Paper · 2026 · Journal of Integrative Agriculture

Heat and drought stress impair nitrogen uptake and allocation in maize by reshaping root morpho-physiology

Ya Guo; Yinglong Chen; Hao Ren; et al.

15
claims
35
verbatim quotes
1
citations
3
research areas
✓
Open access · cc-by-nc-nd

Abstract

Global climate change is increasing the frequency of concurrent heat and drought events, highlighting the urgent need to elucidate their synergistic effects on crop root function and nutrient uptake. We subjected maize plants to normal condition (CK), heat stress (H), drought stress (D), and combined heat and drought stress (HD) at the 12 th fully expanded leaf stage (V12) for five days. The root-to-shoot ratio decreased under H but increased under D. Both single stresses induced “low-cost” root anatomical changes. Under HD, however, these anatomical alterations were most pronounced. They coincided with the strongest oxidative damage, the greatest suppression of root respiration, and the most severe cellular energy deficit. This energy limitation downregulated key nitrogen assimilation enzymes (NR, GS, and GOGAT) and impaired the compensatory upregulation of glutamate dehydrogenase observed under single stresses. Consequently, root nitrogen uptake efficiency declined by 9.0%, 10.4%, and 18.0% under H, D, and HD, respectively. Total plant nitrogen accumulation was lowest under HD, with nitrogen allocation increasingly skewed toward the shoot. Grain yield also lowest under HD. Collectively, these findings demonstrate that combined heat and drought cause oxidative damage, which in turn worsens the energy deficit in roots and suppresses nitrogen assimilation, thereby reducing nitrogen acquisition efficiency in maize.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by-nc-nd. Read at the source ↗

The text the model read, with the source record’s inline markup (italics, sub- and superscripts) rendered and its character entities decoded; quote offsets refer to the raw text.

Claims 15

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Heat decreased maize root-to-shoot ratio.

Taxon
maizeZea mayst0
Treatment
He0
Outcome
root-to-shoot ratioe0
Direction
decreasede0
Comparator
normal condition (CK)e1

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“The root-to-shoot ratio decreased under H but increased under D.”

✓ located verbatim in the source · abstract, characters 391–455 · show in text ↑
e1

“We subjected maize plants to normal condition (CK), heat stress (H), drought stress (D), and combined heat and drought stress (HD) at the 12 th fully expanded leaf stage (V12) for five days.”

✓ located verbatim in the source · abstract, characters 200–390 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Drought increased maize root-to-shoot ratio.

Taxon
maizeZea mayst0
Treatment
De0
Outcome
root-to-shoot ratioe0
Direction
increasede0
Comparator
normal condition (CK)e1

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“The root-to-shoot ratio decreased under H but increased under D.”

✓ located verbatim in the source · abstract, characters 391–455 · show in text ↑
e1

“We subjected maize plants to normal condition (CK), heat stress (H), drought stress (D), and combined heat and drought stress (HD) at the 12 th fully expanded leaf stage (V12) for five days.”

✓ located verbatim in the source · abstract, characters 200–390 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Single stresses induced low-cost root anatomical changes.

Taxon
maizeZea mayst0
Treatment
single stressese0
Outcome
root anatomical changese0

Not reportedComparatorDirection

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Both single stresses induced “low-cost” root anatomical changes.”

✓ located verbatim in the source · abstract, characters 456–520 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Combined stress produced the most pronounced root anatomical changes.

Taxon
maizeZea mayst0
Treatment
HDe0
Outcome
anatomical alterationse0
Direction
most pronouncede0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Under HD, however, these anatomical alterations were most pronounced.”

✓ located verbatim in the source · abstract, characters 521–590 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Combined stress caused the strongest oxidative damage.

Taxon
maizeZea mayst0
Treatment
combined heat and drought stresse1
Outcome
oxidative damagee0
Direction
strongeste0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“They coincided with the strongest oxidative damage, the greatest suppression of root respiration, and the most severe cellular energy deficit.”

✓ located verbatim in the source · abstract, characters 591–733 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
e1

“combined heat and drought stress (HD)”

✓ located verbatim in the source · abstract, characters 293–330 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Combined stress caused the greatest suppression of root respiration.

Taxon
maizeZea mayst0
Treatment
combined heat and drought stresse1
Outcome
root respiratione0
Direction
suppressione0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“They coincided with the strongest oxidative damage, the greatest suppression of root respiration, and the most severe cellular energy deficit.”

✓ located verbatim in the source · abstract, characters 591–733 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
e1

“combined heat and drought stress (HD)”

✓ located verbatim in the source · abstract, characters 293–330 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Combined stress caused the most severe cellular energy deficit.

Taxon
maizeZea mayst0
Treatment
combined heat and drought stresse1
Outcome
cellular energy deficite0
Direction
most severee0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“They coincided with the strongest oxidative damage, the greatest suppression of root respiration, and the most severe cellular energy deficit.”

✓ located verbatim in the source · abstract, characters 591–733 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
e1

“combined heat and drought stress (HD)”

✓ located verbatim in the source · abstract, characters 293–330 · show in text ↑
Mechanismobservedoriginal resultnegativemeasured

Claim · model-written summary

Energy limitation downregulated NR, GS, and GOGAT.

