Gene Details:
- Gene ID: Zm00001eb169360
- Gene Symbol: ZmSig2A
- Gene Name: sigma factor sig2A
- Genome: Zm-B73-REFERENCE-NAM-5.0
- Species: Zea mays
Functional Descriptions:
- A valine residue deletion in ZmSig2A, a sigma factor, accounts for a revertible leaf-color mutation in maize.
- The eal1 mutant with a weak allele of ZmSig2A represents a valuable genetic resource for investigating the regulation of ZmSig2A-mediated chloroplast development in maize.
- ZmSig2A encodes a σ factor essential for chloroplast biogenesis at the early seedling stage of maize.
- We propose that the Val deletion generated a new weak allele of ZmSig2A that cannot completely abolish the ZmSig2A function.
- Ectopic expression of ZmSig2A+ or ZmSig2AΔV driven by the cauliflower mosaic virus 35S promoter rescued the pale green leaf of the sig2 mutant, but ectopic expression of ZmSig2AΔV driven by the SIG2 promoter did not.
Function-related keywords:
- leaf , chloroplast , development , leaf-development , seedling , leaf-color , chloroplast-development , chloroplast-biogenesis
Literature:
- A valine residue deletion in ZmSig2A, a sigma factor, accounts for a revertible leaf-color mutation in maize DOI: 10.1016/j.cj.2021.01.005 ; PMID:
- Subdivision of Light Signaling Networks Contributes to Partitioning of C(4) Photosynthesis. DOI: 10.1104/pp.19.01053 ; PMID: 31862840
Related News:
Gene Resources:
- NCBI ID: 541716
- UniProt accessions: A0A1D6PSU6
Sequences:
cDNA Sequence
- >Zm00001eb169360_T002
CCAGAATTCGGCCCACTAACTCTCACTCCGGGTCGCGCCTGAGCAGGCAGGTCTGAGTTCTGCTCTCTGACTGAGCCTGAGCTCCCCTTATCCAGAGAGGCTAGAACAAACCTGAATTTCATCCCCACCGTCGCCTCGCTCGACCTCCACCTCTCTTTCCTCCCGGCAGCAATGGAGGCGGCACCCAGGTGAGGCGAGCAAGCAGCCAGCCCTGGCTCCCCGATGGCGTGCCTGGCGCCGCAGTTCAAGTGGGCGCCGTCGGCCGCCGCCCACATGCCTTCCTCGTCCTCGTCCTCGTCCTACTCCAGGTGCTCCGCGCTCCGCGTCCACTGCGCCGTCACCTCCGCCGCTGTCGTCGAGGACCGCACCAACGGCAGCGCGGCGCAGCTGAGGCTCGCTTACGCGGCTCCGGCGATTCAGAGAAACTTTGAGGCAACTCTGGCATCAGAAGCACTTCTAAATGAAGAGGCGGTAGTAGCAGCGGCAGCCGCAGAGGCTGTTGCTCTTGCCAGAGCAGCTGCCGAAGCTGCCCAAGAAGTTGTTCAGATAGTACAAAGAAACAACCATCAGCCGGTAATTAGACCAAAAAAGTGTGTGGACAACTACTTGGCAAATGAAATCCTTCGCACAGAGATGCGATCGAGCAGTCCTGACATGTATGCTAATAATCCTTTGATGGAGTATCTGGACTCTTATGGTATCATGGATGCTGAAGATGAATTAGATGCTTATGCACAGTATACTGAGAACATAGCTGTGAAATCTGCTCGTCAATCTGAGAGAAAAGCTAGGAGAACTAGAGCAGCAATAAAAGCTAGCACAACCCTCCATGCTTCACAAAAGGTTGCATTATCCTCAAAGAAGAAGCGGTCCAAGGGTTCCTCGTCTAGTATGAATCCTTTAGGTTCGTTGTGGAAGATGACCGGTAGGAGACTTCTTACAGCCAAGGAAGAGGTTGAATTCTCAGAAGGTATTCAGGATCTTTTGAAGCTTGAGGCGATCCAAGCTGAGCTTATAGAGTACAATGGCGGTCAGCCAACCTTCTCGCAGTGGGCAACAGCAGCTGGAGTTGATGAGAAAACTTTGCGCAAGCGCTTGAATTATGGTATTTATTGCAAGAACAGAATGGTAACATCTAATGTGAGACTTGTAATCTCTATTGCCAGAGAGTTTGAAGGCCCTGGAATGGACTTTTATGATCTTATTCAGGTTTTTTGTTACAGGAAGGAATGCAGGGCCTTATAAGGGGAGCTGAAAAATTTGATTCATCAAAAGGTTTTAGGTTCTCTACGTATTCTCACTGGTGGATCAAACAAGCAATGCGTAAATCTGTCTCAGAGCAATCCCAAATATTTCGCTTGCCTGCTCACATGGTTGAAGCAAGCTACCGTGTAAAGGAGTGTACAAAACGACTTCGCCGTAAGCTTAGAAGACGACCCACCAATGAAGAAATTGCAGTGGACACTGGTATTCCAATTAAACGAGTTGAGGCAGCAGTAAACCTCCCAAAATATAGTGTGTCCCTTGATAGCAAAATTGGTTCCACCGACATGACATATCAGGAGGTCACAGCTGATCCCAGTGCTGAGACAGCTGAAGAGATGCTCAACAGAATGTCCATGAAGAAGGACGTACACATGGCACTAGATACTCTCACCACTCGCGAGAAGCAAGTTGTTGTTCTGAGGTTTGGGCTCGAGGATGGTCGGATAAGAACCCTGCAGGAGATCGGTAACATCATGGGTGTGAGCAGGGAGAGAATCCGGCAGATCGAGTCTGGCGCGTTCCGGAAACTAAGGAGCAAGAAGAAGGTGAAGGCTTTGAAGGATTATCTGGTGCCAGTAGGCAATTGGTGAACCGTTTGTTTGCAGAATTTGTTTTCCTTTTGGTTCCTCTTTTTTTTTTTGAGGATTCTGTACTTGTACCATTTTACGTGGGTCCTCATGTAAAACACGCTATCAGAATACTTGCATCACAAGAGGAAAGAGTAGTTTAGTTCCGTGGGAAATGAAGTAAAATTCACAATTTCTTAGCTGCTGGTTATCTCTTAGAAATGAAATATGTTTGGACAAAACTGTGTAATGCATGAAAACTTTAGAGGCGC - >Zm00001eb169360_T001
CCAGAATTCGGCCCACTAACTCTCACTCCGGGTCGCGCCTGAGCAGGCAGGTCTGAGTTCTGCTCTCTGACTGAGCCTGAGCTCCCCTTATCCAGAGAGGCTAGAACAAACCTGAATTTCATCCCCACCGTCGCCTCGCTCGACCTCCACCTCTCTTTCCTCCCGGCAGCAATGGAGGCGGCACCCAGGTGAGGCGAGCAAGCAGCCAGCCCTGGCTCCCCGATGGCGTGCCTGGCGCCGCAGTTCAAGTGGGCGCCGTCGGCCGCCGCCCACATGCCTTCCTCGTCCTCGTCCTCGTCCTACTCCAGGTGCTCCGCGCTCCGCGTCCACTGCGCCGTCACCTCCGCCGCTGTCGTCGAGGACCGCACCAACGGCAGCGCGGCGCAGCTGAGGCTCGCTTACGCGGCTCCGGCGATTCAGAGAAACTTTGAGGCAACTCTGGCATCAGAAGCACTTCTAAATGAAGAGGCGGTAGTAGCAGCGGCAGCCGCAGAGGCTGTTGCTCTTGCCAGAGCAGCTGCCGAAGCTGCCCAAGAAGTTGTTCAGATAGTACAAAGAAACAACCATCAGCCGGTAATTAGACCAAAAAAGTGTGTGGACAACTACTTGGCAAATGAAATCCTTCGCACAGAGATGCGATCGAGCAGTCCTGACATGTATGCTAATAATCCTTTGATGGAGTATCTGGACTCTTATGGTATCATGGATGCTGAAGATGAATTAGATGCTTATGCACAGTATACTGAGAACATAGCTGTGAAATCTGCTCGTCAATCTGAGAGAAAAGCTAGGAGAACTAGAGCAGCAATAAAAGCTAGCACAACCCTCCATGCTTCACAAAAGGTTGCATTATCCTCAAAGAAGAAGCGGTCCAAGGGTTCCTCGTCTAGTATGAATCCTTTAGGTTCGTTGTGGAAGATGACCGGTAGGAGACTTCTTACAGCCAAGGAAGAGGTTGAATTCTCAGAAGGTATTCAGGATCTTTTGAAGCTTGAGGCGATCCAAGCTGAGCTTATAGAGTACAATGGCGGTCAGCCAACCTTCTCGCAGTGGGCAACAGCAGCTGGAGTTGATGAGAAAACTTTGCGCAAGCGCTTGAATTATGGTATTTATTGCAAGAACAGAATGGTAACATCTAATGTGAGACTTGTAATCTCTATTGCCAGAGAGTTTGAAGGCCCTGGAATGGACTTTTATGATCTTATTCAGGTTTTTTGTTACAGGAAGGAATGCAGGGCCTTATAAGGGGAGCTGAAAAATTTGATTCATCAAAAGGTTTTAGGTTCTCTACGTATTCTCACTGGTGGATCAAACAAGCAATGCGTAAATCTGTCTCAGAGCAATCCCAAATATTTCGCTTGCCTGCTCACATGGTTGAAGCAAGCTACCGTGTAAAGGAGTGTACAAAACGACTTCGCCGTAAGCTTAGAAGACGACCCACCAATGAAGAAATTGCAGTGGACACTGGTATTCCAATTAAACGAGTTGAGGCAGCAGTAAACCTCCCAAAATATAGTGTGTCCCTTGATAGCAAAATTGGTTCCACCGACATGACATATCAGGAGGTCACAGCTGATCCCAGTGCTGAGACAGCTGAAGAGATGCTCAACAGAATGTCCATGAAGAAGGACGTACACATGGCACTAGATACTCTCACCACTCGCGAGAAGCAAGTTGTTGTTCTGAGGTTTGGGCTCGAGGATGGTCGGATAAGAACCCTGCAGGAGATCGGTAACATCATGGGTGTGAGCAGGGAGAGAATCCGGCAGATCGAGTCTGGCGCGTTCCGGAAACTAAGGAGCAAGAAGAAGGTGAAGGCTTTGAAGGATTATCTGGTGCCAGTAGGCAATTGGTGAACCGTTTGTTTGCAGAATTTGTTTTCCTTTTGGTTCCTCTTTTTTTTTTTGAGGATTCTGTACTTGTACCATTTTACGTGGGTCCTCATGTAAAACACGCTATCAGAATACTTGCATCACAAGAGGAAAGAGTAGTTTAGTTCCGTGGGAAATGAAGTAAAATTCACAATTTCTTAGCTGCTGGTTATCTCTTAGAAATGAAATATGTTTGGACAAAACTGTGTAATGCATGAAAACTTTAGAGGCGC - >Zm00001eb169360_T003
CCAGAATTCGGCCCACTAACTCTCACTCCGGGTCGCGCCTGAGCAGGCAGGTCTGAGTTCTGCTCTCTGACTGAGCCTGAGCTCCCCTTATCCAGAGAGGCTAGAACAAACCTGAATTTCATCCCCACCGTCGCCTCGCTCGACCTCCACCTCTCTTTCCTCCCGGCAGCAATGGAGGCGGCACCCAGGTGAGGCGAGCAAGCAGCCAGCCCTGGCTCCCCGATGGCGTGCCTGGCGCCGCAGTTCAAGTGGGCGCCGTCGGCCGCCGCCCACATGCCTTCCTCGTCCTCGTCCTCGTCCTACTCCAGGTGCTCCGCGCTCCGCGTCCACTGCGCCGTCACCTCCGCCGCTGTCGTCGAGGACCGCACCAACGGCAGCGCGGCGCAGCTGAGGCTCGCTTACGCGGCTCCGGCGATTCAGAGAAACTTTGAGGCAACTCTGGCATCAGAAGCACTTCTAAATGAAGAGGCGGTAGTAGCAGCGGCAGCCGCAGAGGCTGTTGCTCTTGCCAGAGCAGCTGCCGAAGCTGCCCAAGAAGTTGTTCAGATAGTACAAAGAAACAACCATCAGCCGGTAATTAGACCAAAAAAGTGTGTGGACAACTACTTGGCAAATGAAATCCTTCGCACAGAGATGCGATCGAGCAGTCCTGACATGTATGCTAATAATCCTTTGATGGAGTATCTGGACTCTTATGGTATCATGGATGCTGAAGATGAATTAGATGCTTATGCACAGTATACTGAGAACATAGCTGTGAAATCTGCTCGTCAATCTGAGAGAAAAGCTAGGAGAACTAGAGCAGCAATAAAAGCTAGCACAACCCTCCATGCTTCACAAAAGGTTGCATTATCCTCAAAGAAGAAGCGGTCCAAGGGTTCCTCGTCTAGTATGAATCCTTTAGGTTCGTTGTGGAAGATGACCGGTAGGAGACTTCTTACAGCCAAGGAAGAGGTTGAATTCTCAGAAGGTATTCAGGATCTTTTGAAGCTTGAGGCGATCCAAGCTGAGCTTATAGAGTACAATGGCGGTCAGCCAACCTTCTCGCAGTGGGCAACAGCAGCTGGAGTTGATGAGAAAACTTTGCGCAAGCGCTTGAATTATGGTATTTATTGCAAGAACAGAATGGTAACATCTAATGTGAGACTTGTAATCTCTATTGCCAGAGAGTTTGAAGGCCCTGGAATGGACTTTTATGATCTTATTCAGGTTTTTTGTTACAGGAAGGAATGCAGGGCCTTATAAGGGGAGCTGAAAAATTTGATTCATCAAAAGGTTTTAGGTTCTCTACGTATTCTCACTGGTGGATCAAACAAGCAATGCGTAAATCTGTCTCAGAGCAATCCCAAATATTTCGCTTGCCTGCTCACATGGTTGAAGCAAGCTACCGTGTAAAGGAGTGTACAAAACGACTTCGCCGTAAGCTTAGAAGACGACCCACCAATGAAGAAATTGCAGTGGACACTGGTATTCCAATTAAACGAGTTGAGGCAGCAGTAAACCTCCCAAAATATAGTGTGTCCCTTGATAGCAAAATTGGTTCCACCGACATGACATATCAGGAGGTCACAGCTGATCCCAGTGCTGAGACAGCTGAAGAGATGCTCAACAGAATGTCCATGAAGAAGGACGTACACATGGCACTAGATACTCTCACCACTCGCGAGAAGCAAGTTGTTGTTCTGAGGTTTGGGCTCGAGGATGGTCGGATAAGAACCCTGCAGGAGATCGGTAACATCATGGGTGTGAGCAGGGAGAGAATCCGGCAGATCGAGTCTGGCGCGTTCCGGAAACTAAGGAGCAAGAAGAAGGTGAAGGCTTTGAAGGATTATCTGGTGCCAGTAGGCAATTGGTGAACCGTTTGTTTGCAGAATTTGTTTTCCTTTTGGTTCCTCTTTTTTTTTTTGAGGATTCTGTACTTGTACCATTTTACGTGGGTCCTCATGTAAAACACGCTATCAGAATACTTGCATCACAAGAGGAAAGAGTAGTTTAGTTCCGTGGGAAATGAAGTAAAATTCACAATTTCTTAGCTGCTGGTTATCTCTTAGAAATGAAATATGTTTGGACAAAACTGTGTAATGCATGAAAACTTTAGAGGCGC
CDS Sequence
- >Zm00001eb169360_T002
