Paper · 2025 · The Plant Journal

Cytokinin mediates age‐dependent drought response by regulating the removal reactive oxygen species in apple

Xumei Jia; Shuo Xu; Jiangtong Wei; et al.

13
claims
19
verbatim quotes
4
citations
4
research areas
—
Not open access

Abstract

Plants exhibit age-dependent drought responses during juvenile-to-adult phase transition. However, the specific regulatory molecular mechanisms remain unknown. In this study, juvenile apple plants exhibited better drought tolerance than adult apple plants because of the age-dependent changes in root vitality, cytokinin (CK) levels, and redox status in roots. The study uncovers CYTOKININ OXIDASE/DEHYDROGENASE5 (MdCKX5) as a negative regulator of drought tolerance in apple roots. Silencing of MdCKX5 caused CK accumulation thereby enhancing drought tolerance by increasing root vitality and preventing the accumulation of reactive oxygen species. In contrast, overexpression of MdCKX5 reduced drought tolerance in apple roots. Yeast one-hybrid, dual-luciferase and electrophoretic mobility shift assays revealed that apple transcription factor MdSPL1 directly binds to the promoters of MdCKX5 thereby repressing its expression. Overexpression of MdSPL1 in apple roots increased the CK content thereby enhancing drought tolerance. Further analysis revealed that MdMYB23, a positive CK-responsive gene, interacts with MdSPL1 to alleviate the repression of MdCKX5 expression by MdSPL1. In general, drought stress significantly downregulated MdMYB23, thereby activating the MdSPL1-mediated repression of MdCKX5 transcription by releasing the MdSPL1. This phenomenon led to enhanced drought tolerance in juvenile apple roots by increasing the CK levels. However, adult apple roots lost the capacity to activate this cascade. These findings provide new insights into the molecular mechanisms of CK-mediated age-dependent drought tolerance during the vegetative phase change in apples.

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Claims 13

Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

Juvenile apple plants had better drought tolerance than adults.

Material
juvenile apple plantse0
Outcome
drought tolerancee0
Direction
bettere0
Comparator
adult apple plantse0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“juvenile apple plants exhibited better drought tolerance than adult apple plants”

✓ located verbatim in the source · abstract, characters 175–255 · show in text ↑
Mechanismobservedoriginal resultnegativemeasured

Claim · model-written summary

MdCKX5 negatively regulates drought tolerance in apple roots.

Organ / stage
rootse0
Gene
MdCKX5e0
Outcome
drought tolerancee0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“MdCKX5) as a negative regulator of drought tolerance in apple roots”

✓ located verbatim in the source · abstract, characters 414–481 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

MdCKX5 silencing accumulated cytokinin.

Treatment
Silencing of MdCKX5e0
Gene
MdCKX5e0
Outcome
CK accumulatione0
Direction
accumulatione0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“Silencing of MdCKX5 caused CK accumulation”

✓ located verbatim in the source · abstract, characters 483–525 · show in text ↑
t0

“apple”

✓ located verbatim in the source · title, characters 103–108 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

MdCKX5 silencing enhanced drought tolerance.

Treatment
Silencing of MdCKX5e0
Gene
MdCKX5e0
Outcome
drought tolerancee0
Direction
enhancinge0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“Silencing of MdCKX5 caused CK accumulation thereby enhancing drought tolerance”

✓ located verbatim in the source · abstract, characters 483–561 · show in text ↑
t0

“apple”

✓ located verbatim in the source · title, characters 103–108 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

MdCKX5 silencing increased root vitality.

Treatment
Silencing of MdCKX5e0
Gene
MdCKX5e0
Outcome
root vitalitye0
Direction
increasinge0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“Silencing of MdCKX5 caused CK accumulation thereby enhancing drought tolerance by increasing root vitality”

✓ located verbatim in the source · abstract, characters 483–589 · show in text ↑
t0

“apple”

✓ located verbatim in the source · title, characters 103–108 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

MdCKX5 silencing prevented reactive oxygen species accumulation.

Treatment
Silencing of MdCKX5e0
Gene
MdCKX5e0
Outcome
accumulation of reactive oxygen speciese0
Direction
preventinge0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“Silencing of MdCKX5 caused CK accumulation thereby enhancing drought tolerance by increasing root vitality and preventing the accumulation of reactive oxygen species”

✓ located verbatim in the source · abstract, characters 483–648 · show in text ↑
t0

“apple”

✓ located verbatim in the source · title, characters 103–108 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

MdCKX5 overexpression reduced drought tolerance.

