Paper · 2026 · Food and Energy Security

Photosynthetic and Transcriptional Regulation Reveals Divergent Mechanisms of Active Adaptation in C 4 Maize and Passive Regulation in C 3 Rice Under Long‐Term Fluctuating Light

Wang‐Shun Zhang; Zi‐Mo Guo; Xiao‐Ling Li; et al.

16
claims
29
verbatim quotes
0
citations
3
research areas
✓
Open access · cc-by

Abstract

ABSTRACT Fluctuating light is a key factor limiting crop photosynthetic efficiency, with C 4 maize ( Zea mays ) and C 3 rice ( Oryza sativa ) crops exhibiting distinct acclimation responses. However, the systemic differences in physiological and transcriptional regulatory mechanisms between C 3 and C 4 crops under long‐term fluctuating light remain poorly understood. Herein, maize variety “Yunrui 408” and rice variety “Dianheyou 615,” both widely cultivated in Southwest China, were used to investigate responses to long‐term fluctuating light (FL) versus steady light (SL) via integrated analyses of phenotypic traits, photosynthetic physiology, transcriptomics, and protein–protein interaction (PPI) networks. Results showed that long‐term FL causes greater yield losses (> 30%) in rice, including reduced 100‐grain weight and shoot dry weight, whereas maize exhibited milder yield suppression (< 20%) and enhanced starch/sucrose accumulation in grains and leaves. Maize maintained stable photosystem function via a synergistic mechanism. A large plastoquinone (PQ) pool buffered electron transport fluctuations, while the transcription factor ZmMYB93 coordinated the upregulation of carbon storage ( ZmAGPS1 ), energy supply ( ZmGAPC3 ), and carbon assimilation ( ZmFBA3 ) pathways to sustain metabolic homeostasis. In contrast, rice showed decreased photosynthesis under long‐term FL, characterized by downregulated core photosynthetic genes ( OsPsaA , OsPsbA ) and ATP synthase genes ( OsAtpB ). This suppression was driven by the PEL‐GLK inhibitory module; subcellular localization assays confirmed that OsPEL1 is a nuclear‐resident protein that physically interacts with GLK transcription factors within the nucleus to repress the transcriptional activation of core photosynthetic genes. Rice also relied on passive cyclic electron flow (CEF) and exhibited increased dark respiration, forming a negative feedback loop that impaired carbon assimilation. Overall, C 4 crop maize exhibits stronger adaptability to long‐term FL than C 3 crop rice. It adopts an active adaptation strategy by integrating PQ pool‐mediated electron transport buffering and ZmMYB93‐coordinated nuclear transcriptional regulation. In contrast, rice adopts a passive and inefficient regulatory strategy, where OsPEL1 ‐mediated nuclear inhibition of GLK and enhanced dark respiration lead to photosynthetic apparatus damage and carbon assimilation suppression, ultimately resulting in severe yield loss. These findings provide a blueprint for improving C 3 crop light‐use efficiency by leveraging the active regulatory mechanisms identified in maize to mitigate yield losses in dynamic field environments.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by. Read at the source ↗

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Claims 16

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Long-term fluctuating light reduced rice yield by over 30%.

Treatment
long‐term FLe0
Outcome
yield lossese0
Direction
lossese0
Value
> 30%e0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“long‐term FL causes greater yield losses (> 30%) in rice”

✓ located verbatim in the source · abstract, characters 736–792 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Long-term fluctuating light reduced rice 100-grain weight and shoot dry weight.

Treatment
long‐term FLe2
Outcome
100‐grain weight and shoot dry weighte0
Direction
reducede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“including reduced 100‐grain weight and shoot dry weight”

✓ located verbatim in the source · abstract, characters 794–849 · show in text ↑
e1

“in rice”

✓ located verbatim in the source · abstract, characters 785–792 · show in text ↑
e2

“long‐term FL causes greater yield losses (> 30%) in rice”

✓ located verbatim in the source · abstract, characters 736–792 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Maize yield suppression under long-term fluctuating light was under 20%.

