Abstract
Heat stress (HS) severely affects plant growth and productivity worldwide. The application of osmoprotectant compounds, such as proline, has been shown to mitigate these effects; however, excessive or uncontrolled proline uptake can lead to pleiotropic effects that compromise plant development. Encapsulation technology offers a promising alternative to achieve a gradual and controlled release of proline, improving its protective efficiency. In this work, proline (Pro) was encapsulated using silica (Si:Pro) and chitosan (Ch:Pro) matrices at different ratios by a spray-drying process, an industrially scalable technique with potential relevance for agricultural applications. The proper homogenization of the Si or Ch matrices with Pro enabled the formulation of suspensions with suitable rheological behavior for efficient atomization. This subsequently allowed effective drying and the obtention of free-flowing particles with high thermal stability compared with non-encapsulated proline, and satisfactory encapsulation efficiencies. These properties allowed the use of Si:Pro and Ch:Pro samples at the three formulated ratios for subsequent biological experiments. In Arabidopsis thaliana , free Pro treatment negatively affected root and rosette growth under non-stress conditions, whereas encapsulated Pro, particularly Si:Pro, mitigated these effects through controlled release. Under HS conditions, plants treated with Si:Pro showed markedly enhanced growth, with root length increasing by ~80%, rosette area by ~100%, and fresh weight by ~20% compared with HS-treated controls. Furthermore, Si:Pro treatment promoted a more balanced hormonal profile, characterized by a roughly twofold increase in salicylic acid (SA), a threefold increase in abscisic acid (ABA), and an approximate 50% reduction in indole-3-acetic acid (IAA) relative to HS-treated plants. Moreover, plants treated with Si:Pro + HS exhibited moderate expression of proline biosynthetic genes, increased ascorbate peroxidase (APX) and glutathione reductase (GR) activities, and decreased lipid peroxidation, collectively supporting an enhanced redox homeostasis. Therefore, silica-based encapsulation of proline provides a controlled delivery system that reduces toxicity, enhances thermotolerance, and maintains physiological and hormonal balance in Arabidopsis thaliana under HS. These findings support its potential as a sustainable and scalable strategy to improve plant resilience, while highlighting the need for further optimization and validation across crop systems and environmental conditions.
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Claims 11
Claim · model-written summary
Free proline reduced root and rosette growth under non-stress conditions.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae0 - Treatment
- free Pro treatmente0
- Outcome
- root and rosette growthe0
- Direction
- negatively affectede0
- Qualifier
- non-stress conditionse0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“In Arabidopsis thaliana , free Pro treatment negatively affected root and rosette growth under non-stress conditions”
Claim · model-written summary
Encapsulated proline, particularly Si:Pro, mitigated the growth harm under non-stress conditions.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae1 - Treatment
- encapsulated Pro, particularly Si:Proe0
- Outcome
- root and rosette growthe0
- Direction
- mitigatede0
- Comparator
- free Pro treatmente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“free Pro treatment negatively affected root and rosette growth under non-stress conditions, whereas encapsulated Pro, particularly Si:Pro, mitigated these effects through controlled release”
e1“In Arabidopsis thaliana”
Claim · model-written summary
Si:Pro increased root length under heat stress by about 80% versus heat-stressed controls.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae1 - Treatment
- Si:Proe0
- Outcome
- root lengthe0
- Direction
- increasinge0
- Value
- ~80%e0
- Comparator
- HS-treated controlse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Under HS conditions, plants treated with Si:Pro showed markedly enhanced growth, with root length increasing by ~80%, rosette area by ~100%, and fresh weight by ~20% compared with HS-treated controls”
e1“In Arabidopsis thaliana”
Claim · model-written summary
Si:Pro increased rosette area under heat stress by about 100% versus heat-stressed controls.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae1 - Treatment
- Si:Proe0
- Outcome
