Abstract
Summary The CBF cold response pathway has a prominent role in cold acclimation. The pathway includes action of three transcription factors, CBF1, 2 and 3 (also known as DREB1b, c and a, respectively), that are rapidly induced in response to low temperature followed by expression of the CBF-targeted genes (the CBF regulon) that act in concert to increase plant-freezing tolerance. The results of transcriptome profiling and mutagenesis experiments, however, indicate that additional cold response pathways exist and may have important roles in life at low temperature. To further understand the roles that the CBF proteins play in configuring the low temperature transcriptome and to identify additional transcription factors with roles in cold acclimation, we used the Affymetrix GeneChip containing probe sets for approximately 24,000 Arabidopsis genes to define a core set of cold-responsive genes and to determine which genes were targets of CBF2 and 6 other transcription factors that appeared to be coordinately regulated with CBF2. A total of 514 genes were placed in the core set of cold-responsive genes, 302 of which were upregulated and 212 downregulated. Hierarchical clustering and bioinformatic analysis indicated that the 514 cold-responsive transcripts could be assigned to one of seven distinct expression classes and identified multiple potential novel cis-acting cold-regulatory elements. Eighty-five cold-induced genes and eight cold-repressed genes were assigned to the CBF2 regulon. An additional nine cold-induced genes and 15 cold-repressed genes were assigned to a regulon controlled by ZAT12. Of the 25 core cold-induced genes that were most highly upregulated (induced over 15-fold), 19 genes (84%) were induced by CBF2 and another two genes (8%) were regulated by both CBF2 and ZAT12. Thus, the large majority (92%) of the most highly induced genes belong to the CBF and ZAT12 regulons. Constitutive expression of ZAT12 in Arabidopsis caused a small, but reproducible, increase in freezing tolerance, indicating a role for the ZAT12 regulon in cold acclimation. In addition, ZAT12 downregulated the expression of the CBF genes indicating a role for ZAT12 in a negative regulatory circuit that dampens expression of the CBF cold response pathway.
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Claims 13
Claim · model-written summary
The core cold-responsive set comprised 514 genes.
- Taxon
- Arabidopsis
Arabidopsise0 - Outcome
- core set of cold-responsive genese1
- Value
- 514 genese1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“A total of 514 genes were placed in the core set of cold-responsive genes”
Claim · model-written summary
Among core cold-responsive genes, 302 were upregulated.
- Taxon
- Arabidopsis
Arabidopsise0 - Outcome
- upregulatede1
- Direction
- upregulatede1
- Value
- 302e1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“302 of which were upregulated”
Claim · model-written summary
Among core cold-responsive genes, 212 were downregulated.
- Taxon
- Arabidopsis
Arabidopsise0 - Outcome
- downregulatede1
- Direction
- downregulatede1
- Value
- 212e1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“212 downregulated”
Claim · model-written summary
Cold-responsive transcripts fell into seven expression classes.
- Taxon
- Arabidopsis
Arabidopsise0 - Outcome
- distinct expression classese1
- Value
- sevene1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“514 cold-responsive transcripts could be assigned to one of seven distinct expression classes”
Claim · model-written summary
Analysis identified multiple potential novel cis-acting cold-regulatory elements.
- Taxon
- Arabidopsis
Arabidopsise0 - Outcome
- multiple potential novel cis-acting cold-regulatory elementse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“identified multiple potential novel cis-acting cold-regulatory elements”
Claim · model-written summary
CBF2 regulon included 85 cold-induced genes.
- Taxon
- Arabidopsis
Arabidopsise0 - Gene
- CBF2e1
- Outcome
- cold-induced genese1
- Value
- Eighty-fivee1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“Eighty-five cold-induced genes and eight cold-repressed genes were assigned to the CBF2 regulon”
Claim · model-written summary
CBF2 regulon included eight cold-repressed genes.
- Taxon
- Arabidopsis
Arabidopsise0 - Gene
- CBF2e1
- Outcome
- cold-repressed genese1
- Value
- eighte1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“Eighty-five cold-induced genes and eight cold-repressed genes were assigned to the CBF2 regulon”
Claim · model-written summary
ZAT12 regulon included nine cold-induced genes.
- Taxon
- Arabidopsis
Arabidopsise0 - Gene
- ZAT12e1
- Outcome
- cold-induced genese1
- Value
- ninee1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“nine cold-induced genes and 15 cold-repressed genes were assigned to a regulon controlled by ZAT12”
Claim · model-written summary
ZAT12 regulon included 15 cold-repressed genes.
- Taxon
- Arabidopsis
Arabidopsise0 - Gene
- ZAT12e1
- Outcome
- cold-repressed genese1
- Value
- 15e1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“nine cold-induced genes and 15 cold-repressed genes were assigned to a regulon controlled by ZAT12”
Claim · model-written summary
Among the 25 most induced core genes, 19 were induced by CBF2.
- Taxon
- Arabidopsis
Arabidopsise0 - Gene
- CBF2e1
- Outcome
- genes (84%) were inducede1
- Value
- 19 genes (84%)e1
- Qualifier
- induced over 15-folde1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“Of the 25 core cold-induced genes that were most highly upregulated (induced over 15-fold), 19 genes (84%) were induced by CBF2”
Claim · model-written summary
Two of the most induced core genes were regulated by both CBF2 and ZAT12.
- Taxon
- Arabidopsis
Arabidopsise0 - Gene
- CBF2 and ZAT12e1
- Outcome
- genes (8%) were regulatede1
- Value
- two genes (8%)e1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“approximately 24,000 Arabidopsis genes”
e1“another two genes (8%) were regulated by both CBF2 and ZAT12”
Claim · model-written summary
Constitutive ZAT12 expression slightly increased freezing tolerance.
- Taxon
- Arabidopsis
Arabidopsise0 - Treatment
- Constitutive expression of ZAT12e0
- Gene
- ZAT12e0
- Outcome
- freezing tolerancee0
- Direction
- increasee0
- Qualifier
- small, but reproduciblee0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Constitutive expression of ZAT12 in Arabidopsis caused a small, but reproducible, increase in freezing tolerance”
Claim · model-written summary
ZAT12 downregulated CBF gene expression.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- ZAT12e0
- Outcome
- expression of the CBF genese0
- Direction
- downregulatede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ZAT12 downregulated the expression of the CBF genes indicating a role for ZAT12 in a negative regulatory circuit”
e1“approximately 24,000 Arabidopsis genes”
The same questions in other papers
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