Taxon
maizeZea mayse0
Treatment
long‐term FLe1
Outcome
yield suppressione0
Direction
suppressione0
Value
< 20%e0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“maize exhibited milder yield suppression (< 20%)”

✓ located verbatim in the source · abstract, characters 859–907 · show in text ↑
e1

“responses to long‐term fluctuating light (FL) versus steady light (SL)”

✓ located verbatim in the source · abstract, characters 507–577 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Fluctuating light increased starch and sucrose accumulation in maize grains and leaves.

Taxon
maizeZea mayse0
Organ / stage
grains and leavese0
Treatment
long‐term FLe1
Outcome
starch/sucrose accumulatione0
Direction
enhancede0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“maize exhibited milder yield suppression (< 20%) and enhanced starch/sucrose accumulation in grains and leaves.”

✓ located verbatim in the source · abstract, characters 859–970 · show in text ↑
e1

“responses to long‐term fluctuating light (FL) versus steady light (SL)”

✓ located verbatim in the source · abstract, characters 507–577 · show in text ↑
Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

Maize maintained stable photosystem function under fluctuating light.

Taxon
MaizeZea mayse0
Outcome
stable photosystem functione0
Direction
stablee0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Maize maintained stable photosystem function via a synergistic mechanism.”

✓ located verbatim in the source · abstract, characters 971–1044 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

A large maize plastoquinone pool buffered electron transport fluctuations.

Taxon
MaizeZea mayse1
Compound
plastoquinonee0
Outcome
electron transport fluctuationse0
Direction
bufferede0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“A large plastoquinone (PQ) pool buffered electron transport fluctuations”

✓ located verbatim in the source · abstract, characters 1045–1117 · show in text ↑
e1

“Maize maintained stable photosystem function”

✓ located verbatim in the source · abstract, characters 971–1015 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

ZmMYB93 coordinated carbon storage, energy supply and carbon assimilation gene upregulation.

Taxon
MaizeZea mayse1
Gene
ZmMYB93e0
Outcome
upregulation of carbon storage ( ZmAGPS1 ), energy supply ( ZmGAPC3 ), and carbon assimilation ( ZmFBA3 ) pathwayse0
Direction
upregulatione0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“the transcription factor ZmMYB93 coordinated the upregulation of carbon storage ( ZmAGPS1 ), energy supply ( ZmGAPC3 ), and carbon assimilation ( ZmFBA3 ) pathways”

✓ located verbatim in the source · abstract, characters 1125–1288 · show in text ↑
e1

“Maize maintained stable photosystem function”

✓ located verbatim in the source · abstract, characters 971–1015 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Rice photosynthesis decreased under long-term fluctuating light.

Treatment
long‐term FLe0
Outcome
photosynthesise0
Direction
decreasede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“rice showed decreased photosynthesis under long‐term FL”

✓ located verbatim in the source · abstract, characters 1336–1391 · show in text ↑
Observationobservedoriginal resultnegativemeasured

Claim · model-written summary

Rice core photosynthetic genes OsPsaA and OsPsbA were downregulated.

Gene
OsPsaA , OsPsbAe0
Outcome
core photosynthetic genese0
Direction
downregulatede0

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“downregulated core photosynthetic genes ( OsPsaA , OsPsbA )”

✓ located verbatim in the source · abstract, characters 1410–1469 · show in text ↑
e1

“rice showed decreased photosynthesis”

✓ located verbatim in the source · abstract, characters 1336–1372 · show in text ↑
Observationobservedoriginal resultnegativemeasured

Claim · model-written summary

Rice ATP synthase gene OsAtpB was downregulated.

Gene
OsAtpBe0
Outcome
ATP synthase genese0
Direction
downregulatede1

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“ATP synthase genes ( OsAtpB )”

✓ located verbatim in the source · abstract, characters 1474–1503 · show in text ↑
e1

“downregulated core photosynthetic genes ( OsPsaA , OsPsbA ) and ATP synthase genes ( OsAtpB )”

✓ located verbatim in the source · abstract, characters 1410–1503 · show in text ↑
e2

“rice showed decreased photosynthesis”

✓ located verbatim in the source · abstract, characters 1336–1372 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

OsPEL1 localized to the nucleus and interacted with GLK transcription factors.

