Abstract

Abiotic stresses, including nitrogen stress (NS), can hamper photosynthesis and cause oxidative damage to plants. Upregulation of the antioxidative defense system and photosynthesis induced by exogenous glycinebetaine (GB) and humic acid (HA) can mitigate the inhibitory effects of NS on plants. In the present investigation, the beneficial effects of exogenously applied GB and HA were examined on growth, leaf N status, photosynthesis, lipid peroxidation, and activities of some key antioxidant enzymes in the seedlings of maize cv. Zhengdan 958 (ZD958) exposed to NS. NS caused a significant reduction in total dry matter of seedlings of ZD958, but both GB and HA proved effective in mitigating this inhibition, hence, the beneficial effects of GB being more pronounced than those of HA. NS led to a considerable decrease in leaf total N and endogenous GB contents, stomatal conductance (g s), net photosynthetic rate (P n), intercellular CO2 concentration (C i), and activities of two key C4 photosynthesis enzymes phosphoenolpyruvate carboxylase (PEPCase) and ribulose-1,5-bisphosphate carboxylase (RuBPCase) as well as of superoxide dismutase (SOD) and peroxidase (POD). This treatment caused an increase in lipid peroxidation, but showed no effect on POD activity. Exogenous application of varying doses of GB resulted in a decrease in lipid peroxidation and C i, and an increase in leaf total N and endogenous glycinebetaine (EGB) content, P n, and activities of RuBPCase, PEPCase, SOD, and catalase (CAT) under NS. In contrast, application of different doses of HA resulted in a decrease in lipid peroxidation, an increase in P n, g s, and C i as well as SOD, CAT, and POD activities without increasing leaf total N and EGB content, and enhanced RuBPCase and PEPCase activities. The present study suggests that exogenous application of GB and HA can induce tolerance in maize plants to NS, but through the regulation of different mechanisms.

Abstract from the paper’s OpenAlex record, shown for indexing and research. Copyright remains with the authors or the publisher. 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 17

Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

Nitrogen stress significantly reduced maize seedling dry matter.

Taxon
ZD958e0
Treatment
NSe0
Outcome
total dry mattere0
Direction
reductione0

Not reportedComparator

Verbatim

e0

“NS caused a significant reduction in total dry matter of seedlings of ZD958”

✓ located verbatim in the source · abstract, characters 571–646 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Both glycinebetaine and humic acid mitigated nitrogen-stress inhibition of seedling dry matter, with glycinebetaine more effective.

Taxon
maizeZea mayst0
Treatment
GB and HAe0
Outcome
this inhibitione0
Direction
mitigatinge0
Comparator
HAe0

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 GB and HA proved effective in mitigating this inhibition, hence, the beneficial effects of GB being more pronounced than those of HA”

✓ located verbatim in the source · abstract, characters 652–789 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Nitrogen stress decreased leaf nitrogen and endogenous glycinebetaine content.

Taxon
maizeZea mayst0
Treatment
NSe0
Outcome
leaf total N and endogenous GB contentse0
Direction
decreasee0

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

“NS led to a considerable decrease in leaf total N and endogenous GB contents”

✓ located verbatim in the source · abstract, characters 791–867 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Nitrogen stress decreased stomatal conductance, net photosynthetic rate and intercellular CO2 concentration.

Taxon
maizeZea mayst0
Treatment
NSe0
Outcome
stomatal conductance (g s), net photosynthetic rate (P n), intercellular CO2 concentration (C i)e0
Direction
decreasee0

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

“NS led to a considerable decrease in leaf total N and endogenous GB contents, stomatal conductance (g s), net photosynthetic rate (P n), intercellular CO2 concentration (C i)”

✓ located verbatim in the source · abstract, characters 791–965 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Nitrogen stress decreased PEPCase and RuBPCase activities.

