Paper · 2025 · Ecotoxicology and Environmental Safety

Exogenous proline enhances salt acclimation in soybean seedlings: Modifying physicochemical properties and controlling proline metabolism through the ornithine-glutamate dual pathway

Lei Yan; Mu Lu; Muhammad Riaz; et al.

14
claims
28
verbatim quotes
24
citations
1
research areas
✓
Open access · cc-by-nc-nd

Abstract

Soil salinization has emerged as a major factor negatively affecting soil quality and plant productivity. Proline, functioning as an osmotic regulator, has been proposed as an effective strategy for enhancing plant tolerance to salt stress. This study aimed to investigate the effects of exogenous proline on salt tolerance in soybeans. A hydroponic experiment was conducted with different salt treatments (without NaCl, -NaCl; with 100 mM NaCl, +NaCl) and with or without 150 mM proline (+Pro, -Pro). The results showed that proline application alleviated salt stress-induced reductions in plant growth, photosynthetic parameters, and chlorophyll content while aiding recovery from leaf chlorosis. Proline treatment improved ion homeostasis by reducing Na + levels and increasing K + and Ca 2+ contents in the leaves. Salt stress increased malondialdehyde (MDA) and reactive oxygen species (ROS) levels, along with leaf peroxidase (POD) and catalase (CAT) activities, while decreasing superoxide dismutase (SOD) activity. Moreover, salt stress obviously enhanced proline accumulation, accompanied by increases in glutamate (Glu), glutamate-1-semialdehyde (GSA), and pyrroline-5-carboxylate (P5C) content, as well as the activities of pyrroline-5-carboxylate synthase (P5CS) and pyrroline-5-carboxylate reductase (P5CR) in the glutamate pathway, while reducing proline dehydrogenase (ProDH) activity. Exogenous proline treatment further elevated proline content and increased key substances and enzyme activities in both the glutamate (Glu and P5C content, P5CS and P5CR activity) and ornithine (Orn content and OAT activity) pathways while also reducing ProDH activity. Collectively, our results revealed that exogenous proline contributed to an attenuation of salt stress in soybeans by regulating both the glutamate and ornithine pathways to stimulate endogenous proline accumulation, mediate Na + /K + homeostasis, and inhibit oxidative damage. • The proline (Pro) metabolism is different between +NaCl and +NaCl +Pro treatments. • Pro improves salt-tolerance by improving the antioxidant defense system. • Pro helped in optimizing Na + /K + homeostasis to aid in photosynthetic protection.

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 14

Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Proline alleviated salt-induced reductions in growth, photosynthesis, and chlorophyll.

Taxon
soybean seedlingsGlycine maxt0
Treatment
proline applicatione0
Outcome
plant growth, photosynthetic parameters, and chlorophyll contente0
Direction
alleviatede0
Comparator
salt stresse0

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“proline application alleviated salt stress-induced reductions in plant growth, photosynthetic parameters, and chlorophyll content”

✓ located verbatim in the source · abstract, characters 526–655 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

Proline aided recovery from leaf chlorosis.

Taxon
soybean seedlingsGlycine maxt0
Outcome
recovery from leaf chlorosise0

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“while aiding recovery from leaf chlorosis”

✓ located verbatim in the source · abstract, characters 656–697 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Proline lowered leaf sodium.

Taxon
soybean seedlingsGlycine maxt0
Organ / stage
leavese0
Treatment
Proline treatmente0
Outcome
Na + levelse0
Direction
reducinge0

Not reportedComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“Proline treatment improved ion homeostasis by reducing Na + levels and increasing K + and Ca 2+ contents in the leaves.”

✓ located verbatim in the source · abstract, characters 699–818 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Proline increased leaf potassium and calcium contents.

Taxon
soybean seedlingsGlycine maxt0
Organ / stage
leavese0
Treatment
Proline treatmente0
Outcome
K + and Ca 2+ contentse0
Direction
increasinge0

Not reportedComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“Proline treatment improved ion homeostasis by reducing Na + levels and increasing K + and Ca 2+ contents in the leaves.”

✓ located verbatim in the source · abstract, characters 699–818 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Salt stress increased MDA and ROS levels.

Taxon
soybean seedlingsGlycine maxt0
Treatment
Salt stresse0
Outcome
malondialdehyde (MDA) and reactive oxygen species (ROS) levelse0
Direction
increasede0

Not reportedComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“Salt stress increased malondialdehyde (MDA) and reactive oxygen species (ROS) levels, along with leaf peroxidase (POD) and catalase (CAT) activities, while decreasing superoxide dismutase (SOD) activity.”

✓ located verbatim in the source · abstract, characters 819–1022 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Salt stress increased leaf POD and CAT activities.

Taxon
soybean seedlingsGlycine maxt0
Treatment
Salt stresse0
Outcome
leaf peroxidase (POD) and catalase (CAT) activitiese0
Direction
increasede0

Not reportedComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“Salt stress increased malondialdehyde (MDA) and reactive oxygen species (ROS) levels, along with leaf peroxidase (POD) and catalase (CAT) activities, while decreasing superoxide dismutase (SOD) activity.”

✓ located verbatim in the source · abstract, characters 819–1022 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Salt stress decreased SOD activity.

Taxon
soybean seedlingsGlycine maxt0
Treatment
Salt stresse0
Outcome
superoxide dismutase (SOD) activitye0
Direction
decreasinge0

Not reportedComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“Salt stress increased malondialdehyde (MDA) and reactive oxygen species (ROS) levels, along with leaf peroxidase (POD) and catalase (CAT) activities, while decreasing superoxide dismutase (SOD) activity.”