Treatment
overexpression of MdCKX5e0
Gene
MdCKX5e0
Outcome
drought tolerancee0
Direction
reducede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“overexpression of MdCKX5 reduced drought tolerance in apple roots”

✓ located verbatim in the source · abstract, characters 663–728 · show in text ↑
Mechanismobservedoriginal resultmeasured

Claim · model-written summary

MdSPL1 directly binds MdCKX5 promoters and represses expression.

Gene
MdSPL1e0
Outcome
repressing its expressione0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“apple transcription factor MdSPL1 directly binds to the promoters of MdCKX5 thereby repressing its expression”

✓ located verbatim in the source · abstract, characters 820–929 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

MdSPL1 overexpression increased cytokinin content.

Treatment
Overexpression of MdSPL1e0
Gene
MdSPL1e0
Outcome
CK contente0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“Overexpression of MdSPL1 in apple roots increased the CK content”

✓ located verbatim in the source · abstract, characters 931–995 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

MdSPL1 overexpression enhanced drought tolerance.

Treatment
Overexpression of MdSPL1e0
Gene
MdSPL1e0
Outcome
drought tolerancee0
Direction
enhancinge0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“Overexpression of MdSPL1 in apple roots increased the CK content thereby enhancing drought tolerance”

✓ located verbatim in the source · abstract, characters 931–1031 · show in text ↑
Mechanismobservedoriginal resultmeasured

Claim · model-written summary

MdMYB23 interacts with MdSPL1 to alleviate MdCKX5 repression.

Gene
MdMYB23e0
Outcome
repression of MdCKX5 expressione0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“MdMYB23, a positive CK-responsive gene, interacts with MdSPL1 to alleviate the repression of MdCKX5 expression by MdSPL1”

✓ located verbatim in the source · abstract, characters 1064–1184 · show in text ↑
t0

“apple”

✓ located verbatim in the source · title, characters 103–108 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

Drought stress downregulated MdMYB23.

Treatment
drought stresse0
Gene
MdMYB23e0
Outcome
MdMYB23e0
Direction
downregulatede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“drought stress significantly downregulated MdMYB23”

✓ located verbatim in the source · abstract, characters 1198–1248 · show in text ↑
t0

“apple”

✓ located verbatim in the source · title, characters 103–108 · show in text ↑
Observationobservedoriginal resultnegativemeasured

Claim · model-written summary

Adult apple roots lost capacity to activate the MdSPL1-MdCKX5 cascade.

Organ / stage
adult apple rootse0
Outcome
lost the capacity to activate this cascadee0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Malus domestica
Malus domestica · TaxonW.carter · CC0 · Wikimedia Commons ↗

Verbatim

e0

“adult apple roots lost the capacity to activate this cascade”

✓ located verbatim in the source · abstract, characters 1461–1521 · show in text ↑

loss of function · physiology: oxidative · Malus domestica (apple)Positive and negative results

4 papers: 2 positive, 2 negative or no effect

1 paper in the same direction as this paper · 2 in the opposite direction

  1. negative
  2. negative
  3. positive
See all 4 papers →

loss of function · physiology: hormones · Malus domestica (apple)Positive and negative results

5 papers: 4 positive, 1 negative or no effect

3 papers in the same direction as this paper · 1 in the opposite direction

  1. negative
  2. positive
  3. positive

1 paper more in this group

See all 5 papers →

loss of function · tolerance: abiotic · Malus domestica (apple)Positive and negative results

8 papers: 5 positive, 3 negative or no effect

4 papers in the same direction as this paper · 3 in the opposite direction

  1. positive
  2. positive
  3. negative

4 papers more in this group

See all 8 papers →

loss of function · growth · Malus domestica (apple)Positive and negative results

9 papers: 5 positive, 4 negative or no effect

4 papers in the same direction as this paper · 4 in the opposite direction

  1. negative
  2. negative
  3. negative

5 papers more in this group

See all 9 papers →

Claims are grouped by organism, treatment and what was measured or targeted. A paper counts as positive when any of its claims in the group is positive.