Taxon
maizeZea mayst0
Treatment
NSe0
Outcome
activities of two key C4 photosynthesis enzymes phosphoenolpyruvate carboxylase (PEPCase) and ribulose-1,5-bisphosphate carboxylase (RuBPCase)e0
Direction
decreasee0

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

“NS led to a considerable decrease in leaf total N and endogenous GB contents, stomatal conductance (g s), net photosynthetic rate (P n), intercellular CO2 concentration (C i), and activities of two key C4 photosynthesis enzymes phosphoenolpyruvate carboxylase (PEPCase) and ribulose-1,5-bisphosphate carboxylase (RuBPCase)”

✓ located verbatim in the source · abstract, characters 791–1113 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Nitrogen stress decreased superoxide dismutase activity.

Taxon
maizeZea mayst0
Treatment
nitrogen stresst1
Outcome
superoxide dismutase (SOD)e0

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

“as well as of superoxide dismutase (SOD) and peroxidase (POD)”

✓ located verbatim in the source · abstract, characters 1114–1175 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Nitrogen stress increased lipid peroxidation.

Taxon
maizeZea mayst0
Treatment
nitrogen stresst1
Outcome
lipid peroxidatione0
Direction
increasee0

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

“This treatment caused an increase in lipid peroxidation”

✓ located verbatim in the source · abstract, characters 1177–1232 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
Intervention effectobservedoriginal resultno effectmeasured

Claim · model-written summary

Nitrogen stress had no reported effect on POD activity.

Taxon
maizeZea mayst0
Treatment
nitrogen stresst1
Outcome
POD activitye0
Direction
no effecte0

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

“but showed no effect on POD activity”

✓ located verbatim in the source · abstract, characters 1234–1270 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Glycinebetaine lowered lipid peroxidation and intercellular CO2 under nitrogen stress.

Taxon
maizeZea mayst0
Treatment
Exogenous application of varying doses of GBe0
Outcome
lipid peroxidation and C ie0
Direction
decreasee0

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

“Exogenous application of varying doses of GB resulted in a decrease in lipid peroxidation and C i”

✓ located verbatim in the source · abstract, characters 1272–1369 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Glycinebetaine increased leaf total nitrogen, endogenous glycinebetaine and photosynthetic rate under nitrogen stress.

Taxon
maizeZea mayst0
Treatment
GBe1
Outcome
leaf total N and endogenous glycinebetaine (EGB) content, P ne0
Direction
increasee0

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

“an increase in leaf total N and endogenous glycinebetaine (EGB) content, P n”

✓ located verbatim in the source · abstract, characters 1375–1451 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
e1

“Exogenous application of varying doses of GB”

✓ located verbatim in the source · abstract, characters 1272–1316 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Glycinebetaine increased RuBPCase, PEPCase and SOD activities under nitrogen stress.

Taxon
maizeZea mayst0
Treatment
GBe1
Outcome
activities of RuBPCase, PEPCase, SODe0

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

“activities of RuBPCase, PEPCase, SOD, and catalase (CAT) under NS”

✓ located verbatim in the source · abstract, characters 1457–1522 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
e1

“Exogenous application of varying doses of GB”

✓ located verbatim in the source · abstract, characters 1272–1316 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Glycinebetaine increased catalase activity under nitrogen stress.

Taxon
maizeZea mayst0
Treatment
GBe1
Outcome
catalase (CAT)e0

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

“activities of RuBPCase, PEPCase, SOD, and catalase (CAT) under NS”

✓ located verbatim in the source · abstract, characters 1457–1522 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
e1

“Exogenous application of varying doses of GB”

✓ located verbatim in the source · abstract, characters 1272–1316 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Humic acid decreased lipid peroxidation under nitrogen stress.

Taxon
maizeZea mayst0
Treatment
application of different doses of HAe0
Outcome
lipid peroxidatione0
Direction
decreasee0

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

“application of different doses of HA resulted in a decrease in lipid peroxidation”

✓ located verbatim in the source · abstract, characters 1537–1618 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Humic acid increased photosynthetic rate, stomatal conductance and intercellular CO2 under nitrogen stress.