ATGGCGTGCCTGGCGCCGCAGTTCAAGTGGGCGCCGTCGGCCGCCGCCCACATGCCTTCCTCGTCCTCGTCCTCGTCCTACTCCAGGTGCTCCGCGCTCCGCGTCCACTGCGCCGTCACCTCCGCCGCTGTCGTCGAGGACCGCACCAACGGCAGCGCGGCGCAGCTGAGGCTCGCTTACGCGGCTCCGGCGATTCAGAGAAACTTTGAGGCAACTCTGGCATCAGAAGCACTTCTAAATGAAGAGGCGGTAGTAGCAGCGGCAGCCGCAGAGGCTGTTGCTCTTGCCAGAGCAGCTGCCGAAGCTGCCCAAGAAGTTGTTCAGATAGTACAAAGAAACAACCATCAGCCGGTAATTAGACCAAAAAAGTGTGTGGACAACTACTTGGCAAATGAAATCCTTCGCACAGAGATGCGATCGAGCAGTCCTGACATGTATGCTAATAATCCTTTGATGGAGTATCTGGACTCTTATGGTATCATGGATGCTGAAGATGAATTAGATGCTTATGCACAGTATACTGAGAACATAGCTGTGAAATCTGCTCGTCAATCTGAGAGAAAAGCTAGGAGAACTAGAGCAGCAATAAAAGCTAGCACAACCCTCCATGCTTCACAAAAGGTTGCATTATCCTCAAAGAAGAAGCGGTCCAAGGGTTCCTCGTCTAGTATGAATCCTTTAGGTTCGTTGTGGAAGATGACCGGTAGGAGACTTCTTACAGCCAAGGAAGAGGTTGAATTCTCAGAAGGTATTCAGGATCTTTTGAAGCTTGAGGCGATCCAAGCTGAGCTTATAGAGTACAATGGCGGTCAGCCAACCTTCTCGCAGTGGGCAACAGCAGCTGGAGTTGATGAGAAAACTTTGCGCAAGCGCTTGAATTATGGTATTTATTGCAAGAACAGAATGGTAACATCTAATGTGAGACTTGTAATCTCTATTGCCAGAGAGTTTGAAGGCCCTGGAATGGACTTTTATGATCTTATTCAGGTTTTTTGTTACAGGAAGGAATGCAGGGCCTTATAA - >Zm00001eb169360_T001
ATGGCGTGCCTGGCGCCGCAGTTCAAGTGGGCGCCGTCGGCCGCCGCCCACATGCCTTCCTCGTCCTCGTCCTCGTCCTACTCCAGGTGCTCCGCGCTCCGCGTCCACTGCGCCGTCACCTCCGCCGCTGTCGTCGAGGACCGCACCAACGGCAGCGCGGCGCAGCTGAGGCTCGCTTACGCGGCTCCGGCGATTCAGAGAAACTTTGAGGCAACTCTGGCATCAGAAGCACTTCTAAATGAAGAGGCGGTAGTAGCAGCGGCAGCCGCAGAGGCTGTTGCTCTTGCCAGAGCAGCTGCCGAAGCTGCCCAAGAAGTTGTTCAGATAGTACAAAGAAACAACCATCAGCCGGTAATTAGACCAAAAAAGTGTGTGGACAACTACTTGGCAAATGAAATCCTTCGCACAGAGATGCGATCGAGCAGTCCTGACATGTATGCTAATAATCCTTTGATGGAGTATCTGGACTCTTATGGTATCATGGATGCTGAAGATGAATTAGATGCTTATGCACAGTATACTGAGAACATAGCTGTGAAATCTGCTCGTCAATCTGAGAGAAAAGCTAGGAGAACTAGAGCAGCAATAAAAGCTAGCACAACCCTCCATGCTTCACAAAAGGTTGCATTATCCTCAAAGAAGAAGCGGTCCAAGGGTTCCTCGTCTAGTATGAATCCTTTAGGTTCGTTGTGGAAGATGACCGGTAGGAGACTTCTTACAGCCAAGGAAGAGGTTGAATTCTCAGAAGGTATTCAGGATCTTTTGAAGCTTGAGGCGATCCAAGCTGAGCTTATAGAGTACAATGGCGGTCAGCCAACCTTCTCGCAGTGGGCAACAGCAGCTGGAGTTGATGAGAAAACTTTGCGCAAGCGCTTGAATTATGGTATTTATTGCAAGAACAGAATGGTAACATCTAATGTGAGACTTGTAATCTCTATTGCCAGAGAGTTTGAAGGCCCTGGAATGGACTTTTATGATCTTATTCAGGTTTTTTGTTACAGGAAGGAATGCAGGGCCTTATAA - >Zm00001eb169360_T003
