Paper · 2026 · Plant Physiology and Biochemistry

A novel role of the SQDG synthase gene in Eucommia ulmoides SQD2 in drought tolerance: Lipid remodeling, chloroplast protection, and antioxidant regulation in transgenic tobaccos

Lidan Guo; Ting Yuan; Xian Gong; et al.

18
claims
43
verbatim quotes
0
citations
2
research areas
✓
Open access · cc-by

Abstract

Sulfoquinovosyldiacylglycerol (SQDG) is a key anionic lipid of thylakoid membranes, but its contribution to drought tolerance remains poorly understood. In this study, the Eucommia ulmoides SQDG synthase gene EuSQD2 was functionally characterized and heterologously expressed in tobacco. EuSQD2 showed preferential expression in young leaves and encoded a conserved SQDG synthase. Transgenic tobacco plants overexpressing EuSQD2 exhibited enhanced drought tolerance, as evidenced by reduced wilting, lower reactive oxygen species accumulation, improved photosynthetic performance, higher water use efficiency, and increased relative water content. Lipidomic analyses revealed substantial SQDG accumulation accompanied by coordinated remodeling of membrane lipids, including elevated phosphatidylglycerol and phosphatidylethanolamine levels. Ultrastructural observations demonstrated that EuSQD2 overexpression effectively preserved chloroplast and thylakoid integrity under drought stress. Moreover, transgenic plants displayed optimized stomatal regulation, enhanced proline accumulation, reduced lipid peroxidation, and increased antioxidant enzyme activities. Transcriptomic analyses indicated enrichment of pathways related to photosynthesis, lipid metabolism, antioxidant defense, and stomatal regulation. These results demonstrate that EuSQD2 enhances drought tolerance by promoting SQDG-mediated membrane stabilization and coordinated physiological and molecular adjustments.

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

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Claims 18

Observationobservedoriginal resultmeasured

Claim · model-written summary

EuSQD2 was preferentially expressed in young leaves of Eucommia ulmoides.

Taxon
Eucommia ulmoidesEucommia ulmoidest0
Organ / stage
young leavese0
Gene
EuSQD2e0
Outcome
preferential expressione0

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

Eucommia ulmoides
Eucommia ulmoides · TaxonSten Porse · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“Eucommia ulmoides”

✓ located verbatim in the source · title, characters 42–59 · show in text ↑
e0

“EuSQD2 showed preferential expression in young leaves”

✓ located verbatim in the source · abstract, characters 288–341 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

EuSQD2 encodes a conserved SQDG synthase.

Taxon
Eucommia ulmoidesEucommia ulmoidest0
Gene
SQDG synthasee1
Outcome
conserved SQDG synthasee1

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

Eucommia ulmoides
Eucommia ulmoides · TaxonSten Porse · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“Eucommia ulmoides”

✓ located verbatim in the source · title, characters 42–59 · show in text ↑
e1

“encoded a conserved SQDG synthase”

✓ located verbatim in the source · abstract, characters 346–379 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Transgenic tobacco overexpressing EuSQD2 showed enhanced drought tolerance.

Treatment
overexpressing EuSQD2e2
Gene
EuSQD2e2
Outcome
drought tolerancee2
Direction
enhancede2

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e2

“Transgenic tobacco plants overexpressing EuSQD2 exhibited enhanced drought tolerance”

✓ located verbatim in the source · abstract, characters 381–465 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

EuSQD2 overexpression reduced wilting of tobacco under drought.

Treatment
overexpressing EuSQD2e2
Outcome
wiltinge3
Direction
reducede3

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e2

“Transgenic tobacco plants overexpressing EuSQD2 exhibited enhanced drought tolerance”

✓ located verbatim in the source · abstract, characters 381–465 · show in text ↑
e3

“reduced wilting”

✓ located verbatim in the source · abstract, characters 483–498 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

EuSQD2 overexpression lowered reactive oxygen species accumulation in tobacco under drought.

Treatment
overexpressing EuSQD2e2
Outcome
reactive oxygen species accumulatione4
Direction
lowere4

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e2

“Transgenic tobacco plants overexpressing EuSQD2 exhibited enhanced drought tolerance”

✓ located verbatim in the source · abstract, characters 381–465 · show in text ↑
e4

“lower reactive oxygen species accumulation”

✓ located verbatim in the source · abstract, characters 500–542 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

EuSQD2 overexpression improved photosynthetic performance of tobacco under drought.

Treatment
overexpressing EuSQD2e2
Outcome
photosynthetic performancee5
Direction
improvede5

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e2

“Transgenic tobacco plants overexpressing EuSQD2 exhibited enhanced drought tolerance”

✓ located verbatim in the source · abstract, characters 381–465 · show in text ↑
e5

“improved photosynthetic performance”

✓ located verbatim in the source · abstract, characters 544–579 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

EuSQD2 overexpression gave higher water use efficiency in tobacco under drought.

Treatment
overexpressing EuSQD2e2
Outcome
water use efficiencye6
Direction
highere6

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e2

“Transgenic tobacco plants overexpressing EuSQD2 exhibited enhanced drought tolerance”

✓ located verbatim in the source · abstract, characters 381–465 · show in text ↑
e6

“higher water use efficiency”

✓ located verbatim in the source · abstract, characters 581–608 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

EuSQD2 overexpression increased relative water content of tobacco under drought.

