Abstract
• Drought and alkaline salt stress disrupted potato growth and leaf microstructure, with the combined stress changes being the most significant. • Drought and alkaline salt stress increased the accumulation of ROS (H 2 O 2 and O 2 - ), osmoregulatory substances (MDA, Pro, SS, AA, GSH, and AsA), and enhanced antioxidant enzyme (SOD, POD, CAT, TPX, APX, GR, GPX, and DHAR) activities. • Drought and alkaline salt stress negatively affected photosynthesis parameters. P n , G s , and T r were significantly reduced, and C i significantly increased after the stress. • Drought and alkaline salt stress increased the contents of ABA, SA, and BRs and decreased the contents of GA 3 and IAA, inducing changes in the expression patterns of hormone-related genes and activating their signaling. • Drought and alkaline salt stress reduced potato tuber yield and quality. The combined stress reduced yield by 54.13 % (2022) and 60.14 % (2023). In the arid and semi-arid zones of Northwest China, soil drought and alkaline salt stress often occur simultaneously and affect plant growth at multiple levels. Potato ( Solanum tuberosum L.) is a food crop sensitive to drought and alkaline salt stresses and is susceptible to yield loss due to environmental impacts. In recent years, most of the research on abiotic stress response in potato has focused on drought and saline single stresses, and the mechanism of potato response to combined drought-alkaline salt stress and its interactions are still unclear. Therefore, a pot experiment was designed in this study and the potato variety ' Atlantic ' was selected as the test material. The effects of drought (25 % PEG-6000), alkaline salt (200 mmol·L -1 NaHCO 3 ) and combined drought-alkaline salt (25 % PEG-6000 + 200 mmol·L -1 NaHCO 3 ) stresses on growth traits, micro- and ultrastructure, reactive oxygen species, osmoregulatory substances, and antioxidant defenses of potato were investigated using no stress (CK) as a control, leaf photosynthesis and endogenous plant hormones, and also analyzed the changes in the expression patterns of genes related to plant hormone signal transduction under different stresses. The results showed that drought, alkaline salt, and combined stress affected growth, leaf anatomy, and photosynthesis, and increased the accumulation of osmoregulatory substances in potato. The scavenging activities of antioxidant compounds and antioxidant enzymes were enhanced in potato, and combined stress treatments significantly damaged potato more than single stresses. In 2022, combined stress caused a marked increase in H 2 O 2 (208.7 %) and O 2 - (455.6 %) content, while in 2023, they increased by 87.5 % and 215.7 %, respectively. SOD, POD, CAT, TPX, APX, GR, GPX and DHAR enzyme activities were increased by 209.13 %, 55.19 %, 152.59 %, 47.13 %, 104.02 %, 347.37 %, 68.45 % and 130.69 % in 2022 compared to CK in the combined stress treatment. In 2023, they increased by 229.81 %, 49.95 %, 160.62 %, 102.16 %, 94.06 %, 505.15 %, 47.00 %, and 121.19 %, respectively. After the stress treatments, the contents of gibberellic acid (GA 3 ) and auxins (IAA) were significantly lower than those in CK, whereas the contents of abscisic acid (ABA), salicylic acid (SA), and brassinosteroids (BRs) increased. Expression of IAA-related genes (AUX1, Aux/IAA, GH3, and SAUR) was up-regulated after stress. ABA-related genes (PYR/PYL, SnRK2, and ABF) were up-regulated after stress, whereas protein phosphatase 2C (PP2C) genes were down-regulated in expression after stress. The GA 3 receptor GID1 and the F-box protein GID2 were up-regulated after stress. Xyloglucosyl transferase TCH4 gene was up-regulated by stress and positively correlated with changes in BRs content. The TGA transcription factor, PR-1 gene, was induced to up-regulate its expression by stress and positively correlated with changes in SA content. Drought, alkaline salt, and combined stress reduced potato tuber yield and quality, which were 54.13 % and 60.14 % lower than CK in combined stress treatments in 2022 and 2023, respectively, which were significantly correlated with changes in physiological and biochemical characteristics and hormone contents of potato plants.
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Claims 20
Claim · model-written summary
Stresses disrupted potato growth and leaf microstructure, most severely when combined.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- Drought and alkaline salt stresse1
- Outcome
- potato growth and leaf microstructuree1
- Direction
- disruptede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Drought and alkaline salt stress disrupted potato growth and leaf microstructure, with the combined stress changes being the most significant”
Claim · model-written summary
Stress increased H 2 O 2 and O 2 - accumulation.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- Drought and alkaline salt stresse1
- Outcome
- accumulation of ROS (H 2 O 2 and O 2 - )e1
- Direction
- increasede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Drought and alkaline salt stress increased the accumulation of ROS (H 2 O 2 and O 2 - )”
Claim · model-written summary
Stress increased osmoregulatory substance accumulation.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- Drought and alkaline salt stresse1
- Outcome
- osmoregulatory substances (MDA, Pro, SS, AA, GSH, and AsA)e1
- Direction
- increasede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Drought and alkaline salt stress increased the accumulation of ROS (H 2 O 2 and O 2 - ), osmoregulatory substances (MDA, Pro, SS, AA, GSH, and AsA)”
Claim · model-written summary
Stress enhanced antioxidant enzyme activities.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- Drought and alkaline salt stresse1
- Outcome
- antioxidant enzyme (SOD, POD, CAT, TPX, APX, GR, GPX, and DHAR) activitiese1
- Direction
- enhancede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Drought and alkaline salt stress increased the accumulation of ROS (H 2 O 2 and O 2 - ), osmoregulatory substances (MDA, Pro, SS, AA, GSH, and AsA), and enhanced antioxidant enzyme (SOD, POD, CAT, TPX, APX, GR, GPX, and DHAR) activities”
Claim · model-written summary
Stress significantly reduced P n, G s and T r.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- Drought and alkaline salt stresse1
- Outcome
- P n , G s , and T re1
- Direction
- reducede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Drought and alkaline salt stress negatively affected photosynthesis parameters. P n , G s , and T r were significantly reduced”
Claim · model-written summary
Stress significantly increased C i.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- the stresse1
- Outcome
- C ie1
- Direction
- increasede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“C i significantly increased after the stress”
Claim · model-written summary
Stress increased ABA, SA and BRs contents.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- Drought and alkaline salt stresse1
- Outcome
- contents of ABA, SA, and BRse1
- Direction
- increasede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Drought and alkaline salt stress increased the contents of ABA, SA, and BRs”
Claim · model-written summary
Stress significantly decreased GA 3 and IAA contents.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- stress treatmentse1
- Outcome
- contents of gibberellic acid (GA 3 ) and auxins (IAA)e1
- Direction
- lowere1
- Comparator
- CKe1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“After the stress treatments, the contents of gibberellic acid (GA 3 ) and auxins (IAA) were significantly lower than those in CK”
Claim · model-written summary
IAA-related genes were up-regulated after stress.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- stresse1
- Gene
- IAA-related genes (AUX1, Aux/IAA, GH3, and SAUR)e1
- Outcome
- Expressione1
- Direction
- up-regulatede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Expression of IAA-related genes (AUX1, Aux/IAA, GH3, and SAUR) was up-regulated after stress”
Claim · model-written summary
PYR/PYL, SnRK2 and ABF genes were up-regulated after stress.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- stresse1
- Gene
- ABA-related genes (PYR/PYL, SnRK2, and ABF)e1
- Outcome
- ABA-related genese1
- Direction
- up-regulatede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“ABA-related genes (PYR/PYL, SnRK2, and ABF) were up-regulated after stress”
Claim · model-written summary
PP2C genes were down-regulated after stress.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- stresse1
- Gene
- protein phosphatase 2C (PP2C) genese1
- Outcome
- expressione1
- Direction
- down-regulatede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“protein phosphatase 2C (PP2C) genes were down-regulated in expression after stress”
Claim · model-written summary
GID1 and GID2 were up-regulated after stress.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- stresse1
- Gene
- GID1 and the F-box protein GID2e1
- Outcome
- GID1 and the F-box protein GID2e1
- Direction
- up-regulatede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“The GA 3 receptor GID1 and the F-box protein GID2 were up-regulated after stress”
Claim · model-written summary
TCH4 was up-regulated by stress.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- stresse1
- Gene
- TCH4 genee1
- Outcome
- TCH4 genee1
- Direction
- up-regulatede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Xyloglucosyl transferase TCH4 gene was up-regulated by stress”
Claim · model-written summary
TCH4 changes were positively correlated with BRs content.
- Taxon
- potato
Solanum tuberosumt0 - Gene
- TCH4 genee1
- Outcome
- changes in BRs contente1
- Direction
- positively correlatede1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Xyloglucosyl transferase TCH4 gene was up-regulated by stress and positively correlated with changes in BRs content”
Claim · model-written summary
TGA and PR-1 expression was up-regulated by stress.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- stresse1
- Gene
- TGA transcription factor, PR-1 genee1
- Outcome
- expressione1
- Direction
- up-regulatee1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“The TGA transcription factor, PR-1 gene, was induced to up-regulate its expression by stress”
Claim · model-written summary
TGA and PR-1 expression correlated positively with SA content changes.
- Taxon
- potato
Solanum tuberosumt0 - Gene
- TGA transcription factor, PR-1 genee1
- Outcome
- changes in SA contente1
- Direction
- positively correlatede1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“The TGA transcription factor, PR-1 gene, was induced to up-regulate its expression by stress and positively correlated with changes in SA content”
Claim · model-written summary
Combined stress raised H 2 O 2 content by 208.7 % in 2022 and 87.5 % in 2023.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- combined stresse1
- Outcome
- H 2 O 2e1
- Direction
- increasee1
- Value
- 208.7 %e1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“In 2022, combined stress caused a marked increase in H 2 O 2 (208.7 %) and O 2 - (455.6 %) content, while in 2023, they increased by 87.5 % and 215.7 %, respectively”
Claim · model-written summary
Combined stress raised O 2 - content by 455.6 % in 2022 and 215.7 % in 2023.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- combined stresse1
- Outcome
- O 2 -e1
- Direction
- increasee1
- Value
- 455.6 %e1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“In 2022, combined stress caused a marked increase in H 2 O 2 (208.7 %) and O 2 - (455.6 %) content, while in 2023, they increased by 87.5 % and 215.7 %, respectively”
Claim · model-written summary
Combined stress reduced tuber yield by 54.13 % in 2022 and 60.14 % in 2023.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- combined stresse1
- Outcome
- potato tuber yielde1
- Direction
- lowere1
- Value
- 54.13 %e1
- Comparator
- CKe1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Drought, alkaline salt, and combined stress reduced potato tuber yield and quality, which were 54.13 % and 60.14 % lower than CK in combined stress treatments in 2022 and 2023, respectively”
Claim · model-written summary
Drought, alkaline salt and combined stress reduced potato tuber quality.
- Taxon
- potato
Solanum tuberosumt0 - Treatment
- Drought, alkaline salt, and combined stresse1
- Outcome
- potato tuber yield and qualitye1
- Direction
- reducede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“potato (Solanum tuberosum L.)”
e1“Drought, alkaline salt, and combined stress reduced potato tuber yield and quality”
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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.