Paper · 2026 · BMC Plant Biology

Drill seeding and 5-ALA improve photosynthetic efficiency and ion balance of rice seedlings under salt stress

Shaobiao Duan; Liming Zhao; Renren Jiang; et al.

15
claims
40
verbatim quotes
0
citations
2
research areas
✓
Open access · cc-by-nc-nd

Abstract

Rice (Oryza sativa L.) seedling raising is a vital step in rice cultivation. Salt stress severely impairs the growth of rice seedlings, whereas 5-aminolevulinic acid (5-ALA) plays a crucial role in plant responses to various abiotic stresses. In this study, we combined agronomic practices with chemical regulation. We selected two sowing methods, namely drill sowing and broadcast sowing. Treatments were applied at the three-leaf stage. These included foliar application of 40 mg·L− 1 5-aminolevulinic acid (5-ALA) and application of 0.4% (w/w) NaCl based on dry soil weight. This resulted in a soil electrical conductivity of 5500 ± 50 µS·cm− 1. Three biological replicates were set for each treatment, and sampling was conducted on the 3rd, 6th, and 9th days after salt stress imposition. The results showed that, under salt stress, excessive accumulation of Na+ and Cl− in rice seedling leaves damaged the photosynthetic apparatus and significantly inhibited seedling growth. Exogenous foliar application of 5-ALA promoted the synthesis of photosynthetic pigments by increasing the content of precursors for chloroplast synthesis. Meanwhile, 5-ALA increased K+ and Ca2+ levels significantly and reduced Na+ content by 26.96%. It also enhanced antioxidant enzyme activity and alleviated oxidative damage. As a result, gas exchange and chloroplast fluorescence parameters in seedlings improved, the photosynthetic rate increased by 49.39%, and carbohydrate accumulation was enhanced. Compared with broadcast sowing, drill sowing combined with 5-ALA application resulted in higher levels of chloroplast synthesis precursors and greater carbohydrate accumulation. These reserves provided additional energy for chlorophyll synthesis and for the repair of the photosynthetic apparatus under salt stress. This enhanced peroxide-scavenging capacity, maintained photosynthetic apparatus stability, improved ion homeostasis, and gave drill sowing stronger salt tolerance than broadcast sowing. Taken together, drill sowing and foliar application of 5-ALA significantly maintain the stability of the photosynthetic system and ion homeostasis in rice seedlings under salt stress. This enhances salt tolerance. These findings lay a theoretical foundation for future field trials and the use of saline-alkali soils.

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

Associationobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

Under salt stress, excessive Na+ accumulation significantly inhibited rice seedling growth.

Treatment
salt stresse3
Compound
Na+e1
Outcome
seedling growthe2
Direction
inhibitede2

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e1

“excessive accumulation of Na+”

✓ located verbatim in the source · abstract, characters 837–866 · show in text ↑
e2

“significantly inhibited seedling growth”

✓ located verbatim in the source · abstract, characters 940–979 · show in text ↑
e3

“The results showed that, under salt stress”

✓ located verbatim in the source · abstract, characters 793–835 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Foliar 5-ALA promoted synthesis of photosynthetic pigments in rice seedlings.

Treatment
foliar application of 5-ALAe3
Outcome
synthesis of photosynthetic pigmentse3
Direction
promotede3

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e3

“Exogenous foliar application of 5-ALA promoted the synthesis of photosynthetic pigments by increasing the content of precursors for chloroplast synthesis”

✓ located verbatim in the source · abstract, characters 981–1134 · show in text ↑
Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

Claim · model-written summary

5-ALA significantly increased K+ and Ca2+ levels in rice seedlings under salt stress.

Treatment
5-ALAe4
Outcome
K+ and Ca2+ levelse4
Direction
increasede4

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e4

“Meanwhile, 5-ALA increased K+ and Ca2+ levels significantly and reduced Na+ content by 26.96%”

✓ located verbatim in the source · abstract, characters 1136–1229 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

5-ALA reduced Na+ content in rice seedlings by 26.96%.

Treatment
5-ALAe4
Outcome
Na+ contente4
Direction
reducede4
Value
26.96%e4

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e4

“Meanwhile, 5-ALA increased K+ and Ca2+ levels significantly and reduced Na+ content by 26.96%”

✓ located verbatim in the source · abstract, characters 1136–1229 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

5-ALA enhanced antioxidant enzyme activity in rice seedlings under salt stress.

