Abstract
Phytoremediation technology can help achieve moderate cost and considerable effect with respect to the remediation of heavy metal (HM) pollution in soil and water. Many previous studies have suggested the role of nitrogen (N) in the alleviation of effects of HM on plants. Herein, we sought to determine the molecular mechanisms by which additional N supplementation mitigates cadmium (Cd) toxicity in poplars using a combination of physiological, transcriptomic and phosphoproteomic analyses. The application of N can alleviate the toxicity of Cd to Populus by reducing chlorophyll degradation, maintaining the stability of ions inside and outside the cell membrane and increasing the soluble sugar content. Plant samples from the control, Cd stress and Cd_N treatments were used for an integrated analysis of the transcriptome, as well as for phosphoproteomics analysis. Moreover, 1314 differentially expressed genes and 119 differentially expressed kinase genes were discovered. Application of additional N under Cd stress promoted the phosphorylation process. Furthermore, 51 significantly enriched phosphorylated protein sites and 23 differentially expressed kinases were identified using phosphoproteomic and proteomic analyses. Importantly, transcriptomic and phosphoproteomic analyses jointly determined that the application of N could activate corresponding gene expression [UDP-glucose-dehydrogenase (UGD), GAUT, PME, pectin lyase, UDP-glucose-pyrophosphorylase 2 (UGP2), sucrose phosphate synthase (SPS), SUS and SPP2] and protein phosphorylation (UGP2 and SPS) in the sugar and starch synthesis pathways, which promoted the synthesis of sucrose and soluble sugar and subsequently alleviated the damage caused by Cd.
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Claims 11
Claim · model-written summary
Nitrogen reduced cadmium toxicity in Populus by reducing chlorophyll degradation.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The application of N can alleviate the toxicity of Cd to Populus by reducing chlorophyll degradation”
Claim · model-written summary
Nitrogen maintained ion stability inside and outside the cell membrane under cadmium stress.
- Treatment
- Nt0
- Outcome
- stability of ions inside and outside the cell membranee0
- Direction
- maintaininge0
- Comparator
- cadmiumt0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“maintaining the stability of ions inside and outside the cell membrane”
t0“External application of nitrogen alleviates toxicity of cadmium on poplars via starch and sucrose metabolism”
Claim · model-written summary
Nitrogen increased soluble sugar content under cadmium stress.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“increasing the soluble sugar content”
t0“External application of nitrogen alleviates toxicity of cadmium on poplars via starch and sucrose metabolism”
Claim · model-written summary
The analysis identified 1314 differentially expressed genes.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“1314 differentially expressed genes and 119 differentially expressed kinase genes were discovered”
t0“External application of nitrogen alleviates toxicity of cadmium on poplars via starch and sucrose metabolism”
Claim · model-written summary
The analysis identified 119 differentially expressed kinase genes.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“1314 differentially expressed genes and 119 differentially expressed kinase genes were discovered”
t0“External application of nitrogen alleviates toxicity of cadmium on poplars via starch and sucrose metabolism”
Claim · model-written summary
Additional nitrogen under cadmium stress promoted phosphorylation.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Application of additional N under Cd stress promoted the phosphorylation process”
t0“poplars”
Claim · model-written summary
The analyses identified 51 significantly enriched phosphorylated protein sites.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“51 significantly enriched phosphorylated protein sites and 23 differentially expressed kinases were identified”
t0“poplars”
Claim · model-written summary
The analyses identified 23 differentially expressed kinases.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“51 significantly enriched phosphorylated protein sites and 23 differentially expressed kinases were identified”
t0“poplars”
Claim · model-written summary
Nitrogen activated expression of genes in sugar and starch synthesis pathways.
- Treatment
- Ne0
- Gene
- UDP-glucose-dehydrogenase (UGD), GAUT, PME, pectin lyase, UDP-glucose-pyrophosphorylase 2 (UGP2), sucrose phosphate synthase (SPS), SUS and SPP2e0
- Outcome
- gene expressione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the application of N could activate corresponding gene expression [UDP-glucose-dehydrogenase (UGD), GAUT, PME, pectin lyase, UDP-glucose-pyrophosphorylase 2 (UGP2), sucrose phosphate synthase (SPS), SUS and SPP2]”
t0“poplars”
Claim · model-written summary
Nitrogen activated UGP2 and SPS phosphorylation in sugar and starch synthesis pathways.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“protein phosphorylation (UGP2 and SPS) in the sugar and starch synthesis pathways, which promoted the synthesis of sucrose and soluble sugar and subsequently alleviated the damage caused by Cd”
t0“poplars”
Claim · model-written summary
Pathway changes promoted sucrose and soluble sugar synthesis and alleviated cadmium damage.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“which promoted the synthesis of sucrose and soluble sugar and subsequently alleviated the damage caused by Cd”
t0“poplars”
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See all 5 papers →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.