ATGGCGTGCCTGGCGCCGCAGTTCAAGTGGGCGCCGTCGGCCGCCGCCCACATGCCTTCCTCGTCCTCGTCCTCGTCCTACTCCAGGTGCTCCGCGCTCCGCGTCCACTGCGCCGTCACCTCCGCCGCTGTCGTCGAGGACCGCACCAACGGCAGCGCGGCGCAGCTGAGGCTCGCTTACGCGGCTCCGGCGATTCAGAGAAACTTTGAGGCAACTCTGGCATCAGAAGCACTTCTAAATGAAGAGGCGGTAGTAGCAGCGGCAGCCGCAGAGGCTGTTGCTCTTGCCAGAGCAGCTGCCGAAGCTGCCCAAGAAGTTGTTCAGATAGTACAAAGAAACAACCATCAGCCGGTAATTAGACCAAAAAAGTGTGTGGACAACTACTTGGCAAATGAAATCCTTCGCACAGAGATGCGATCGAGCAGTCCTGACATGTATGCTAATAATCCTTTGATGGAGTATCTGGACTCTTATGGTATCATGGATGCTGAAGATGAATTAGATGCTTATGCACAGTATACTGAGAACATAGCTGTGAAATCTGCTCGTCAATCTGAGAGAAAAGCTAGGAGAACTAGAGCAGCAATAAAAGCTAGCACAACCCTCCATGCTTCACAAAAGGTTGCATTATCCTCAAAGAAGAAGCGGTCCAAGGGTTCCTCGTCTAGTATGAATCCTTTAGGTTCGTTGTGGAAGATGACCGGTAGGAGACTTCTTACAGCCAAGGAAGAGGTTGAATTCTCAGAAGGTATTCAGGATCTTTTGAAGCTTGAGGCGATCCAAGCTGAGCTTATAGAGTACAATGGCGGTCAGCCAACCTTCTCGCAGTGGGCAACAGCAGCTGGAGTTGATGAGAAAACTTTGCGCAAGCGCTTGAATTATGGTATTTATTGCAAGAACAGAATGGTAACATCTAATGTGAGACTTGTAATCTCTATTGCCAGAGAGTTTGAAGGCCCTGGAATGGACTTTTATGATCTTATTCAGGTTTTTTGTTACAGGAAGGAATGCAGGGCCTTATAA
Protein Sequence
- >Zm00001eb169360_P002
MACLAPQFKWAPSAAAHMPSSSSSSSYSRCSALRVHCAVTSAAVVEDRTNGSAAQLRLAYAAPAIQRNFEATLASEALLNEEAVVAAAAAEAVALARAAAEAAQEVVQIVQRNNHQPVIRPKKCVDNYLANEILRTEMRSSSPDMYANNPLMEYLDSYGIMDAEDELDAYAQYTENIAVKSARQSERKARRTRAAIKASTTLHASQKVALSSKKKRSKGSSSSMNPLGSLWKMTGRRLLTAKEEVEFSEGIQDLLKLEAIQAELIEYNGGQPTFSQWATAAGVDEKTLRKRLNYGIYCKNRMVTSNVRLVISIAREFEGPGMDFYDLIQVFCYRKECRAL - >Zm00001eb169360_P001
MACLAPQFKWAPSAAAHMPSSSSSSSYSRCSALRVHCAVTSAAVVEDRTNGSAAQLRLAYAAPAIQRNFEATLASEALLNEEAVVAAAAAEAVALARAAAEAAQEVVQIVQRNNHQPVIRPKKCVDNYLANEILRTEMRSSSPDMYANNPLMEYLDSYGIMDAEDELDAYAQYTENIAVKSARQSERKARRTRAAIKASTTLHASQKVALSSKKKRSKGSSSSMNPLGSLWKMTGRRLLTAKEEVEFSEGIQDLLKLEAIQAELIEYNGGQPTFSQWATAAGVDEKTLRKRLNYGIYCKNRMVTSNVRLVISIAREFEGPGMDFYDLIQVFCYRKECRAL - >Zm00001eb169360_P003
MACLAPQFKWAPSAAAHMPSSSSSSSYSRCSALRVHCAVTSAAVVEDRTNGSAAQLRLAYAAPAIQRNFEATLASEALLNEEAVVAAAAAEAVALARAAAEAAQEVVQIVQRNNHQPVIRPKKCVDNYLANEILRTEMRSSSPDMYANNPLMEYLDSYGIMDAEDELDAYAQYTENIAVKSARQSERKARRTRAAIKASTTLHASQKVALSSKKKRSKGSSSSMNPLGSLWKMTGRRLLTAKEEVEFSEGIQDLLKLEAIQAELIEYNGGQPTFSQWATAAGVDEKTLRKRLNYGIYCKNRMVTSNVRLVISIAREFEGPGMDFYDLIQVFCYRKECRAL
Source: Genetic information comes from the article