Treatment
overexpressing EuSQD2e2
Outcome
relative water contente7
Direction
increasede7

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e2

“Transgenic tobacco plants overexpressing EuSQD2 exhibited enhanced drought tolerance”

✓ located verbatim in the source · abstract, characters 381–465 · show in text ↑
e7

“increased relative water content”

✓ located verbatim in the source · abstract, characters 614–646 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

EuSQD2 overexpression increased SQDG accumulation in tobacco.

Treatment
overexpressing EuSQD2e1
Outcome
SQDG accumulatione0
Direction
increaseinferrede0

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e0

“Lipidomic analyses revealed substantial SQDG accumulation”

✓ located verbatim in the source · abstract, characters 648–705 · show in text ↑
e1

“Transgenic tobacco plants overexpressing EuSQD2”

✓ located verbatim in the source · abstract, characters 381–428 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

EuSQD2 overexpression elevated phosphatidylglycerol and phosphatidylethanolamine levels in tobacco.

Treatment
overexpressing EuSQD2e1
Outcome
phosphatidylglycerol and phosphatidylethanolamine levelse0
Direction
elevatede0

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e0

“elevated phosphatidylglycerol and phosphatidylethanolamine levels”

✓ located verbatim in the source · abstract, characters 774–839 · show in text ↑
e1

“Transgenic tobacco plants overexpressing EuSQD2”

✓ located verbatim in the source · abstract, characters 381–428 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

EuSQD2 overexpression preserved chloroplast and thylakoid integrity in tobacco under drought stress.

Treatment
EuSQD2 overexpressione0
Gene
EuSQD2e0
Outcome
chloroplast and thylakoid integritye0
Direction
preservede0
Qualifier
under drought stresse0

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e0

“EuSQD2 overexpression effectively preserved chloroplast and thylakoid integrity under drought stress”

✓ located verbatim in the source · abstract, characters 888–988 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Transgenic tobacco showed optimized stomatal regulation.

Treatment
transgenic plantse0
Outcome
stomatal regulatione0
Direction
optimizede0

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e0

“transgenic plants displayed optimized stomatal regulation, enhanced proline accumulation, reduced lipid peroxidation, and increased antioxidant enzyme activities”

✓ located verbatim in the source · abstract, characters 1000–1161 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Transgenic tobacco showed enhanced proline accumulation.

Treatment
transgenic plantse0
Outcome
proline accumulatione0
Direction
enhancede0

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e0

“transgenic plants displayed optimized stomatal regulation, enhanced proline accumulation, reduced lipid peroxidation, and increased antioxidant enzyme activities”

✓ located verbatim in the source · abstract, characters 1000–1161 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Transgenic tobacco showed reduced lipid peroxidation.

Treatment
transgenic plantse0
Outcome
lipid peroxidatione0
Direction
reducede0

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e0

“transgenic plants displayed optimized stomatal regulation, enhanced proline accumulation, reduced lipid peroxidation, and increased antioxidant enzyme activities”

✓ located verbatim in the source · abstract, characters 1000–1161 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Transgenic tobacco showed increased antioxidant enzyme activities.

Treatment
transgenic plantse0
Outcome
antioxidant enzyme activitiese0
Direction
increasede0

Not reportedComparator

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e0

“transgenic plants displayed optimized stomatal regulation, enhanced proline accumulation, reduced lipid peroxidation, and increased antioxidant enzyme activities”

✓ located verbatim in the source · abstract, characters 1000–1161 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Transcriptomic analyses showed enrichment of pathways related to photosynthesis, lipid metabolism and antioxidant defense in tobacco.

Outcome
enrichment of pathways related to photosynthesis, lipid metabolism, antioxidant defensee0

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e0

“Transcriptomic analyses indicated enrichment of pathways related to photosynthesis, lipid metabolism, antioxidant defense, and stomatal regulation”

✓ located verbatim in the source · abstract, characters 1163–1309 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Transcriptomic analyses showed enrichment of pathways related to stomatal regulation in tobacco.

Outcome
stomatal regulatione0
Qualifier
Transcriptomic analyses indicated enrichment of pathways related to photosynthesis, lipid metabolism, antioxidant defense, and stomatal regulatione0

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e0

“Transcriptomic analyses indicated enrichment of pathways related to photosynthesis, lipid metabolism, antioxidant defense, and stomatal regulation”

✓ located verbatim in the source · abstract, characters 1163–1309 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivebasis n/a

Claim · model-written summary

EuSQD2 enhances drought tolerance in tobacco by promoting SQDG-mediated membrane stabilization.

Gene
EuSQD2e0
Compound
SQDGe0
Outcome
drought tolerancee0
Qualifier
SQDG-mediated membrane stabilizatione0

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

Nicotiana tabacum
Nicotiana tabacum · TaxonJom / Joachim Müllerchen · CC BY 2.5 · Wikimedia Commons ↗

Verbatim

t0

“transgenic tobaccos”

✓ located verbatim in the source · title, characters 159–178 · show in text ↑
e0

“These results demonstrate that EuSQD2 enhances drought tolerance by promoting SQDG-mediated membrane stabilization”

✓ located verbatim in the source · abstract, characters 1311–1425 · show in text ↑