Abstract
TCH4 is a xyloglucan endotransglucosylase/hydrolase (XTH) family member. Extensive studies have shown that XTHs are very important in cell wall homeostasis for plant growth and development. Boron (B), as an essential micronutrient for plants, plays an essential role in the cross-linking of cell wall pectin. However, the effect of B on cell wall organization is unclear. This study aimed to explore the mechanism of plant adaption to B stress by investigating the role of TCH4 in cell wall homeostasis. We conducted both plate and hydroponic cultures of wild-type Col-0 and overexpression and gene knockout lines of XTH22/TCH4 to analyze the phenotype, components, and characteristics of the cell wall using immunofluorescence, atomic force microscopy (AFM), and transmission electron microscopy (TEM). B deficiency induces the expression of TCH4. The overexpression lines of TCH4 presented more sensitivity to B deficiency than the wild-type Col-0, while the knockout lines of TCH4 were more resistant to low B stress. Up-regulation of TCH4 influenced the ratio of chelator-soluble pectin to alkali-soluble pectin and decreased the degree of methylesterification of pectin under B-deficient conditions. Moreover, we found that B deficiency disturbed the arrangement of cellulose, enlarged the gap between cellulose microfibrils, and decreased the mechanical strength of the cell wall, leading to the formation of a thickened and deformed triangular region of the cell wall. These symptoms were more profound in the TCH4 overexpression lines. Consistently, compared with Col-0, the O2− and MDA contents in the TCH4 overexpression lines increased under B-deficient conditions. This study identified the B-deficiency-induced TCH4 gene, which regulates cell wall homeostasis to influence plant growth under B-deficient conditions.
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Claims 13
Claim · model-written summary
Boron deficiency induced TCH4 expression.
- Taxon
- Arabidopsis
Arabidopsist0 - Treatment
- B deficiencye0
- Gene
- TCH4e0
- Outcome
- expression of TCH4e0
- Direction
- inducese0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“B deficiency induces the expression of TCH4”
t0“Arabidopsis”
Claim · model-written summary
TCH4 overexpression lines were more sensitive to boron deficiency than wild-type Col-0.
- Taxon
- Arabidopsis
Arabidopsist0 - Material
- overexpression lines of TCH4e0
- Treatment
- B deficiencye0
- Gene
- TCH4e0
- Outcome
- sensitivity to B deficiencye0
- Direction
- moree0
- Comparator
- wild-type Col-0e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The overexpression lines of TCH4 presented more sensitivity to B deficiency than the wild-type Col-0”
t0“Arabidopsis”
Claim · model-written summary
TCH4 knockout lines were more resistant to low boron stress than wild-type Col-0.
- Taxon
- Arabidopsis
Arabidopsist0 - Material
- knockout lines of TCH4e0
- Treatment
- low B stresse0
- Gene
- TCH4e0
- Outcome
- resistant to low B stresse0
- Direction
- moree0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the knockout lines of TCH4 were more resistant to low B stress”
t0“Arabidopsis”
Claim · model-written summary
TCH4 upregulation altered the ratio of chelator-soluble to alkali-soluble pectin under boron deficiency.
- Taxon
- Arabidopsis
Arabidopsist0 - Gene
- TCH4e0
- Outcome
- ratio of chelator-soluble pectin to alkali-soluble pectine0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Up-regulation of TCH4 influenced the ratio of chelator-soluble pectin to alkali-soluble pectin”
t0“Arabidopsis”
Claim · model-written summary
TCH4 upregulation decreased pectin methylesterification under boron deficiency.
- Taxon
- Arabidopsis
Arabidopsist0 - Treatment
- B-deficient conditionse0
- Gene
- TCH4e0
- Outcome
- degree of methylesterification of pectine0
- Direction
- decreasede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Up-regulation of TCH4 influenced the ratio of chelator-soluble pectin to alkali-soluble pectin and decreased the degree of methylesterification of pectin under B-deficient conditions”
t0“Arabidopsis”
Claim · model-written summary
Boron deficiency disturbed cellulose arrangement.
- Taxon
- Arabidopsis
Arabidopsist0 - Treatment
- B deficiencye0
- Outcome
- arrangement of cellulosee0
- Direction
- disturbede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“B deficiency disturbed the arrangement of cellulose”
t0“Arabidopsis”
Claim · model-written summary
Boron deficiency enlarged the gap between cellulose microfibrils.
- Taxon
- Arabidopsis
Arabidopsist0 - Treatment
- B deficiencye0
- Outcome
- gap between cellulose microfibrilse0
- Direction
- enlargede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“B deficiency disturbed the arrangement of cellulose, enlarged the gap between cellulose microfibrils”
t0“Arabidopsis”
Claim · model-written summary
Boron deficiency decreased cell wall mechanical strength.
- Taxon
- Arabidopsis
Arabidopsist0 - Treatment
- B deficiencye0
- Outcome
- mechanical strength of the cell walle0
- Direction
- decreasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“B deficiency disturbed the arrangement of cellulose, enlarged the gap between cellulose microfibrils, and decreased the mechanical strength of the cell wall”
t0“Arabidopsis”
Claim · model-written summary
Boron deficiency produced a thickened, deformed triangular cell wall region.
- Taxon
- Arabidopsis
Arabidopsist0 - Outcome
- thickened and deformed triangular region of the cell walle0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“leading to the formation of a thickened and deformed triangular region of the cell wall”
t0“Arabidopsis”
Claim · model-written summary
Cell wall symptoms under boron deficiency were more profound in TCH4 overexpression lines.
- Taxon
- Arabidopsis
Arabidopsist0 - Material
- TCH4 overexpression linese0
- Outcome
- These symptomse0
- Direction
- more profounde0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“These symptoms were more profound in the TCH4 overexpression lines”
t0“Arabidopsis”
Claim · model-written summary
Under boron deficiency, TCH4 overexpression lines had increased O2− content compared with Col-0.
- Taxon
- Arabidopsis
Arabidopsist0 - Material
- TCH4 overexpression linese0
- Treatment
- B-deficient conditionse0
- Outcome
- O2−e0
- Direction
- increasede0
- Comparator
- Col-0e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“compared with Col-0, the O2− and MDA contents in the TCH4 overexpression lines increased under B-deficient conditions”
t0“Arabidopsis”
Claim · model-written summary
Under boron deficiency, TCH4 overexpression lines had increased MDA content compared with Col-0.
- Taxon
- Arabidopsis
Arabidopsist0 - Material
- TCH4 overexpression linese0
- Treatment
- B-deficient conditionse0
- Outcome
- MDA contentse0
- Direction
- increasede0
- Comparator
- Col-0e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“compared with Col-0, the O2− and MDA contents in the TCH4 overexpression lines increased under B-deficient conditions”
t0“Arabidopsis”
Claim · model-written summary
The authors identify boron deficiency induced TCH4 as a regulator of cell wall homeostasis and plant growth.
- Taxon
- Arabidopsis
Arabidopsist0 - Gene
- TCH4e0
- Outcome
- plant growthe0
- Qualifier
- cell wall homeostasise0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“B-deficiency-induced TCH4 gene, which regulates cell wall homeostasis to influence plant growth under B-deficient conditions”
t0“Arabidopsis”