Paper · 2026 · Plants

Humic Acid Modulates Photosynthetic Responses to PEG-Induced Drought in Ocimum basilicum L.

Martin A. Stefanov; Georgi D. Rashkov; Preslava B. Borisova; et al.

20
claims
49
verbatim quotes
0
citations
2
research areas
✓
Open access · cc-by

Abstract

Drought is a major environmental constraint that disrupts photosynthetic processes. This study investigated the effects of foliar-applied commercial humic acid (HA) at different concentrations (1, 3 and 5 mg/mL) on the photosynthetic apparatus of sweet basil (Ocimum basilicum L. Italiano classico) under PEG-induced stress. The responses of the photosynthetic machinery were evaluated using chlorophyll a fluorescence analyses (JIP-test and PAM), leaf pigment composition, and assessments of membrane integrity. Drought stress caused pronounced alterations on both the donor and acceptor sides of photosystem II (PSII), including impaired QA− reoxidation, reduced open PSII reaction centers (qP), diminished electron transport (ETo/RC, REo/RC), and substantial declines in performance indices (PIABS, PItotal). Energy dissipation increased (DI0/RC), with regulated energy losses (ΦNPQ) rising more strongly than non-regulated losses (ΦNO). Drought also elevated oxidative stress markers (MDA and H2O2), leading to enhanced membrane injury. Among the tested concentrations, 5 mg/mL HA provided the most effective protection against drought stress. This treatment mitigated PEG-induced damage on both PSII donor and acceptor sides and increased the proportion of open reaction centers (qP). Improved PSII photochemistry corresponded with more efficient QA− reoxidation, facilitated its interaction with plastoquinone, and caused the overall stabilization of photosynthetic functions under drought. The protective effects of HA were also evident for both PSI subpopulations. The enhanced tolerance was associated with the activation of antioxidant enzymes (CAT, SOD, APX) and the increased levels of anthocyanins and total phenolic content (TPC). In contrast, lower HA concentrations (1 and 3 mg/mL) provided insufficient protection. This study clearly demonstrates that HA enhances drought tolerance in basil in a concentration-dependent manner by protecting the structural and functional integrity of the photosynthetic apparatus, supporting its potential use as a foliar treatment to improve crop resilience under water-limited conditions.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by. 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 20

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought impaired PSII donor and acceptor function, including QA− reoxidation.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
Drought stresse0
Outcome
QA− reoxidatione0
Direction
impairede0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress caused pronounced alterations on both the donor and acceptor sides of photosystem II (PSII), including impaired QA− reoxidation”

✓ located verbatim in the source · abstract, characters 513–655 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought reduced open PSII centers.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
Drought stresse2
Outcome
open PSII reaction centers (qP)e0
Direction
reducede0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“reduced open PSII reaction centers (qP)”

✓ located verbatim in the source · abstract, characters 657–696 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
e2

“Drought stress caused pronounced alterations”

✓ located verbatim in the source · abstract, characters 513–557 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought diminished electron transport.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
Drought stresse2
Outcome
electron transport (ETo/RC, REo/RC)e0
Direction
diminishede0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“diminished electron transport (ETo/RC, REo/RC)”

✓ located verbatim in the source · abstract, characters 698–744 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
e2

“Drought stress caused pronounced alterations”

✓ located verbatim in the source · abstract, characters 513–557 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought reduced PSII performance indices.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
Drought stresse2
Outcome
performance indices (PIABS, PItotal)e0
Direction
declinese0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“substantial declines in performance indices (PIABS, PItotal)”

✓ located verbatim in the source · abstract, characters 750–810 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
e2

“Drought stress caused pronounced alterations”

✓ located verbatim in the source · abstract, characters 513–557 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought increased energy dissipation.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Outcome
Energy dissipatione0
Direction
increasede0

Not reportedTreatmentComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Energy dissipation increased (DI0/RC)”

✓ located verbatim in the source · abstract, characters 812–849 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Regulated energy losses rose more strongly than non-regulated losses.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Outcome
regulated energy losses (ΦNPQ)e0
Direction
rising more stronglye0
Comparator
non-regulated losses (ΦNO)e0

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“regulated energy losses (ΦNPQ) rising more strongly than non-regulated losses (ΦNO)”

✓ located verbatim in the source · abstract, characters 856–939 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought elevated MDA and H2O2 oxidative-stress markers.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
Droughte0
Outcome
oxidative stress markers (MDA and H2O2)e0
Direction
elevatede0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought also elevated oxidative stress markers (MDA and H2O2)”

✓ located verbatim in the source · abstract, characters 941–1002 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought enhanced membrane injury.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
Droughte2
Outcome
membrane injurye0
Direction
enhancede0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“leading to enhanced membrane injury”

✓ located verbatim in the source · abstract, characters 1004–1039 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
e2

“Drought also elevated oxidative stress markers”

✓ located verbatim in the source · abstract, characters 941–987 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

5 mg/mL humic acid provided the strongest drought protection.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
5 mg/mL HAe0
Outcome
protection against drought stresse0
Direction
most effectivee0
Comparator
tested concentrationse0

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Among the tested concentrations, 5 mg/mL HA provided the most effective protection against drought stress”

✓ located verbatim in the source · abstract, characters 1041–1146 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

5 mg/mL humic acid mitigated drought injury to PSII donor and acceptor sides.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume2
Treatment
5 mg/mL HAe1
Outcome
PEG-induced damage on both PSII donor and acceptor sidese0
Direction
mitigatede0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“This treatment mitigated PEG-induced damage on both PSII donor and acceptor sides”