Treatment
foliar application of 5-ALAe3
Outcome
antioxidant enzyme activitye5
Direction
enhancede5

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e3

“Exogenous foliar application of 5-ALA promoted the synthesis of photosynthetic pigments by increasing the content of precursors for chloroplast synthesis”

✓ located verbatim in the source · abstract, characters 981–1134 · show in text ↑
e5

“It also enhanced antioxidant enzyme activity and alleviated oxidative damage”

✓ located verbatim in the source · abstract, characters 1231–1307 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

5-ALA alleviated oxidative damage in rice seedlings under salt stress.

Treatment
foliar application of 5-ALAe3
Outcome
oxidative damagee5
Direction
alleviatede5

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e3

“Exogenous foliar application of 5-ALA promoted the synthesis of photosynthetic pigments by increasing the content of precursors for chloroplast synthesis”

✓ located verbatim in the source · abstract, characters 981–1134 · show in text ↑
e5

“It also enhanced antioxidant enzyme activity and alleviated oxidative damage”

✓ located verbatim in the source · abstract, characters 1231–1307 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

5-ALA improved gas exchange and chloroplast fluorescence parameters of rice seedlings.

Treatment
foliar application of 5-ALAe3
Outcome
gas exchange and chloroplast fluorescence parameterse6
Direction
improvede6

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e3

“Exogenous foliar application of 5-ALA promoted the synthesis of photosynthetic pigments by increasing the content of precursors for chloroplast synthesis”

✓ located verbatim in the source · abstract, characters 981–1134 · show in text ↑
e6

“As a result, gas exchange and chloroplast fluorescence parameters in seedlings improved, the photosynthetic rate increased by 49.39%, and carbohydrate accumulation was enhanced”

✓ located verbatim in the source · abstract, characters 1309–1485 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

5-ALA increased the photosynthetic rate of rice seedlings by 49.39%.

Treatment
foliar application of 5-ALAe3
Outcome
photosynthetic ratee6
Direction
increasede6
Value
49.39%e6

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e3

“Exogenous foliar application of 5-ALA promoted the synthesis of photosynthetic pigments by increasing the content of precursors for chloroplast synthesis”

✓ located verbatim in the source · abstract, characters 981–1134 · show in text ↑
e6

“As a result, gas exchange and chloroplast fluorescence parameters in seedlings improved, the photosynthetic rate increased by 49.39%, and carbohydrate accumulation was enhanced”

✓ located verbatim in the source · abstract, characters 1309–1485 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

5-ALA enhanced carbohydrate accumulation in rice seedlings.

Treatment
foliar application of 5-ALAe3
Outcome
carbohydrate accumulatione6
Direction
enhancede6

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e3

“Exogenous foliar application of 5-ALA promoted the synthesis of photosynthetic pigments by increasing the content of precursors for chloroplast synthesis”

✓ located verbatim in the source · abstract, characters 981–1134 · show in text ↑
e6

“As a result, gas exchange and chloroplast fluorescence parameters in seedlings improved, the photosynthetic rate increased by 49.39%, and carbohydrate accumulation was enhanced”

✓ located verbatim in the source · abstract, characters 1309–1485 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Drill sowing with 5-ALA gave higher levels of chloroplast synthesis precursors than broadcast sowing.

Treatment
drill sowing combined with 5-ALA applicatione7
Outcome
levels of chloroplast synthesis precursorse7
Direction
highere7
Comparator
broadcast sowinge7

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e7

“Compared with broadcast sowing, drill sowing combined with 5-ALA application resulted in higher levels of chloroplast synthesis precursors and greater carbohydrate accumulation”

✓ located verbatim in the source · abstract, characters 1487–1663 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Drill sowing with 5-ALA gave greater carbohydrate accumulation than broadcast sowing.

Treatment
drill sowing combined with 5-ALA applicatione7
Outcome
carbohydrate accumulatione7
Direction
greatere7
Comparator
broadcast sowinge7

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e7

“Compared with broadcast sowing, drill sowing combined with 5-ALA application resulted in higher levels of chloroplast synthesis precursors and greater carbohydrate accumulation”

✓ located verbatim in the source · abstract, characters 1487–1663 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Drill sowing gave rice stronger salt tolerance than broadcast sowing.