✓ located verbatim in the source · abstract, characters 819–1022 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Salt stress raised proline, Glu, GSA, and P5C content.

Taxon
soybean seedlingsGlycine maxt0
Treatment
salt stresse0
Outcome
proline accumulation, accompanied by increases in glutamate (Glu), glutamate-1-semialdehyde (GSA), and pyrroline-5-carboxylate (P5C) contente0
Direction
enhancede0

Not reportedComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“salt stress obviously enhanced proline accumulation, accompanied by increases in glutamate (Glu), glutamate-1-semialdehyde (GSA), and pyrroline-5-carboxylate (P5C) content”

✓ located verbatim in the source · abstract, characters 1033–1204 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Salt stress increased P5CS and P5CR activities.

Taxon
soybean seedlingsGlycine maxt0
Outcome
activities of pyrroline-5-carboxylate synthase (P5CS) and pyrroline-5-carboxylate reductase (P5CR)e0
Direction
increaseinferrede0t0

Not reportedTreatmentComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“the activities of pyrroline-5-carboxylate synthase (P5CS) and pyrroline-5-carboxylate reductase (P5CR) in the glutamate pathway, while reducing proline dehydrogenase (ProDH) activity.”

✓ located verbatim in the source · abstract, characters 1217–1400 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Salt stress reduced ProDH activity.

Taxon
soybean seedlingsGlycine maxt0
Outcome
proline dehydrogenase (ProDH) activitye0
Direction
reducinge0

Not reportedTreatmentComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“while reducing proline dehydrogenase (ProDH) activity.”

✓ located verbatim in the source · abstract, characters 1346–1400 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous proline further increased endogenous proline content.

Taxon
soybean seedlingsGlycine maxt0
Treatment
Exogenous proline treatmente0
Outcome
proline contente0
Direction
elevatede0

Not reportedComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“Exogenous proline treatment further elevated proline content and increased key substances and enzyme activities in both the glutamate (Glu and P5C content, P5CS and P5CR activity) and ornithine (Orn content and OAT activity) pathways while also reducing ProDH activity.”

✓ located verbatim in the source · abstract, characters 1401–1670 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous proline increased Glu and P5C contents and P5CS and P5CR activities.

Taxon
soybean seedlingsGlycine maxt0
Treatment
Exogenous proline treatmente0
Outcome
Glu and P5C content, P5CS and P5CR activitye0
Direction
increasede0

Not reportedComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“Exogenous proline treatment further elevated proline content and increased key substances and enzyme activities in both the glutamate (Glu and P5C content, P5CS and P5CR activity) and ornithine (Orn content and OAT activity) pathways while also reducing ProDH activity.”

✓ located verbatim in the source · abstract, characters 1401–1670 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous proline increased Orn content and OAT activity.

Taxon
soybean seedlingsGlycine maxt0
Treatment
Exogenous proline treatmente0
Outcome
Orn content and OAT activitye0
Direction
increasede0

Not reportedComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“Exogenous proline treatment further elevated proline content and increased key substances and enzyme activities in both the glutamate (Glu and P5C content, P5CS and P5CR activity) and ornithine (Orn content and OAT activity) pathways while also reducing ProDH activity.”

✓ located verbatim in the source · abstract, characters 1401–1670 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous proline reduced ProDH activity.

Taxon
soybean seedlingsGlycine maxt0
Treatment
Exogenous proline treatmente0
Outcome
ProDH activitye0
Direction
reducinge0

Not reportedComparator

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

Glycine max
Glycine max · TaxonPublic domain · Wikimedia Commons ↗

Verbatim

e0

“Exogenous proline treatment further elevated proline content and increased key substances and enzyme activities in both the glutamate (Glu and P5C content, P5CS and P5CR activity) and ornithine (Orn content and OAT activity) pathways while also reducing ProDH activity.”

✓ located verbatim in the source · abstract, characters 1401–1670 · show in text ↑
t0

“soybean seedlings”

✓ located verbatim in the source · title, characters 47–64 · show in text ↑

proline · nutrient: content · Glycine max (soybean)Positive and negative results

2 papers: 1 positive, 1 negative or no effect

1 in the opposite direction

  1. no effect
See all 2 papers →

proline · physiology: photosynthesis · Glycine max (soybean)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 →

proline · physiology: osmolytes · Glycine max (soybean)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 →

proline · molecular: metabolites · Glycine max (soybean)Results agree

3 papers: 3 positive, 0 negative or no effect

2 papers in the same direction as this paper

  1. positive
  2. positive
See all 3 papers →

salinity · physiology: osmolytes · Glycine max (soybean)Results differ

11 papers: 6 positive, 5 negative or no effect

5 papers in the same direction as this paper · 5 in the opposite direction

  1. negative
  2. negative
  3. negative

7 papers more in this group

See all 11 papers →

salinity · physiology: antioxidant enzymes · Glycine max (soybean)Results differ

20 papers: 10 positive, 10 negative or no effect

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

  1. positive
  2. positive
  3. positive

16 papers more in this group

See all 20 papers →

salinity · physiology: oxidative · Glycine max (soybean)Results differ

41 papers: 4 positive, 37 negative or no effect

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

  1. positive
  2. positive
  3. positive

37 papers more in this group

See all 41 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.