✓ located verbatim in the source · abstract, characters 1148–1229 · show in text ↑
e1

“5 mg/mL HA”

✓ located verbatim in the source · abstract, characters 1074–1084 · show in text ↑
e2

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

5 mg/mL humic acid increased open reaction centers.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume2
Treatment
5 mg/mL HAe1
Outcome
proportion of open reaction centers (qP)e0
Direction
increasede0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“increased the proportion of open reaction centers (qP)”

✓ located verbatim in the source · abstract, characters 1234–1288 · show in text ↑
e1

“5 mg/mL HA”

✓ located verbatim in the source · abstract, characters 1074–1084 · show in text ↑
e2

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Improved PSII photochemistry accompanied more efficient QA− reoxidation.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
5 mg/mL HAe2
Outcome
QA− reoxidatione0
Direction
more efficiente0
Qualifier
Improved PSII photochemistrye0

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Improved PSII photochemistry corresponded with more efficient QA− reoxidation”

✓ located verbatim in the source · abstract, characters 1290–1367 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
e2

“5 mg/mL HA”

✓ located verbatim in the source · abstract, characters 1074–1084 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

Improved PSII photochemistry facilitated QA− interaction with plastoquinone.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume2
Outcome
interaction with plastoquinonee0
Qualifier
Improved PSII photochemistrye1

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“facilitated its interaction with plastoquinone”

✓ located verbatim in the source · abstract, characters 1369–1415 · show in text ↑
e1

“Improved PSII photochemistry corresponded with more efficient QA− reoxidation”

✓ located verbatim in the source · abstract, characters 1290–1367 · show in text ↑
e2

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

Improved PSII photochemistry stabilized photosynthetic function under drought.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume2
Outcome
stabilization of photosynthetic functions under droughte0
Qualifier
Improved PSII photochemistrye1

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“caused the overall stabilization of photosynthetic functions under drought”

✓ located verbatim in the source · abstract, characters 1421–1495 · show in text ↑
e1

“Improved PSII photochemistry”

✓ located verbatim in the source · abstract, characters 1290–1318 · show in text ↑
e2

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Humic acid protected both PSI subpopulations.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
HAe0
Outcome
protective effectse0
Direction
protectivee0
Qualifier
both PSI subpopulationse0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“The protective effects of HA were also evident for both PSI subpopulations”

✓ located verbatim in the source · abstract, characters 1497–1571 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Enhanced tolerance was associated with CAT, SOD, and APX antioxidant enzyme activation.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Gene
antioxidant enzymes (CAT, SOD, APX)e0
Outcome
enhanced tolerancee0
Direction
enhancede0

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“The enhanced tolerance was associated with the activation of antioxidant enzymes (CAT, SOD, APX)”

✓ located verbatim in the source · abstract, characters 1573–1669 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Enhanced tolerance was associated with increased anthocyanin and phenolic content.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Compound
anthocyaninse0
Outcome
enhanced tolerancee0
Direction
enhancede0
Qualifier
increased levels of anthocyanins and total phenolic content (TPC)e0

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“The enhanced tolerance was associated with the activation of antioxidant enzymes (CAT, SOD, APX) and the increased levels of anthocyanins and total phenolic content (TPC)”

✓ located verbatim in the source · abstract, characters 1573–1743 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

The 1 mg/mL humic acid dose provided insufficient drought protection.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
1e0
Outcome
insufficient protectione0
Direction
insufficiente0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“lower HA concentrations (1 and 3 mg/mL) provided insufficient protection”

✓ located verbatim in the source · abstract, characters 1758–1830 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

The 3 mg/mL humic acid dose provided insufficient drought protection.

Taxon
sweet basil (Ocimum basilicum L. Italiano classico)Ocimum basilicume1
Treatment
3 mg/mLe0
Outcome
insufficient protectione0
Direction
insufficiente0

Not reportedComparator

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“lower HA concentrations (1 and 3 mg/mL) provided insufficient protection”

✓ located verbatim in the source · abstract, characters 1758–1830 · show in text ↑
e1

“sweet basil (Ocimum basilicum L. Italiano classico)”

✓ located verbatim in the source · abstract, characters 247–298 · show in text ↑
Recommendationauthor interpretationoriginal resultpositivebasis n/a

Claim · model-written summary

Authors propose foliar humic acid to improve basil resilience under water limitation.

Outcome
potential use as a foliar treatment to improve crop resilience under water-limited conditionse0

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

Ocimum basilicum
Ocimum basilicum · TaxonCastielli · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“supporting its potential use as a foliar treatment to improve crop resilience under water-limited conditions”

✓ located verbatim in the source · abstract, characters 2031–2139 · show in text ↑
e1

“enhances drought tolerance in basil”

✓ located verbatim in the source · abstract, characters 1872–1907 · show in text ↑

other · physiology: photosynthesis · Ocimum basilicum (basil)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 →

other · control · Ocimum basilicum (basil)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 →

drought · physiology: oxidative · Ocimum basilicum (basil)Results agree

5 papers: 0 positive, 5 negative or no effect

4 papers in the same direction as this paper

  1. negative
  2. negative
  3. negative

1 paper more in this group

See all 5 papers →

drought · physiology: photosynthesis · Ocimum basilicum (basil)Results agree

7 papers: 0 positive, 7 negative or no effect

6 papers in the same direction as this paper

  1. negative
  2. negative
  3. negative

3 papers more in this group

See all 7 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.