Taxon
maizeZea mayst0
Treatment
HAe1
Outcome
P n, g s, and C ie0
Direction
increasee0

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

“an increase in P n, g s, and C i”

✓ located verbatim in the source · abstract, characters 1620–1652 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
e1

“application of different doses of HA”

✓ located verbatim in the source · abstract, characters 1537–1573 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Humic acid increased SOD, CAT and POD activities under nitrogen stress.

Taxon
maizeZea mayst0
Treatment
HAe1
Outcome
SOD, CAT, and POD activitiese0

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

“SOD, CAT, and POD activities without increasing leaf total N and EGB content”

✓ located verbatim in the source · abstract, characters 1664–1740 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
e1

“application of different doses of HA”

✓ located verbatim in the source · abstract, characters 1537–1573 · show in text ↑
Intervention effectobservedoriginal resultno effectmeasured

Claim · model-written summary

Humic acid did not increase leaf nitrogen or endogenous glycinebetaine content.

Taxon
maizeZea mayst0
Treatment
HAe1
Outcome
leaf total N and EGB contente0
Direction
without increasinge0

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

“without increasing leaf total N and EGB content”

✓ located verbatim in the source · abstract, characters 1693–1740 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
e1

“application of different doses of HA”

✓ located verbatim in the source · abstract, characters 1537–1573 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Humic acid enhanced RuBPCase and PEPCase activities.

Taxon
maizeZea mayst0
Treatment
HAe1
Outcome
RuBPCase and PEPCase activitiese0
Direction
enhancede0

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

“enhanced RuBPCase and PEPCase activities”

✓ located verbatim in the source · abstract, characters 1746–1786 · show in text ↑
t0

“Exogenous glycinebetaine and humic acid improve growth, nitrogen status, photosynthesis, and antioxidant defense system and confer tolerance to nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 0–178 · show in text ↑
t1

“nitrogen stress in maize seedlings”

✓ located verbatim in the source · title, characters 144–178 · show in text ↑
e1

“application of different doses of HA”

✓ located verbatim in the source · abstract, characters 1537–1573 · show in text ↑

glycine betaine · disease · Zea mays (maize)Results agree

2 papers: 2 positive, 0 negative or no effect

1 paper in the same direction as this paper

  1. positive
See all 2 papers →

glycine betaine · physiology: photosynthesis · Zea mays (maize)Results agree

5 papers: 5 positive, 0 negative or no effect

4 papers in the same direction as this paper

  1. positive
  2. positive
  3. positive

1 paper more in this group

See all 5 papers →

glycine betaine · physiology: antioxidant enzymes · Zea mays (maize)Results agree

8 papers: 8 positive, 0 negative or no effect

7 papers in the same direction as this paper

  1. positive
  2. positive
  3. positive

4 papers more in this group

See all 8 papers →

glycine betaine · physiology: oxidative · Zea mays (maize)Results agree

15 papers: 15 positive, 0 negative or no effect

14 papers in the same direction as this paper

  1. positive
  2. positive
  3. positive

11 papers more in this group

See all 15 papers →

other · disease · Zea mays (maize)Positive and negative results

12 papers: 11 positive, 1 negative or no effect

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

  1. no effect
  2. positive
  3. positive

8 papers more in this group

See all 12 papers →

nitrogen · physiology: antioxidant enzymes · Zea mays (maize)Positive and negative results

17 papers: 14 positive, 3 negative or no effect

2 papers in the same direction as this paper · 14 in the opposite direction

  1. positive
  2. positive
  3. positive

13 papers more in this group

See all 17 papers →

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

17 papers: 16 positive, 1 negative or no effect

16 in the opposite direction

  1. positive
  2. positive
  3. positive

13 papers more in this group

See all 17 papers →

other · molecular: metabolites · Zea mays (maize)Positive and negative results

39 papers: 37 positive, 2 negative or no effect

36 papers in the same direction as this paper · 2 in the opposite direction

  1. no effect
  2. no effect
  3. positive

35 papers more in this group

See all 39 papers →

8 of 14 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.