Paper · 2023 · PLoS ONE

Changes in growth, physiology, and photosynthetic capacity of spinach (Spinacia oleracea L.) under different nitrate levels

Kangning Han; Jing Zhang; Cheng Wang; et al.

14
claims
23
verbatim quotes
21
citations
1
research areas
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Open access · cc-by

Abstract

Nitrate content is an essential indicator of the quality of vegetables but can cause stress at high levels. This study aimed to elucidate the regulatory mechanisms of nitrate stress tolerance in spinach (Spinacia oleracea L.). We studied the effects of exogenous application of 15 (control), 50, 100, 150, 200, and 250 mM NO3- on spinach growth, physiology, and photosynthesis. The results showed that all the nitrate treatments inhibited the growth of the aerial parts of spinach compared to the control. In contrast, low nitrate levels (50 and 100 mM) promoted spinach root formation, but this effect was inhibited at high levels (150, 200, and 250 mM). Treatment with 150 mM NO3- significantly decreased the root growth vigor. Low nitrate levels increased the chlorophyll content in spinach leaves, whereas high levels had the opposite effect. High nitrate levels also weakened the net photosynthetic rate (Pn), the actual photochemical efficiency of PSII Y(II), and increased non-photochemical quenching (NPQ), reducing photosynthetic performance. Nitrate stress increased the activity of nitrate reductase (NR) and promoted the accumulation of nitrate in spinach leaves, exceeding the health-tolerance limit for nitrate in vegetables, highlighting the necessity of mitigating nitrate stress to ensure food safety. Starting with the 150 mM NO3- treatment, the proline and malondialdehyde content in spinach leaves and roots increased significantly as the nitrate levels increased. Treatment with 150 mM NO3- significantly increased soluble protein and flavonoid contents, while the activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), and ascorbate peroxidase (APX) were significantly reduced in leaves. However, spinach could resist nitrate stress by regulating the synthesis of osmoregulatory substances such as proline, thus showing some nitrate tolerance. These results provide insights into the physiological regulatory mechanisms of nitrate stress tolerance and its mitigation in spinach, an essential vegetable crop.

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Claims 14

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Nitrate treatments inhibited spinach aerial growth versus the control.

Treatment
nitrate treatmentse0
Outcome
growth of the aerial partse0
Direction
inhibitede0
Comparator
controle0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“all the nitrate treatments inhibited the growth of the aerial parts of spinach compared to the control”

✓ located verbatim in the source · abstract, characters 402–504 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Low nitrate levels promoted spinach root formation.

Treatment
low nitrate levels (50 and 100 mM)e0
Outcome
root formatione0
Direction
promotede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“low nitrate levels (50 and 100 mM) promoted spinach root formation”

✓ located verbatim in the source · abstract, characters 519–585 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

High nitrate levels inhibited spinach root formation.

Treatment
high levels (150, 200, and 250 mM)e0
Outcome
this effecte0
Direction
inhibitede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“this effect was inhibited at high levels (150, 200, and 250 mM)”

✓ located verbatim in the source · abstract, characters 591–654 · show in text ↑
t0

“spinach (Spinacia oleracea L.)”

✓ located verbatim in the source · title, characters 62–92 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

The 150 mM nitrate treatment significantly decreased spinach root growth vigor.

Treatment
150 mM NO3-e0
Outcome
root growth vigore0
Direction
decreasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Treatment with 150 mM NO3- significantly decreased the root growth vigor”

✓ located verbatim in the source · abstract, characters 656–728 · show in text ↑
t0

“spinach (Spinacia oleracea L.)”

✓ located verbatim in the source · title, characters 62–92 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Low nitrate increased spinach leaf chlorophyll content.

Treatment
Low nitrate levelse0
Outcome
chlorophyll contente0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Low nitrate levels increased the chlorophyll content in spinach leaves”

✓ located verbatim in the source · abstract, characters 730–800 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

High nitrate decreased spinach leaf chlorophyll content.

Treatment
high levelse0
Outcome
chlorophyll contente1
Direction
opposite effecte0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“high levels had the opposite effect”

✓ located verbatim in the source · abstract, characters 810–845 · show in text ↑
e1

“chlorophyll content in spinach leaves”

✓ located verbatim in the source · abstract, characters 763–800 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

High nitrate weakened spinach net photosynthetic rate and PSII efficiency.

Treatment
High nitrate levelse0
Outcome
net photosynthetic rate (Pn), the actual photochemical efficiency of PSII Y(II)e0
Direction
weakenede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“High nitrate levels also weakened the net photosynthetic rate (Pn), the actual photochemical efficiency of PSII Y(II)”

✓ located verbatim in the source · abstract, characters 847–964 · show in text ↑
t0

“spinach (Spinacia oleracea L.)”

