Abstract
Phosphorus (P) is an important nutrient affecting nodulation and nitrogen fixation in soybeans. To further investigate the relationship of phosphorus with soybean nodulation and nitrogen fixation, the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment. From the unfolded cotyledon stage to the initial flowering stage, one side of each dual-root soybean system was irrigated with nutrient solution containing 1 mg/L, 31 mg/L, or 61 mg/L of phosphorus (phosphorus-application side), and the other side was irrigated with a phosphorus-free nutrient solution (phosphorus-free side), to study the effect of local phosphorus supply on nodulation and nitrogen fixation in soybean. The results are described as follows: (1) Increasing the phosphorus supply increased the nodules weight, nitrogenase activity, ureide content, number of bacteroids, number of infected cells, and relative expression levels of nodule nitrogen fixation key genes (GmEXPB2, GmSPX5, nifH, nifD, nifK, GmALN1, GmACP1, GmUR5, GmPUR5, and GmHIUH5) in root nodules on the phosphorus-application side. Although the phosphorus-application and phosphorus-free sides demonstrated similar changing trends, the phosphorus-induced increases were more prominent on the phosphorus-application side, which indicated that phosphorus supply systematically regulates nodulation and nitrogen fixation in soybean. (2) When the level of phosphorus supply was increased from 1 mg/L to 31 mg/L, the increase on the P– side root was significant, and nodule phosphorus content increased by 57.14–85.71% and 68.75–75.00%, respectively; ARA and SNA were 218.64–383.33% and 11.41–16.11%, respectively, while ureide content was 118.18–156.44%. When the level of phosphorus supply was increased from 31mg/L to 61mg/L, the increase in the regulation ability of root and nodule phosphorus content, ARA, SNA, and ureide content were low for roots, and the value for nodules was lower than when the phosphorus level increased from 1 mg/L to 31 mg/L. (3) A high-concentration phosphorus supply on one side of a dual-root soybean plant significantly increased the phosphorus content in the aboveground tissues, as well as the roots and nodules on both sides. In the roots on the phosphorus-free side, the nodules were prioritized for receiving the phosphorus transported from the aboveground tissues to maintain their phosphorus content and functionality.
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Claims 13
Claim · model-written summary
Increasing phosphorus supply increased nodule weight, nitrogenase activity and ureide content in dual-root soybean on the phosphorus-application side.
- Taxon
- soybean
Glycine maxe5 - Organ / stage
- nodulese1
- Treatment
- Increasing the phosphorus supplye1
- Outcome
- nodules weight, nitrogenase activity, ureide contente1
- Direction
- increasede1
- Comparator
- phosphorus-free sidee4
- Setting
- sand culturee5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Increasing the phosphorus supply increased the nodules weight, nitrogenase activity, ureide content”
e4“and the other side was irrigated with a phosphorus-free nutrient solution (phosphorus-free side)”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
Claim · model-written summary
Increasing phosphorus supply increased the number of bacteroids and infected cells in soybean nodules on the phosphorus-application side.
- Taxon
- soybean
Glycine maxe5 - Treatment
- Increasing the phosphorus supplye1
- Outcome
- number of bacteroids, number of infected cellse2
- Direction
- increasede1
- Comparator
- phosphorus-free sidee4
- Setting
- sand culturee5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Increasing the phosphorus supply increased the nodules weight, nitrogenase activity, ureide content”
e2“number of bacteroids, number of infected cells”
e4“and the other side was irrigated with a phosphorus-free nutrient solution (phosphorus-free side)”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
Claim · model-written summary
Increasing phosphorus supply increased relative expression of nodule nitrogen fixation key genes in root nodules on the phosphorus-application side.
- Taxon
- soybean
Glycine maxe5 - Organ / stage
- root nodulese3
- Treatment
- Increasing the phosphorus supplye1
- Gene
- GmEXPB2, GmSPX5, nifH, nifD, nifK, GmALN1, GmACP1, GmUR5, GmPUR5, and GmHIUH5e3
- Outcome
- relative expression levels of nodule nitrogen fixation key genese3
- Direction
- increasede1
- Comparator
- phosphorus-free sidee4
- Setting
- sand culturee5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Increasing the phosphorus supply increased the nodules weight, nitrogenase activity, ureide content”
e3“relative expression levels of nodule nitrogen fixation key genes (GmEXPB2, GmSPX5, nifH, nifD, nifK, GmALN1, GmACP1, GmUR5, GmPUR5, and GmHIUH5) in root nodules on the phosphorus-application side”
e4“and the other side was irrigated with a phosphorus-free nutrient solution (phosphorus-free side)”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
Claim · model-written summary
Phosphorus-induced increases were more prominent on the phosphorus-application side than on the phosphorus-free side in soybean.
- Taxon
- soybean
Glycine maxe5 - Outcome
- phosphorus-induced increasese6
- Direction
- more prominente6
- Comparator
- phosphorus-free sidee4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e6“the phosphorus-induced increases were more prominent on the phosphorus-application side”
e4“and the other side was irrigated with a phosphorus-free nutrient solution (phosphorus-free side)”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
Claim · model-written summary
The authors conclude that phosphorus supply systematically regulates nodulation and nitrogen fixation in soybean.
