping of QTL related to Chr7 confirmed the existence of factors that were required for the resistance of Pb1 in the 22 to 23 Mbp region of the rice genome.
  • We clarified how the K209 cultivar is vulnerable to the blast disease despite possessing Pb1 and found the DNA marker responsible for the quantitative resistance of Pb1.
  • We found that the expression of WRKY45 gene downstream of Pb1 was weakly induced by rice blast inoculation at the full heading stage in K209.

Literature:

Gene Resources:

  • NCBI ID:
  • UniProt accessions:

Sequences:

cDNA Sequence
  • >LOC_Os11g37050.1
    GGGAGCCATCAGCGGCCATGACCGAGCTCGCGGCTGGCGCCGTGAGCTCACTGCTGGTGGTCATCCGCAACGAGGCTGTGCTGCTTGGCGGCGTCCGGGACGATGTGCAGTTCATCAAGGAGGAGATGGAGAGCATGAACAGCTTCCTAGGGCACCTAGCCAGGAGCGCGCCGCAGGGTGGAGAGCACGACGAGCAGGTGCGCACCTGGATGAACCAGGTGCGCCTGCTCGCGCAGGACTGCAACAACTGCATCGACCTCTACCTCTACAGCGGGAACCCGGAGATCCACCGTGCCAAAGGTAGACTCCGGCGACACCTCTGGTGGGTCTACTGGTCCCTGCGCAAGATGGTCGCGCAGCACCGAGCGGCCATCCAGCTCCGCCAGCTCAAGGACCGGGCGCGGGACGTCGGCGAGCGACGACTGAGGTATGGCGTTGAGATCCCAGCCACGACGAAGGCGGCCGCACCTGATGCAACAGGTGGCTATGCTGCTGGGGACGATGAAGAAGAAGATGAAGATGACCGCGAAGGACAATTTGCTGTGGCAACACCAACGTTGGCCCATCATTCTGCTCGATGGCCCGTCTTTGAGCCCCCTTCTCTGGACGACTACGTGAAGGCCAAGTTATTGGAGTGGATTGAGGGAGTTCCTGGAAACGCCATCGTGACGTTGTCCATCGCCATTGTAGCACCAGATGCAGATAATAAGGAAGTCCTTGCTATTGCACATGAAACTTTGGTTGCTCCAGACTACTACTACCGTCGCAGTATCATGGTCAACGTCCCGGCGGTGCACCTTGATTTTTTACCGCTTCGACCCAAGGAGGTCCTCTACTACATCTTGCGGGAGCTCGAGCGGGAAGAAGCGGCCGGATCGCAGAAGCAGCCCACAGATCAAGGTGAATGGGAGGAGGAGGATCCTGACCCTTGGCAAGATTACTACAAAAAATGTGGCATTTACCGTAGCAAAAAAGGTGTGCTCGATAAAATCAAAAGGAATATAAAAAAGATGAATATTTACGAAAAGCTTGATAAGATCAAGAGCGACATTCGAGAAGGACAACACAAGAGCGGCAAGCTGTTGCTGCTTAAGCTACAAGAGAAAGGTGCGGATCAGGTGGACCTACATGTACTCCTTCAGCTGTTGGTGCTCCAGTCTCAGCAAGACCAAGCGAAGAACAAAGCAGTAGACACCCATAAATTACCAGAGTGGAACGACAACCTCATAGAAAAATTAGCCATGAGGCTGAAGGATCATATGGAAGCAGATGAAAAGACCAAGAAGCTTAATGAGCAAACTGGAGTAGAAGAAGAAACAGCAGTCAGACAAGGCGGAGGAGGAGAGAGGAAGGAGGATGAGAAAGACGAAAGAGGAGACGGAGAAGAAGAGGGGAAGGAGGAGAGGAGGGACATGGAGAAAGGAGAAGAGAGGAAGGAGCAGCAGCAGGAGGAGCAGGAGAAAGAAGGGAGGAAGGAGGAACAGAACGAGGTAAGAAAGGAGACAGAAGGAAGAAAAGAACAAGTAGCAGGAGAGGAGGAAGAAAAAGAAGACCATGATGCTGATAATGATGAAGACAGCAATGATGATGATGATGATGAGGAGGAGGAGGAGGAGGACGACGATGATGAAGAAGAACCAATTCATCTTCATGAAGATCAATATGAACAAATCCTACGAGAAGTGTTCACAAAGAACGCCAGCAGCAAGGCCCAGGAGCAAGATAAATTGGTGGCCGAGCAAGCTACAAAGACTGCAGCCACTACATTGGATGAGGAACGAATCAAACAAATGATCAACGAAGCAAAACAAGATGTCTTAAGGGAGCTGCGGGGACGCGAAACTGATAAGAATCAAGCAACAGGTGAACCTGATGTTCCACCTGATAAGAATCAAGCAACAGGTCAACATGCAGTTGTTCTGGATCAAAACGAAGAGGCTTATTTTGAAGAAGTAGAGCAGAAGATTGAAGAAATCAAGCAGGAACTTAAAGAGCAGCTGAAGATCAAATGGATCGTGGACAAGATTAAACATCACTTACAGGATCAGTGCCCCCTCATTATCCTAAAATTTGATCAGATGATGGATGGATCCCGATGGGAGGAGATCAGAAAGGCTCTGAGCCTGTTAGAATTGAGTGCAGATGCATTGATCTTCACCACAGGGAGCACAGAACAGGCTAAAGGATATTGCTATCCACCACGAGAACCTATAGATCACTGTTCTCTTGTTGGTCTCTACTATTATACGGTGCTCAAGCTTACTAGCAAGCACAAGAATGAAGACAACGATAACGCACAGATTTTTCGTGGCATCTTGGAGGAGTGTGAGGGGCATGAATTCTGCATGAAGATCTTCACTCATGCTGTGTATGCTAATCCCAAGAGGAGCAATGAAGAGTTAAGGAAGCTACACAGCACCCTGCAGTCTCCAAAAAAATCATTCGACACCATAGCTAAGAAGATGTTCATGTACTCTTACAATGATCTACCTAAAGAATACAAGTCATGCTTGCTGTACCTAGCTATCTTCCCCAAGGGACAGAAGATTAGGCGGTCAACCTTGATTGCAAGGTGGGTTGCAGAAGGGTTGACATTCAAGGAAGATTGGCCCAGCTCTGTGTATCAGGCAAATCGATGTTTTGATGCCCTCATCCGTCGGTGGCTTGTTTATCCTGATGATATTAGTGCCACAGGAAAGATCAAGAGCTGTGTGGTAGGCGATCCAGTTCATGGGTTCATTACCGCAATTGCCAGAAAACAACATATTGTGGAGACACGCCTATCACATCACTTGGCTCGCCACTTCTCCATTTTCAATGATCTTCGACTCCGCAGCTCTGATAGAATTGGCACGTTCTTCCAAGGCCTCTCACGATCATCTCGAGTATCCCTACTCAAGGTGCTAGATCTAGAAGGTTGTCAGTGCTTTGCTAGTAAGAATCAGCGGTACCTCAAGGACATCTGCAACAAGATGTTACTGCTCAAATATCTGAGCCTAAAGGGAACAGATATTACCCAGCTGCCCAGTGAAATCAACTGCCTCCGCGAGCTAGAGGTATTGGATATCCGAGAAACCAAGGTGCCTGCAAATGCAACAGTAAATGTCCTGCTCTTGAAGCTGAAGCGTCTACTTGCTGGTGCTAGTCAGATTGATCCAACTCCAAGAAATTTTGTCACTAACGTCCGGATTCCTTCCAGGATAGACAAGATGATAAACATAGAGGTACTATCTAATGTCAAGGCCCAGCACCATGATAATTTGGAAGATATTGGGAAGCTATGCCAGTTGAGGAAGCTAGGTGTGGTTATTGATGGTAAGAAAAGTCACCTTGGGAGTTTGCTTAAAGCGATCAGTGACCTACATGCAAGCCTCCGTTCTCTGTCAATCACTATTCCCACAACCACACTCGAGGTTACTCCTTCAAGCCCAGAGTTACAAGATATTGCATCTCGCCTAAAACACCATCCTGAGTTTCTTGAGAGTCTAAGCATCAGTGGAGCCAAGCATCTTTTTCCATTGTTGACCGAAGGTGGTAATAAGAAACTTGCCAAGGTAACTCTAAGCAACACCCCACTGAACCAAGATGATCTGAAGTTCTTTGCCCAGCTGCCCATGTTACAGTGTGTTAGGCTCCGACACATTTCATGCACCGAGAGTGTGCTCAACTTCAAGAAAGATGATTTCAAATGCCTCAAGTACCTTCTTATTGAGGGCTCAAACTTGACTAATATTACTTTTGAGGATGAGGCAGCCTGTGAGCTCGAGAAGATGGTTTTATCTTCCACTTGCATAGAGTCTATTTCTGGAGTTCATGGGCTTCCAAAATTCGAAGAGCTTGAGTTGAACAGCAGCAGCTGCGGAAGGTTGCTATCATCATGTTTTTACAATGTTGAACGAATAGCCAAGCTGACTCTTCGTGGTACATTGCTGAAGCAAGGTGATCTACGAATCATCGCCAGGGAACTAAATATATGCTGCCTAGTGCTCTTGGAAAACTCTTTTGACATAAGCCAGAACCAGATTACCTTCGAAAAAGAAGAGTTCATATGGCTCAAACTTCTTAGTGTTGATTGCTCTACCATCACCAAGATTAACTTCATCACTGGATCAGCTCCTAGGCTCAAGAAGATCGTCTGGTCATCTTTCACCTCTCTCTCTGGCATCAACAACCTTCCTAGGTTGAAGGAGCTCGAGTTCAATGGATACTCAGTCCCAAATGATGTGGAAGAAGCCATTAAAAACAATAAAAGCATAAATCTTAAACATAATAAACCATAAGCTCAAGACAAAGCAAAATGGAGATGAATAAGAGGATGAAGACAATGCTGGGACGTTCCCATTCCGCTGGAAGAAACAGGTTTGTCACCAGAGGAGCATCTCCTTTTATCCCATGGTGTGGTTTGTGTGTGCTAGTGTGGATTGACATTCACTACTTGTCTGTTTAGTTCTTTCTGCTGTGCGTGAGCGTGTTTTTATGTTCTGCATGATGGAATAATTCTTGTTATCCCTTGTTTTTCCATCCTGCTCTCTGTTTCCTCCTCACTCTGTTTTGCTCTGTTTCCTTTGGTTCGAGTCAGTGTAAGCTCGAGGCTTGCTTGTTCAAGCCGGAGCGAATCCTTAACTAGTTGGCTTGACTTGCACCGGCGTGTGGGATGGCACACGATGCAGCGGGCATGGAGGGCAAGTTAGTCATGTCATTGGCACGGCATGCCCCGACGTGCGGGATGGCGCACAAAGAGGACGGGCGTGATGCGTGCAGTAGCATCAAAACCACTATTTTGTTTGTATTCAAGTTTCAATACAATCCAGAAAATGCTGCCACTTGACAGCGCCAAAAACTCTCTTCCTGTTCGGGTTCTTCCTCATCAAGTTCATAGCGACTTCGTTTGTTTGGTTTGTTTCTGTCACTGTGATTGTTTCAACCGAGTGTCTGGTTACTTGCAATGCATGAGTGCCTTGGGCCTCTGATTGAGCTAAAATGGTGGTGTCTATGCCCGAATTCGAATTTGTGTGGTTCTCAT
CDS Sequence
  • >LOC_Os11g37050.1