- rosette areae0
- Direction
- increasinge0
- Value
- ~100%e0
- Comparator
- HS-treated controlse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Under HS conditions, plants treated with Si:Pro showed markedly enhanced growth, with root length increasing by ~80%, rosette area by ~100%, and fresh weight by ~20% compared with HS-treated controls”
e1“In Arabidopsis thaliana”
Claim · model-written summary
Si:Pro increased fresh weight under heat stress by about 20% versus heat-stressed controls.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae1 - Treatment
- Si:Proe0
- Outcome
- fresh weighte0
- Direction
- increasinge0
- Value
- ~20%e0
- Comparator
- HS-treated controlse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Under HS conditions, plants treated with Si:Pro showed markedly enhanced growth, with root length increasing by ~80%, rosette area by ~100%, and fresh weight by ~20% compared with HS-treated controls”
e1“In Arabidopsis thaliana”
Claim · model-written summary
Si:Pro approximately doubled salicylic acid under heat stress.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae1 - Treatment
- Si:Pro treatmente0
- Outcome
- salicylic acid (SA)e0
- Direction
- increasee0
- Value
- roughly twofolde0
- Comparator
- HS-treated plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Si:Pro treatment promoted a more balanced hormonal profile, characterized by a roughly twofold increase in salicylic acid (SA), a threefold increase in abscisic acid (ABA), and an approximate 50% reduction in indole-3-acetic acid (IAA) relative to HS-treated plants”
e1“In Arabidopsis thaliana”
Claim · model-written summary
Si:Pro tripled abscisic acid under heat stress.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae1 - Treatment
- Si:Pro treatmente0
- Outcome
- abscisic acid (ABA)e0
- Direction
- increasee0
- Value
- threefolde0
- Comparator
- HS-treated plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Si:Pro treatment promoted a more balanced hormonal profile, characterized by a roughly twofold increase in salicylic acid (SA), a threefold increase in abscisic acid (ABA), and an approximate 50% reduction in indole-3-acetic acid (IAA) relative to HS-treated plants”
e1“In Arabidopsis thaliana”
Claim · model-written summary
Si:Pro reduced indole-3-acetic acid by about 50% under heat stress.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae1 - Treatment
- Si:Pro treatmente0
- Outcome
- indole-3-acetic acid (IAA)e0
- Direction
- reductione0
- Value
- approximate 50%e0
- Comparator
- HS-treated plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Si:Pro treatment promoted a more balanced hormonal profile, characterized by a roughly twofold increase in salicylic acid (SA), a threefold increase in abscisic acid (ABA), and an approximate 50% reduction in indole-3-acetic acid (IAA) relative to HS-treated plants”
e1“In Arabidopsis thaliana”
Claim · model-written summary
Si:Pro plus heat stress yielded moderate expression of proline biosynthetic genes.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae1 - Treatment
- Si:Pro + HSe0
- Gene
- proline biosynthetic genese0
- Outcome
- moderate expression of proline biosynthetic genese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plants treated with Si:Pro + HS exhibited moderate expression of proline biosynthetic genes”
e1“In Arabidopsis thaliana”
Claim · model-written summary
Si:Pro plus heat stress increased ascorbate peroxidase and glutathione reductase activities.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae1 - Treatment
- Si:Pro + HSe0
- Outcome
- ascorbate peroxidase (APX) and glutathione reductase (GR) activitiese0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plants treated with Si:Pro + HS exhibited moderate expression of proline biosynthetic genes, increased ascorbate peroxidase (APX) and glutathione reductase (GR) activities, and decreased lipid peroxidation”
e1“In Arabidopsis thaliana”
Claim · model-written summary
Si:Pro plus heat stress decreased lipid peroxidation.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae1 - Treatment
- Si:Pro + HSe0
- Outcome
- lipid peroxidatione0
- Direction
- decreasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plants treated with Si:Pro + HS exhibited moderate expression of proline biosynthetic genes, increased ascorbate peroxidase (APX) and glutathione reductase (GR) activities, and decreased lipid peroxidation”
e1“In Arabidopsis thaliana”
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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.