- Taxon
- soybean
Glycine maxe7 - Treatment
- phosphorus supplye7
- Outcome
- nodulation and nitrogen fixatione7
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“phosphorus supply systematically regulates nodulation and nitrogen fixation in soybean”
Claim · model-written summary
Raising phosphorus supply from 1 to 31 mg/L increased nodule phosphorus content by 57.14–85.71% and 68.75–75.00%, respectively.
- Taxon
- soybean
Glycine maxe5 - Organ / stage
- nodulee0
- Treatment
- increased from 1 mg/L to 31 mg/Le1
- Outcome
- nodule phosphorus contente0
- Direction
- increasede0
- Value
- 57.14–85.71% and 68.75–75.00%e0
- Qualifier
- respectivelye0
- Comparator
- 1 mg/Le1
- Setting
- sand culturee5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“When the level of phosphorus supply was increased from 1 mg/L to 31 mg/L”
e0“nodule phosphorus content increased by 57.14–85.71% and 68.75–75.00%, respectively”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
Claim · model-written summary
Raising phosphorus supply from 1 to 31 mg/L increased ARA by 218.64–383.33%.
- Taxon
- soybean
Glycine maxe5 - Treatment
- increased from 1 mg/L to 31 mg/Le1
- Outcome
- ARAe2
- Direction
- increaseinferrede1
- Value
- 218.64–383.33%e2
- Comparator
- 1 mg/Le1
- Setting
- sand culturee5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“When the level of phosphorus supply was increased from 1 mg/L to 31 mg/L”
e2“ARA and SNA were 218.64–383.33% and 11.41–16.11%, respectively”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
Claim · model-written summary
Raising phosphorus supply from 1 to 31 mg/L increased SNA by 11.41–16.11%.
- Taxon
- soybean
Glycine maxe5 - Treatment
- increased from 1 mg/L to 31 mg/Le1
- Outcome
- SNAe2
- Direction
- increaseinferrede1
- Value
- 11.41–16.11%e2
- Comparator
- 1 mg/Le1
- Setting
- sand culturee5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“When the level of phosphorus supply was increased from 1 mg/L to 31 mg/L”
e2“ARA and SNA were 218.64–383.33% and 11.41–16.11%, respectively”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
Claim · model-written summary
Raising phosphorus supply from 1 to 31 mg/L increased ureide content by 118.18–156.44%.
- Taxon
- soybean
Glycine maxe5 - Treatment
- increased from 1 mg/L to 31 mg/Le1
- Outcome
- ureide contente3
- Direction
- increaseinferrede1
- Value
- 118.18–156.44%e3
- Comparator
- 1 mg/Le1
- Setting
- sand culturee5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“When the level of phosphorus supply was increased from 1 mg/L to 31 mg/L”
e3“ureide content was 118.18–156.44%”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
Claim · model-written summary
Raising phosphorus supply from 31 to 61 mg/L gave a smaller gain in root and nodule phosphorus content, ARA, SNA and ureide content than the 1 to 31 mg/L step, and a lower value for nodules.
- Taxon
- soybean
Glycine maxe5 - Organ / stage
- nodulese0
- Treatment
- increased from 31mg/L to 61mg/Le0
- Outcome
- regulation ability of root and nodule phosphorus content, ARA, SNA, and ureide contente0
- Direction
- lowe0
- Comparator
- increased from 1 mg/L to 31 mg/Le0
- Setting
- sand culturee5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“When the level of phosphorus supply was increased from 31mg/L to 61mg/L, the increase in the regulation ability of root and nodule phosphorus content, ARA, SNA, and ureide content were low for roots, and the value for nodules was lower than when the phosphorus level increased from 1 mg/L to 31 mg/L”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
Claim · model-written summary
A high phosphorus supply on one side of dual-root soybean significantly increased phosphorus content in the aboveground tissues, roots and nodules on both sides.
- Taxon
- soybean
Glycine maxe0 - Organ / stage
- aboveground tissuese0
- Treatment
- A high-concentration phosphorus supply on one sidee0
- Outcome
- phosphorus content in the aboveground tissues, as well as the roots and nodules on both sidese0
- Direction
- significantly increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“A high-concentration phosphorus supply on one side of a dual-root soybean plant significantly increased the phosphorus content in the aboveground tissues, as well as the roots and nodules on both sides”
Claim · model-written summary
The authors interpret that nodules on the phosphorus-free side are prioritized for phosphorus from the aboveground tissues to maintain their phosphorus content and functionality.
- Taxon
- soybean
Glycine maxe5 - Organ / stage
- nodulese0
- Outcome
- phosphorus content and functionalitye0
- Qualifier
- prioritized for receiving the phosphorus transported from the aboveground tissuese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“In the roots on the phosphorus-free side, the nodules were prioritized for receiving the phosphorus transported from the aboveground tissues to maintain their phosphorus content and functionality”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
Claim · model-written summary
Nodulation and nitrogen fixation traits on the phosphorus-application and phosphorus-free sides showed similar changing trends in soybean.
- Taxon
- soybean
Glycine maxe5 - Outcome
- changing trendse0
- Direction
- similare0
- Comparator
- phosphorus-free sidee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the phosphorus-application and phosphorus-free sides demonstrated similar changing trends”
e5“the seedling grafting technique was applied in this study to prepare dual-root soybean systems for a sand culture experiment”
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