Treatment
drill sowinge0
Outcome
salt tolerancee0
Direction
strongere0
Comparator
broadcast sowinge0

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e0

“This enhanced peroxide-scavenging capacity, maintained photosynthetic apparatus stability, improved ion homeostasis, and gave drill sowing stronger salt tolerance than broadcast sowing”

✓ located verbatim in the source · abstract, characters 1803–1987 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Drill sowing with 5-ALA enhanced peroxide-scavenging capacity of rice seedlings under salt stress.

Treatment
drill sowing combined with 5-ALA applicatione7
Outcome
peroxide-scavenging capacitye0
Direction
enhancede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e7

“Compared with broadcast sowing, drill sowing combined with 5-ALA application resulted in higher levels of chloroplast synthesis precursors and greater carbohydrate accumulation”

✓ located verbatim in the source · abstract, characters 1487–1663 · show in text ↑
e0

“This enhanced peroxide-scavenging capacity, maintained photosynthetic apparatus stability, improved ion homeostasis, and gave drill sowing stronger salt tolerance than broadcast sowing”

✓ located verbatim in the source · abstract, characters 1803–1987 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Drill sowing with 5-ALA maintained photosynthetic apparatus stability in rice seedlings under salt stress.

Treatment
drill sowing combined with 5-ALA applicatione7
Outcome
photosynthetic apparatus stabilitye0
Direction
maintainede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e7

“Compared with broadcast sowing, drill sowing combined with 5-ALA application resulted in higher levels of chloroplast synthesis precursors and greater carbohydrate accumulation”

✓ located verbatim in the source · abstract, characters 1487–1663 · show in text ↑
e0

“This enhanced peroxide-scavenging capacity, maintained photosynthetic apparatus stability, improved ion homeostasis, and gave drill sowing stronger salt tolerance than broadcast sowing”

✓ located verbatim in the source · abstract, characters 1803–1987 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Drill sowing with 5-ALA improved ion homeostasis in rice seedlings under salt stress.

Treatment
drill sowing combined with 5-ALA applicatione7
Outcome
ion homeostasise0
Direction
improvede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice seedlings under salt stress”

✓ located verbatim in the source · title, characters 77–109 · show in text ↑
e7

“Compared with broadcast sowing, drill sowing combined with 5-ALA application resulted in higher levels of chloroplast synthesis precursors and greater carbohydrate accumulation”

✓ located verbatim in the source · abstract, characters 1487–1663 · show in text ↑
e0

“This enhanced peroxide-scavenging capacity, maintained photosynthetic apparatus stability, improved ion homeostasis, and gave drill sowing stronger salt tolerance than broadcast sowing”

✓ located verbatim in the source · abstract, characters 1803–1987 · show in text ↑

5-aminolevulinic acid · physiology: oxidative · Oryza sativa (rice)Positive and negative results

4 papers: 3 positive, 1 negative or no effect

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

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

5-aminolevulinic acid · physiology: photosynthesis · Oryza sativa (rice)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 →

5-aminolevulinic acid · soil: biology · Oryza sativa (rice)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 →

application · soil: biology · Oryza sativa (rice)Positive and negative results

15 papers: 12 positive, 3 negative or no effect

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

  1. negative
  2. no effect
  3. no effect

11 papers more in this group

See all 15 papers →

transplanting · quality: sugar starch · Oryza sativa (rice)Positive and negative results

19 papers: 15 positive, 4 negative or no effect

14 papers in the same direction as this paper · 3 in the opposite direction · 1 reporting no difference from an alternative

  1. negative
  2. negative
  3. no effect

15 papers more in this group

See all 19 papers →

transplanting · physiology: photosynthesis · Oryza sativa (rice)Positive and negative results

21 papers: 15 positive, 6 negative or no effect

14 papers in the same direction as this paper · 6 in the opposite direction

  1. negative
  2. no effect
  3. negative

17 papers more in this group

See all 21 papers →

application · quality: sugar starch · Oryza sativa (rice)Positive and negative results

24 papers: 21 positive, 3 negative or no effect

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

  1. no effect
  2. negative
  3. no effect

20 papers more in this group

See all 24 papers →

application · physiology: oxidative · Oryza sativa (rice)Positive and negative results

36 papers: 35 positive, 1 negative or no effect

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

  1. positive
  2. no effect
  3. positive

32 papers more in this group

See all 36 papers →

8 of 11 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.