Gene
OsPEL1e0
Outcome
physically interacts with GLK transcription factors within the nucleuse0

Not reportedDirection

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“OsPEL1 is a nuclear‐resident protein that physically interacts with GLK transcription factors within the nucleus”

✓ located verbatim in the source · abstract, characters 1614–1726 · show in text ↑
e1

“rice showed decreased photosynthesis”

✓ located verbatim in the source · abstract, characters 1336–1372 · show in text ↑
Mechanismauthor interpretationoriginal resultnegativemeasured

Claim · model-written summary

The PEL-GLK module suppressed activation of core photosynthetic genes in rice.

Gene
OsPEL1e0
Outcome
transcriptional activation of core photosynthetic genese0
Direction
represse0

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“the PEL‐GLK inhibitory module; subcellular localization assays confirmed that OsPEL1 is a nuclear‐resident protein that physically interacts with GLK transcription factors within the nucleus to repress the transcriptional activation of core photosynthetic genes”

✓ located verbatim in the source · abstract, characters 1536–1797 · show in text ↑
e1

“rice showed decreased photosynthesis”

✓ located verbatim in the source · abstract, characters 1336–1372 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Rice relied on passive cyclic electron flow under fluctuating light.

Outcome
passive cyclic electron flow (CEF)e0

Not reportedDirection

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“Rice also relied on passive cyclic electron flow (CEF)”

✓ located verbatim in the source · abstract, characters 1799–1853 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Rice dark respiration increased under fluctuating light.

Treatment
long‐term FLe1
Outcome
dark respiratione0
Direction
increasede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“Rice also relied on passive cyclic electron flow (CEF) and exhibited increased dark respiration”

✓ located verbatim in the source · abstract, characters 1799–1894 · show in text ↑
e1

“responses to long‐term fluctuating light (FL) versus steady light (SL)”

✓ located verbatim in the source · abstract, characters 507–577 · show in text ↑
Mechanismauthor interpretationoriginal resultnegativemeasured

Claim · model-written summary

Enhanced dark respiration impaired rice carbon assimilation.

Outcome
carbon assimilatione0
Direction
impairede0
Qualifier
increased dark respiratione0

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“exhibited increased dark respiration, forming a negative feedback loop that impaired carbon assimilation”

✓ located verbatim in the source · abstract, characters 1858–1962 · show in text ↑
e1

“Rice also relied on passive cyclic electron flow”

✓ located verbatim in the source · abstract, characters 1799–1847 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Maize had stronger long-term fluctuating-light adaptability than rice.

Taxon
maizeZea mayse0
Outcome
adaptability to long‐term FLe0
Direction
strongere0
Comparator
ricee0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“C 4 crop maize exhibits stronger adaptability to long‐term FL than C 3 crop rice.”

✓ located verbatim in the source · abstract, characters 1973–2054 · show in text ↑

light · physiology: photosynthesis · Oryza sativa (rice)Results differ

47 papers: 24 positive, 23 negative or no effect

22 papers in the same direction as this paper · 24 in the opposite direction

  1. positive
  2. negative
  3. positive

43 papers more in this group

See all 47 papers →

light · yield · Oryza sativa (rice)Results differ

68 papers: 36 positive, 32 negative or no effect

31 papers in the same direction as this paper · 36 in the opposite direction

  1. positive
  2. positive
  3. positive

64 papers more in this group

See all 68 papers →

light · control · Zea mays (maize)Positive and negative results

2 papers: 1 positive, 1 negative or no effect

1 in the opposite direction

  1. positive
See all 2 papers →

light · physiology: respiration · Oryza sativa (rice)Positive and negative results

3 papers: 1 positive, 2 negative or no effect

1 paper in the same direction as this paper · 1 in the opposite direction

  1. positive
  2. no effect
See all 3 papers →

light · quality: sugar starch · Zea mays (maize)Positive and negative results

11 papers: 6 positive, 5 negative or no effect

5 papers in the same direction as this paper · 5 in the opposite direction

  1. no effect
  2. no effect
  3. negative

7 papers more in this group

See all 11 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.