    ATGACCGAGCTCGCGGCTGGCGCCGTGAGCTCACTGCTGGTGGTCATCCGCAACGAGGCTGTGCTGCTTGGCGGCGTCCGGGACGATGTGCAGTTCATCAAGGAGGAGATGGAGAGCATGAACAGCTTCCTAGGGCACCTAGCCAGGAGCGCGCCGCAGGGTGGAGAGCACGACGAGCAGGTGCGCACCTGGATGAACCAGGTGCGCCTGCTCGCGCAGGACTGCAACAACTGCATCGACCTCTACCTCTACAGCGGGAACCCGGAGATCCACCGTGCCAAAGGTAGACTCCGGCGACACCTCTGGTGGGTCTACTGGTCCCTGCGCAAGATGGTCGCGCAGCACCGAGCGGCCATCCAGCTCCGCCAGCTCAAGGACCGGGCGCGGGACGTCGGCGAGCGACGACTGAGGTATGGCGTTGAGATCCCAGCCACGACGAAGGCGGCCGCACCTGATGCAACAGGTGGCTATGCTGCTGGGGACGATGAAGAAGAAGATGAAGATGACCGCGAAGGACAATTTGCTGTGGCAACACCAACGTTGGCCCATCATTCTGCTCGATGGCCCGTCTTTGAGCCCCCTTCTCTGGACGACTACGTGAAGGCCAAGTTATTGGAGTGGATTGAGGGAGTTCCTGGAAACGCCATCGTGACGTTGTCCATCGCCATTGTAGCACCAGATGCAGATAATAAGGAAGTCCTTGCTATTGCACATGAAACTTTGGTTGCTCCAGACTACTACTACCGTCGCAGTATCATGGTCAACGTCCCGGCGGTGCACCTTGATTTTTTACCGCTTCGACCCAAGGAGGTCCTCTACTACATCTTGCGGGAGCTCGAGCGGGAAGAAGCGGCCGGATCGCAGAAGCAGCCCACAGATCAAGGTGAATGGGAGGAGGAGGATCCTGACCCTTGGCAAGATTACTACAAAAAATGTGGCATTTACCGTAGCAAAAAAGGTGTGCTCGATAAAATCAAAAGGAATATAAAAAAGATGAATATTTACGAAAAGCTTGATAAGATCAAGAGCGACATTCGAGAAGGACAACACAAGAGCGGCAAGCTGTTGCTGCTTAAGCTACAAGAGAAAGGTGCGGATCAGGTGGACCTACATGTACTCCTTCAGCTGTTGGTGCTCCAGTCTCAGCAAGACCAAGCGAAGAACAAAGCAGTAGACACCCATAAATTACCAGAGTGGAACGACAACCTCATAGAAAAATTAGCCATGAGGCTGAAGGATCATATGGAAGCAGATGAAAAGACCAAGAAGCTTAATGAGCAAACTGGAGTAGAAGAAGAAACAGCAGTCAGACAAGGCGGAGGAGGAGAGAGGAAGGAGGATGAGAAAGACGAAAGAGGAGACGGAGAAGAAGAGGGGAAGGAGGAGAGGAGGGACATGGAGAAAGGAGAAGAGAGGAAGGAGCAGCAGCAGGAGGAGCAGGAGAAAGAAGGGAGGAAGGAGGAACAGAACGAGGTAAGAAAGGAGACAGAAGGAAGAAAAGAACAAGTAGCAGGAGAGGAGGAAGAAAAAGAAGACCATGATGCTGATAATGATGAAGACAGCAATGATGATGATGATGATGAGGAGGAGGAGGAGGAGGACGACGATGATGAAGAAGAACCAATTCATCTTCATGAAGATCAATATGAACAAATCCTACGAGAAGTGTTCACAAAGAACGCCAGCAGCAAGGCCCAGGAGCAAGATAAATTGGTGGCCGAGCAAGCTACAAAGACTGCAGCCACTACATTGGATGAGGAACGAATCAAACAAATGATCAACGAAGCAAAACAAGATGTCTTAAGGGAGCTGCGGGGACGCGAAACTGATAAGAATCAAGCAACAGGTGAACCTGATGTTCCACCTGATAAGAATCAAGCAACAGGTCAACATGCAGTTGTTCTGGATCAAAACGAAGAGGCTTATTTTGAAGAAGTAGAGCAGAAGATTGAAGAAATCAAGCAGGAACTTAAAGAGCAGCTGAAGATCAAATGGATCGTGGACAAGATTAAACATCACTTACAGGATCAGTGCCCCCTCATTATCCTAAAATTTGATCAGATGATGGATGGATCCCGATGGGAGGAGATCAGAAAGGCTCTGAGCCTGTTAGAATTGAGTGCAGATGCATTGATCTTCACCACAGGGAGCACAGAACAGGCTAAAGGATATTGCTATCCACCACGAGAACCTATAGATCACTGTTCTCTTGTTGGTCTCTACTATTATACGGTGCTCAAGCTTACTAGCAAGCACAAGAATGAAGACAACGATAACGCACAGATTTTTCGTGGCATCTTGGAGGAGTGTGAGGGGCATGAATTCTGCATGAAGATCTTCACTCATGCTGTGTATGCTAATCCCAAGAGGAGCAATGAAGAGTTAAGGAAGCTACACAGCACCCTGCAGTCTCCAAAAAAATCATTCGACACCATAGCTAAGAAGATGTTCATGTACTCTTACAATGATCTACCTAAAGAATACAAGTCATGCTTGCTGTACCTAGCTATCTTCCCCAAGGGACAGAAGATTAGGCGGTCAACCTTGATTGCAAGGTGGGTTGCAGAAGGGTTGACATTCAAGGAAGATTGGCCCAGCTCTGTGTATCAGGCAAATCGATGTTTTGATGCCCTCATCCGTCGGTGGCTTGTTTATCCTGATGATATTAGTGCCACAGGAAAGATCAAGAGCTGTGTGGTAGGCGATCCAGTTCATGGGTTCATTACCGCAATTGCCAGAAAACAACATATTGTGGAGACACGCCTATCACATCACTTGGCTCGCCACTTCTCCATTTTCAATGATCTTCGACTCCGCAGCTCTGATAGAATTGGCACGTTCTTCCAAGGCCTCTCACGATCATCTCGAGTATCCCTACTCAAGGTGCTAGATCTAGAAGGTTGTCAGTGCTTTGCTAGTAAGAATCAGCGGTACCTCAAGGACATCTGCAACAAGATGTTACTGCTCAAATATCTGAGCCTAAAGGGAACAGATATTACCCAGCTGCCCAGTGAAATCAACTGCCTCCGCGAGCTAGAGGTATTGGATATCCGAGAAACCAAGGTGCCTGCAAATGCAACAGTAAATGTCCTGCTCTTGAAGCTGAAGCGTCTACTTGCTGGTGCTAGTCAGATTGATCCAACTCCAAGAAATTTTGTCACTAACGTCCGGATTCCTTCCAGGATAGACAAGATGATAAACATAGAGGTACTATCTAATGTCAAGGCCCAGCACCATGATAATTTGGAAGATATTGGGAAGCTATGCCAGTTGAGGAAGCTAGGTGTGGTTATTGATGGTAAGAAAAGTCACCTTGGGAGTTTGCTTAAAGCGATCAGTGACCTACATGCAAGCCTCCGTTCTCTGTCAATCACTATTCCCACAACCACACTCGAGGTTACTCCTTCAAGCCCAGAGTTACAAGATATTGCATCTCGCCTAAAACACCATCCTGAGTTTCTTGAGAGTCTAAGCATCAGTGGAGCCAAGCATCTTTTTCCATTGTTGACCGAAGGTGGTAATAAGAAACTTGCCAAGGTAACTCTAAGCAACACCCCACTGAACCAAGATGATCTGAAGTTCTTTGCCCAGCTGCCCATGTTACAGTGTGTTAGGCTCCGACACATTTCATGCACCGAGAGTGTGCTCAACTTCAAGAAAGATGATTTCAAATGCCTCAAGTACCTTCTTATTGAGGGCTCAAACTTGACTAATATTACTTTTGAGGATGAGGCAGCCTGTGAGCTCGAGAAGATGGTTTTATCTTCCACTTGCATAGAGTCTATTTCTGGAGTTCATGGGCTTCCAAAATTCGAAGAGCTTGAGTTGAACAGCAGCAGCTGCGGAAGGTTGCTATCATCATGTTTTTACAATGTTGAACGAATAGCCAAGCTGACTCTTCGTGGTACATTGCTGAAGCAAGGTGATCTACGAATCATCGCCAGGGAACTAAATATATGCTGCCTAGTGCTCTTGGAAAACTCTTTTGACATAAGCCAGAACCAGATTACCTTCGAAAAAGAAGAGTTCATATGGCTCAAACTTCTTAGTGTTGATTGCTCTACCATCACCAAGATTAACTTCATCACTGGATCAGCTCCTAGGCTCAAGAAGATCGTCTGGTCATCTTTCACCTCTCTCTCTGGCATCAACAACCTTCCTAGGTTGAAGGAGCTCGAGTTCAATGGATACTCAGTCCCAAATGATGTGGAAGAAGCCATTAAAAACAATAAAAGCATAAATCTTAAACATAATAAACCATAA
Protein Sequence
  • >LOC_Os11g37050.1
    MTELAAGAVSSLLVVIRNEAVLLGGVRDDVQFIKEEMESMNSFLGHLARSAPQGGEHDEQVRTWMNQVRLLAQDCNNCIDLYLYSGNPEIHRAKGRLRRHLWWVYWSLRKMVAQHRAAIQLRQLKDRARDVGERRLRYGVEIPATTKAAAPDATGGYAAGDDEEEDEDDREGQFAVATPTLAHHSARWPVFEPPSLDDYVKAKLLEWIEGVPGNAIVTLSIAIVAPDADNKEVLAIAHETLVAPDYYYRRSIMVNVPAVHLDFLPLRPKEVLYYILRELEREEAAGSQKQPTDQGEWEEEDPDPWQDYYKKCGIYRSKKGVLDKIKRNIKKMNIYEKLDKIKSDIREGQHKSGKLLLLKLQEKGADQVDLHVLLQLLVLQSQQDQAKNKAVDTHKLPEWNDNLIEKLAMRLKDHMEADEKTKKLNEQTGVEEETAVRQGGGGERKEDEKDERGDGEEEGKEERRDMEKGEERKEQQQEEQEKEGRKEEQNEVRKETEGRKEQVAGEEEEKEDHDADNDEDSNDDDDDEEEEEEDDDDEEEPIHLHEDQYEQILREVFTKNASSKAQEQDKLVAEQATKTAATTLDEERIKQMINEAKQDVLRELRGRETDKNQATGEPDVPPDKNQATGQHAVVLDQNEEAYFEEVEQKIEEIKQELKEQLKIKWIVDKIKHHLQDQCPLIILKFDQMMDGSRWEEIRKALSLLELSADALIFTTGSTEQAKGYCYPPREPIDHCSLVGLYYYTVLKLTSKHKNEDNDNAQIFRGILEECEGHEFCMKIFTHAVYANPKRSNEELRKLHSTLQSPKKSFDTIAKKMFMYSYNDLPKEYKSCLLYLAIFPKGQKIRRSTLIARWVAEGLTFKEDWPSSVYQANRCFDALIRRWLVYPDDISATGKIKSCVVGDPVHGFITAIARKQHIVETRLSHHLARHFSIFNDLRLRSSDRIGTFFQGLSRSSRVSLLKVLDLEGCQCFASKNQRYLKDICNKMLLLKYLSLKGTDITQLPSEINCLRELEVLDIRETKVPANATVNVLLLKLKRLLAGASQIDPTPRNFVTNVRIPSRIDKMINIEVLSNVKAQHHDNLEDIGKLCQLRKLGVVIDGKKSHLGSLLKAISDLHASLRSLSITIPTTTLEVTPSSPELQDIASRLKHHPEFLESLSISGAKHLFPLLTEGGNKKLAKVTLSNTPLNQDDLKFFAQLPMLQCVRLRHISCTESVLNFKKDDFKCLKYLLIEGSNLTNITFEDEAACELEKMVLSSTCIESISGVHGLPKFEELELNSSSCGRLLSSCFYNVERIAKLTLRGTLLKQGDLRIIARELNICCLVLLENSFDISQNQITFEKEEFIWLKLLSVDCSTITKINFITGSAPRLKKIVWSSFTSLSGINNLPRLKELEFNGYSVPNDVEEAIKNNKSINLKHNKP*