Abstract
BACKGROUND: NAC transcription factors belong to a large family of plant-specific transcription factors with more than 100 family members in monocot and dicot species. To date, the majority of the studied NAC proteins are involved in the response to abiotic stress, to biotic stress and in the regulation of developmental processes. Maize NAC transcription factors involved in the biotic stress response have not yet been identified. RESULTS: We have found that two NAC transcription factors, ZmNAC41 and ZmNAC100, are transcriptionally induced both during the initial biotrophic as well as the ensuing necrotrophic colonization of maize leaves by the hemibiotrophic ascomycete fungus C. graminicola. ZmNAC41 transcripts were also induced upon infection with C. graminicola mutants that are defective in host penetration, while the induction of ZmNAC100 did not occur in such interactions. While ZmNAC41 transcripts accumulated specifically in response to jasmonate (JA), ZmNAC100 transcripts were also induced by the salicylic acid analog 2,6-dichloroisonicotinic acid (INA).To assess the phylogenetic relation of ZmNAC41 and ZmNAC100, we studied the family of maize NAC transcription factors based on the recently annotated B73 genome information. We identified 116 maize NAC transcription factor genes that clustered into 12 clades. ZmNAC41 and ZmNAC100 both belong to clade G and appear to have arisen by a recent gene duplication event. Including four other defence-related NAC transcription factors of maize and functionally characterized Arabidopsis and rice NAC transcription factors, we observed an enrichment of NAC transcription factors involved in host defense regulation in clade G. In silico analyses identified putative binding elements for the defence-induced ERF, Myc2, TGA and WRKY transcription factors in the promoters of four out of the six defence-related maize NAC transcription factors, while one of the analysed maize NAC did not contain any of these potential binding sites. CONCLUSIONS: Our study provides a systematic in silico analysis of maize NAC transcription factors in which we propose a nomenclature for maize genes encoding NAC transcription factors, based on their chromosomal position. We have further identified five pathogen-responsive maize NAC transcription factors that harbour putative binding elements for other defence-associated transcription factors in the proximal promoter region, indicating an involvement of the described NACs in the maize defence network. Our phylogenetic analysis has revealed that the majority of the yet described pathogen responsive NAC proteins from all plant species belong to clade G and suggests that they are phylogenetically related.
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Claims 11
Claim · model-written summary
ZmNAC41 was induced during biotrophic and necrotrophic C. graminicola colonization of maize leaves.
- Organ / stage
- maize leavese0
- Interactor
- C. graminicola
Colletotrichum graminicolae0pathogen - Gene
- ZmNAC41e0
- Outcome
- transcriptionally inducede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

_G.W._Wilson_1235209.jpg?width=500)
Verbatim
t0“maize”
e0“two NAC transcription factors, ZmNAC41 and ZmNAC100, are transcriptionally induced both during the initial biotrophic as well as the ensuing necrotrophic colonization of maize leaves by the hemibiotrophic ascomycete fungus C. graminicola”
Claim · model-written summary
ZmNAC100 was induced during biotrophic and necrotrophic C. graminicola colonization of maize leaves.
- Organ / stage
- maize leavese0
- Interactor
- C. graminicola
Colletotrichum graminicolae0pathogen - Gene
- ZmNAC100e0
- Outcome
- transcriptionally inducede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

_G.W._Wilson_1235209.jpg?width=500)
Verbatim
t0“maize”
e0“two NAC transcription factors, ZmNAC41 and ZmNAC100, are transcriptionally induced both during the initial biotrophic as well as the ensuing necrotrophic colonization of maize leaves by the hemibiotrophic ascomycete fungus C. graminicola”
Claim · model-written summary
ZmNAC41 transcripts were induced by penetration-defective C. graminicola mutants.
- Interactor
- C. graminicola mutants
Colletotrichum graminicolae0pathogen - Gene
- ZmNAC41e0
- Outcome
- transcripts were also inducede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

_G.W._Wilson_1235209.jpg?width=500)
Verbatim
t0“maize”
e0“ZmNAC41 transcripts were also induced upon infection with C. graminicola mutants that are defective in host penetration, while the induction of ZmNAC100 did not occur in such interactions.”
Claim · model-written summary
ZmNAC100 induction did not occur with penetration-defective C. graminicola mutants.
- Interactor
- C. graminicola mutants
Colletotrichum graminicolae0pathogen - Gene
- ZmNAC100e0
- Outcome
- induction of ZmNAC100 did not occure0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

_G.W._Wilson_1235209.jpg?width=500)
Verbatim
t0“maize”
e0“ZmNAC41 transcripts were also induced upon infection with C. graminicola mutants that are defective in host penetration, while the induction of ZmNAC100 did not occur in such interactions.”
Claim · model-written summary
ZmNAC41 transcripts accumulated specifically in response to jasmonate.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“maize”
e0“ZmNAC41 transcripts accumulated specifically in response to jasmonate (JA), ZmNAC100 transcripts were also induced by the salicylic acid analog 2,6-dichloroisonicotinic acid (INA).”
Claim · model-written summary
ZmNAC100 transcripts were induced by INA.
- Treatment
- 2,6-dichloroisonicotinic acid (INA)e0
- Gene
- ZmNAC100e0
- Outcome
- transcripts were also inducede0
- Direction
- inducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“maize”
e0“ZmNAC41 transcripts accumulated specifically in response to jasmonate (JA), ZmNAC100 transcripts were also induced by the salicylic acid analog 2,6-dichloroisonicotinic acid (INA).”
Claim · model-written summary
The maize genome analysis identified 116 NAC transcription factor genes in 12 clades.
- Gene
- NAC transcription factor genese0
- Outcome
- NAC transcription factor genes that clustered into 12 cladese0
- Value
- 116e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“maize”
e0“We identified 116 maize NAC transcription factor genes that clustered into 12 clades.”
Claim · model-written summary
ZmNAC41 and ZmNAC100 belong to clade G and appear recently duplicated.
- Gene
- ZmNAC41 and ZmNAC100e0
- Outcome
- belong to clade Ge0
- Qualifier
- appear to have arisen by a recent gene duplication evente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“maize”
e0“ZmNAC41 and ZmNAC100 both belong to clade G and appear to have arisen by a recent gene duplication event.”
Claim · model-written summary
Clade G was enriched for host-defense NAC transcription factors.
- Gene
- NAC transcription factorse0
- Outcome
- enrichment of NAC transcription factors involved in host defense regulation in clade Ge0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“maize”
e0“we observed an enrichment of NAC transcription factors involved in host defense regulation in clade G.”
Claim · model-written summary
Four of six defense-related maize NAC promoters had putative binding elements for defense-induced transcription factors.
- Gene
- ERF, Myc2, TGA and WRKY transcription factorse0
- Outcome
- putative binding elementse0
- Value
- four out of the sixe0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“maize”
e0“In silico analyses identified putative binding elements for the defence-induced ERF, Myc2, TGA and WRKY transcription factors in the promoters of four out of the six defence-related maize NAC transcription factors”
Claim · model-written summary
One analyzed maize NAC lacked these potential binding sites.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“maize”
e0“one of the analysed maize NAC did not contain any of these potential binding sites.”
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