✓ located verbatim in the source · title, characters 62–92 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

High nitrate increased spinach non-photochemical quenching.

Treatment
High nitrate levelse0
Outcome
non-photochemical quenching (NPQ)e0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“High nitrate levels also weakened the net photosynthetic rate (Pn), the actual photochemical efficiency of PSII Y(II), and increased non-photochemical quenching (NPQ)”

✓ located verbatim in the source · abstract, characters 847–1013 · show in text ↑
t0

“spinach (Spinacia oleracea L.)”

✓ located verbatim in the source · title, characters 62–92 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Nitrate stress increased nitrate reductase activity in spinach.

Treatment
Nitrate stresse0
Gene
nitrate reductase (NR)e0
Outcome
activity of nitrate reductase (NR)e0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Nitrate stress increased the activity of nitrate reductase (NR)”

✓ located verbatim in the source · abstract, characters 1052–1115 · show in text ↑
t0

“spinach (Spinacia oleracea L.)”

✓ located verbatim in the source · title, characters 62–92 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Nitrate stress promoted nitrate accumulation in spinach leaves beyond the vegetable health tolerance limit.

Treatment
nitratee0
Outcome
accumulation of nitrate in spinach leavese0
Direction
promotede0
Qualifier
exceeding the health-tolerance limit for nitrate in vegetablese0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“promoted the accumulation of nitrate in spinach leaves, exceeding the health-tolerance limit for nitrate in vegetables”

✓ located verbatim in the source · abstract, characters 1120–1238 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

From 150 mM nitrate, proline and malondialdehyde content in spinach leaves and roots increased significantly as nitrate rose.

Organ / stage
leaves and rootse0
Treatment
150 mM NO3- treatmente0
Outcome
proline and malondialdehyde contente0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Starting with the 150 mM NO3- treatment, the proline and malondialdehyde content in spinach leaves and roots increased significantly as the nitrate levels increased”

✓ located verbatim in the source · abstract, characters 1319–1483 · show in text ↑
Intervention effectobservedoriginal resultsignificant (as stated)measured

Claim · model-written summary

The 150 mM nitrate treatment significantly increased spinach soluble protein and flavonoid contents.

Treatment
150 mM NO3-e0
Outcome
soluble protein and flavonoid contentse0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Treatment with 150 mM NO3- significantly increased soluble protein and flavonoid contents”

✓ located verbatim in the source · abstract, characters 1485–1574 · show in text ↑
t0

“spinach (Spinacia oleracea L.)”

✓ located verbatim in the source · title, characters 62–92 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

The 150 mM nitrate treatment significantly reduced four antioxidant enzyme activities in spinach leaves.

Organ / stage
leavese0
Treatment
150 mM NO3-e1
Outcome
activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), and ascorbate peroxidase (APX)e0
Direction
reducede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“the activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), and ascorbate peroxidase (APX) were significantly reduced in leaves”

✓ located verbatim in the source · abstract, characters 1582–1729 · show in text ↑
e1

“Treatment with 150 mM NO3- significantly increased soluble protein and flavonoid contents”

✓ located verbatim in the source · abstract, characters 1485–1574 · show in text ↑
t0

“spinach (Spinacia oleracea L.)”

✓ located verbatim in the source · title, characters 62–92 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivebasis n/a

Claim · model-written summary

The authors interpret proline regulation as contributing to spinach nitrate tolerance.

Outcome
nitrate tolerancee0
Qualifier
regulating the synthesis of osmoregulatory substances such as prolinee0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Spinacia oleracea
Spinacia oleracea · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“spinach could resist nitrate stress by regulating the synthesis of osmoregulatory substances such as proline, thus showing some nitrate tolerance”

✓ located verbatim in the source · abstract, characters 1740–1885 · show in text ↑

nitrogen · physiology: pigments · Spinacia oleracea (spinach)Results differ

19 papers: 16 positive, 3 negative or no effect

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  1. no effect
  2. negative
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nitrogen · physiology: photosynthesis · Spinacia oleracea (spinach)Positive and negative results

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nitrogen · nutrient: content · Spinacia oleracea (spinach)Positive and negative results

35 papers: 21 positive, 14 negative or no effect

13 papers in the same direction as this paper · 21 in the opposite direction

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nitrogen · growth · Spinacia oleracea (spinach)Positive and negative results

40 papers: 28 positive, 12 negative or no effect

27 papers in the same direction as this paper · 11 in the opposite direction · 1 reporting no difference from an alternative

  1. negative
  2. positive
  3. positive

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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.