Abstract
Nitrogen (N) and phosphorus (P) are essential phytomacronutrients, and deficiencies in these two elements limit growth and yield in apple ( Malus domestica Borkh.). The rootstock plays a key role in the nutrient uptake and environmental adaptation of apple. The objective of this study was to investigate the effects of N and/or P deficiency on hydroponically-grown dwarfing rootstock ‘M9-T337’ seedlings, particularly the roots, by performing an integrated physiological, transcriptomics-, and metabolomics-based analyses. Compared to N and P sufficiency, N and/or P deficiency inhibited aboveground growth, increased the partitioning of total N and total P in roots, enhanced the total number of tips, length, volume, and surface area of roots, and improved the root-to-shoot ratio. P and/or N deficiency inhibited NO3 − influx into roots, and H + pumps played a important role in the response to P and/or N deficiency. Conjoint analysis of differentially expressed genes and differentially accumulated metabolites in roots revealed that N and/or P deficiency altered the biosynthesis of cell wall components such as cellulose, hemicellulose, lignin, and pectin. The expression of MdEXPA4 and MdEXLB1 , two cell wall expansin genes, were shown to be induced by N and/or P deficiency. Overexpression of MdEXPA4 enhanced root development and improved tolerance to N and/or P deficiency in transgenic Arabidopsis thaliana plants. In addition, overexpression of MdEXLB1 in transgenic Solanum lycopersicum seedlings increased the root surface area and promoted acquisition of N and P, thereby facilitating plant growth and adaptation to N and/or P deficiency. Collectively, these results provided a reference for improving root architecture in dwarfing rootstock and furthering our understanding of integration between N and P signaling pathways.
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Claims 15
Claim · model-written summary
N and/or P deficiency inhibited aboveground growth.
- Taxon
- apple
Malus domesticat0 - Treatment
- N and/or P deficiencye0
- Outcome
- aboveground growthe0
- Direction
- inhibitede0
- Comparator
- N and P sufficiencye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Compared to N and P sufficiency, N and/or P deficiency inhibited aboveground growth”
t0“apple dwarfing rootstock”
Claim · model-written summary
Deficiency increased root partitioning of total N and P.
- Taxon
- apple
Malus domesticat0 - Treatment
- N and/or P deficiencye0
- Outcome
- partitioning of total N and total P in rootse0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“N and/or P deficiency inhibited aboveground growth, increased the partitioning of total N and total P in roots”
t0“apple”
Claim · model-written summary
Deficiency enhanced root tip number, length and volume.
- Taxon
- apple
Malus domesticat0 - Treatment
- N and/or P deficiencye0
- Outcome
- total number of tips, length, volumee0
- Direction
- enhancede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“N and/or P deficiency inhibited aboveground growth, increased the partitioning of total N and total P in roots, enhanced the total number of tips, length, volume, and surface area of roots”
t0“apple dwarfing rootstock”
Claim · model-written summary
Deficiency enhanced root surface area.
- Taxon
- apple
Malus domesticat0 - Treatment
- N and/or P deficiencye0
- Outcome
- surface area of rootse0
- Direction
- enhancede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“N and/or P deficiency inhibited aboveground growth, increased the partitioning of total N and total P in roots, enhanced the total number of tips, length, volume, and surface area of roots”
t0“apple dwarfing rootstock”
Claim · model-written summary
Deficiency improved root-to-shoot ratio.
- Taxon
- apple
Malus domesticat0 - Treatment
- N and/or P deficiencye0
- Outcome
- root-to-shoot ratioe0
- Direction
- improvede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“N and/or P deficiency inhibited aboveground growth, increased the partitioning of total N and total P in roots, enhanced the total number of tips, length, volume, and surface area of roots, and improved the root-to-shoot ratio”
t0“apple dwarfing rootstock”
Claim · model-written summary
P and/or N deficiency inhibited nitrate influx into roots.