Taxon
maizeZea mayst0
Gene
NR, GS, and GOGATe0
Outcome
key nitrogen assimilation enzymese0
Qualifier
energy limitatione0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“This energy limitation downregulated key nitrogen assimilation enzymes (NR, GS, and GOGAT) and impaired the compensatory upregulation of glutamate dehydrogenase observed under single stresses.”

✓ located verbatim in the source · abstract, characters 734–926 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Mechanismobservedoriginal resultnegativemeasured

Claim · model-written summary

Energy limitation impaired compensatory glutamate dehydrogenase upregulation.

Taxon
maizeZea mayst0
Gene
glutamate dehydrogenasee0
Outcome
compensatory upregulation of glutamate dehydrogenasee0
Qualifier
energy limitatione0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“This energy limitation downregulated key nitrogen assimilation enzymes (NR, GS, and GOGAT) and impaired the compensatory upregulation of glutamate dehydrogenase observed under single stresses.”

✓ located verbatim in the source · abstract, characters 734–926 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

H reduced maize root nitrogen uptake efficiency by 9.0%.

Taxon
maizeZea mayst0
Treatment
He0
Outcome
root nitrogen uptake efficiencye0
Direction
declinede0
Value
9.0%e0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Consequently, root nitrogen uptake efficiency declined by 9.0%, 10.4%, and 18.0% under H, D, and HD, respectively.”

✓ located verbatim in the source · abstract, characters 927–1041 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

D reduced maize root nitrogen uptake efficiency by 10.4%.

Taxon
maizeZea mayst0
Treatment
De0
Outcome
root nitrogen uptake efficiencye0
Direction
declinede0
Value
10.4%e0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Consequently, root nitrogen uptake efficiency declined by 9.0%, 10.4%, and 18.0% under H, D, and HD, respectively.”

✓ located verbatim in the source · abstract, characters 927–1041 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

HD reduced maize root nitrogen uptake efficiency by 18.0%.

Taxon
maizeZea mayst0
Treatment
HDe0
Outcome
root nitrogen uptake efficiencye0
Direction
declinede0
Value
18.0%e0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Consequently, root nitrogen uptake efficiency declined by 9.0%, 10.4%, and 18.0% under H, D, and HD, respectively.”

✓ located verbatim in the source · abstract, characters 927–1041 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Total maize nitrogen accumulation was lowest under combined stress.

Taxon
maizeZea mayst0
Treatment
HDe0
Outcome
Total plant nitrogen accumulatione0
Direction
loweste0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Total plant nitrogen accumulation was lowest under HD, with nitrogen allocation increasingly skewed toward the shoot.”

✓ located verbatim in the source · abstract, characters 1042–1159 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Combined stress increasingly skewed nitrogen allocation toward the shoot.

Taxon
maizeZea mayst0
Treatment
HDe0
Outcome
nitrogen allocatione0
Direction
increasingly skewed toward the shoote0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Total plant nitrogen accumulation was lowest under HD, with nitrogen allocation increasingly skewed toward the shoot.”

✓ located verbatim in the source · abstract, characters 1042–1159 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Maize grain yield was lowest under combined stress.

Taxon
maizeZea mayst0
Treatment
HDe0
Outcome
Grain yielde0
Direction
loweste0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Grain yield also lowest under HD.”

✓ located verbatim in the source · abstract, characters 1160–1193 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 65–70 · show in text ↑

drought · nutrient: content · Zea mays (maize)Results differ

38 papers: 11 positive, 27 negative or no effect

26 papers in the same direction as this paper · 11 in the opposite direction

  1. positive
  2. positive
  3. positive

34 papers more in this group

See all 38 papers →

heat · physiology: respiration · Zea mays (maize)Positive and negative results

2 papers: 1 positive, 1 negative or no effect

1 in the opposite direction

  1. positive
See all 2 papers →

heat · nutrient: content · Zea mays (maize)Positive and negative results

8 papers: 4 positive, 4 negative or no effect

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

  1. positive
  2. positive
  3. positive

4 papers more in this group

See all 8 papers →

heat · physiology: oxidative · Zea mays (maize)Positive and negative results

23 papers: 3 positive, 20 negative or no effect

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

  1. negative
  2. positive
  3. positive

19 papers more in this group

See all 23 papers →

heat · yield · Zea mays (maize)Positive and negative results

85 papers: 17 positive, 68 negative or no effect

67 papers in the same direction as this paper · 17 in the opposite direction

  1. positive
  2. positive
  3. positive

81 papers more in this group

See all 85 papers →

drought · physiology: oxidative · Zea mays (maize)Positive and negative results

96 papers: 8 positive, 88 negative or no effect

87 papers in the same direction as this paper · 8 in the opposite direction

  1. negative
  2. positive
  3. positive

92 papers more in this group

See all 96 papers →

drought · yield · Zea mays (maize)Positive and negative results

353 papers: 53 positive, 300 negative or no effect

299 papers in the same direction as this paper · 53 in the opposite direction

  1. negative
  2. positive
  3. positive

349 papers more in this group

See all 353 papers →

drought · nutrient: efficiency · Zea mays (maize)Results agree

2 papers: 0 positive, 2 negative or no effect

1 paper in the same direction as this paper

  1. negative
See all 2 papers →

8 of 9 groups shown: named compounds and organisms first, then groups whose results differ (checked claim by claim), then those with positive and negative results, then the smaller groups.

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.