- Taxon
- apple
Malus domesticat0 - Treatment
- P and/or N deficiencye0
- Outcome
- NO3 − influx into rootse0
- Direction
- inhibitede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“P and/or N deficiency inhibited NO3 − influx into roots”
t0“apple”
Claim · model-written summary
H+ pumps were implicated in the response to deficiency.
- Taxon
- apple
Malus domesticat0 - Gene
- H + pumpse0
- Outcome
- response to P and/or N deficiencye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“H + pumps played a important role in the response to P and/or N deficiency”
t0“apple”
Claim · model-written summary
N and/or P deficiency altered cell wall component biosynthesis.
- Taxon
- apple
Malus domesticat0 - Treatment
- N and/or P deficiencye0
- Outcome
- biosynthesis of cell wall components such as cellulose, hemicellulose, lignin, and pectine0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“N and/or P deficiency altered the biosynthesis of cell wall components such as cellulose, hemicellulose, lignin, and pectin”
t0“apple”
Claim · model-written summary
Deficiency induced MdEXPA4 expression.
- Taxon
- apple
Malus domesticat0 - Treatment
- N and/or P deficiencye0
- Gene
- MdEXPA4e0
- Outcome
- expression of MdEXPA4e0
- Direction
- inducede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The expression of MdEXPA4 and MdEXLB1 , two cell wall expansin genes, were shown to be induced by N and/or P deficiency”
t0“apple”
Claim · model-written summary
Deficiency induced MdEXLB1 expression.
- Taxon
- apple
Malus domesticat0 - Treatment
- N and/or P deficiencye0
- Gene
- MdEXLB1e0
- Outcome
- MdEXLB1e0
- Direction
- inducede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The expression of MdEXPA4 and MdEXLB1 , two cell wall expansin genes, were shown to be induced by N and/or P deficiency”
t0“apple”
Claim · model-written summary
MdEXPA4 overexpression enhanced Arabidopsis root development.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae0 - Treatment
- Overexpression of MdEXPA4e0
- Gene
- MdEXPA4e0
- Outcome
- root developmente0
- Direction
- enhancede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Overexpression of MdEXPA4 enhanced root development and improved tolerance to N and/or P deficiency in transgenic Arabidopsis thaliana plants”
Claim · model-written summary
MdEXPA4 overexpression improved Arabidopsis deficiency tolerance.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianae0 - Treatment
- Overexpression of MdEXPA4e0
- Gene
- MdEXPA4e0
- Outcome
- tolerance to N and/or P deficiencye0
- Direction
- improvede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Overexpression of MdEXPA4 enhanced root development and improved tolerance to N and/or P deficiency in transgenic Arabidopsis thaliana plants”
Claim · model-written summary
MdEXLB1 overexpression increased tomato root surface area.
- Taxon
- Solanum lycopersicum
Solanum lycopersicume0 - Treatment
- overexpression of MdEXLB1e0
- Gene
- MdEXLB1e0
- Outcome
- root surface areae0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“overexpression of MdEXLB1 in transgenic Solanum lycopersicum seedlings increased the root surface area”
Claim · model-written summary
MdEXLB1 overexpression promoted N and P acquisition in tomato.
- Taxon
- Solanum lycopersicum
Solanum lycopersicume0 - Treatment
- overexpression of MdEXLB1e0
- Gene
- MdEXLB1e0
- Outcome
- acquisition of N and Pe0
- Direction
- promotede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“overexpression of MdEXLB1 in transgenic Solanum lycopersicum seedlings increased the root surface area and promoted acquisition of N and P”
Claim · model-written summary
MdEXLB1 overexpression facilitated tomato growth and adaptation to deficiency.
- Taxon
- Solanum lycopersicum
Solanum lycopersicume0 - Treatment
- overexpression of MdEXLB1e0
- Gene
- MdEXLB1e0
- Outcome
- plant growth and adaptation to N and/or P deficiencye0
- Direction
- facilitatinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“overexpression of MdEXLB1 in transgenic Solanum lycopersicum seedlings increased the root surface area and promoted acquisition of N and P, thereby facilitating plant growth and adaptation to N and/or P deficiency”
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