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.
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Claims 16
Claim · model-written summary
Long-term fluctuating light reduced rice yield by over 30%.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“long‐term FL causes greater yield losses (> 30%) in rice”
Claim · model-written summary
Long-term fluctuating light reduced rice 100-grain weight and shoot dry weight.
- Taxon
- rice
Oryza sativae1
- 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.
Verbatim
e0“including reduced 100‐grain weight and shoot dry weight”
e1“in rice”
e2“long‐term FL causes greater yield losses (> 30%) in rice”
Claim · model-written summary
Maize yield suppression under long-term fluctuating light was under 20%.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“maize exhibited milder yield suppression (< 20%)”
e1“responses to long‐term fluctuating light (FL) versus steady light (SL)”
Claim · model-written summary
Fluctuating light increased starch and sucrose accumulation in maize grains and leaves.
- 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.

Verbatim
e0“maize exhibited milder yield suppression (< 20%) and enhanced starch/sucrose accumulation in grains and leaves.”
e1“responses to long‐term fluctuating light (FL) versus steady light (SL)”
Claim · model-written summary
Maize maintained stable photosystem function under fluctuating light.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Maize maintained stable photosystem function via a synergistic mechanism.”
Claim · model-written summary
A large maize plastoquinone pool buffered electron transport fluctuations.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“A large plastoquinone (PQ) pool buffered electron transport fluctuations”
e1“Maize maintained stable photosystem function”
Claim · model-written summary
ZmMYB93 coordinated carbon storage, energy supply and carbon assimilation gene upregulation.
- 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.

Verbatim
e0“the transcription factor ZmMYB93 coordinated the upregulation of carbon storage ( ZmAGPS1 ), energy supply ( ZmGAPC3 ), and carbon assimilation ( ZmFBA3 ) pathways”
e1“Maize maintained stable photosystem function”
Claim · model-written summary
Rice photosynthesis decreased under long-term fluctuating light.
- Taxon
- rice
Oryza sativae0
- Treatment
- long‐term FLe0
- Outcome
- photosynthesise0
- Direction
- decreasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“rice showed decreased photosynthesis under long‐term FL”
Claim · model-written summary
Rice core photosynthetic genes OsPsaA and OsPsbA were downregulated.
- Taxon
- rice
Oryza sativae1
- Gene
- OsPsaA , OsPsbAe0
- Outcome
- core photosynthetic genese0
- Direction
- downregulatede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“downregulated core photosynthetic genes ( OsPsaA , OsPsbA )”
e1“rice showed decreased photosynthesis”
Claim · model-written summary
Rice ATP synthase gene OsAtpB was downregulated.
- Taxon
- rice
Oryza sativae2
- Gene
- OsAtpBe0
- Outcome
- ATP synthase genese0
- Direction
- downregulatede1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“ATP synthase genes ( OsAtpB )”
e1“downregulated core photosynthetic genes ( OsPsaA , OsPsbA ) and ATP synthase genes ( OsAtpB )”
e2“rice showed decreased photosynthesis”
Claim · model-written summary
OsPEL1 localized to the nucleus and interacted with GLK transcription factors.
- Taxon
- rice
Oryza sativae1
- 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.
Verbatim
e0“OsPEL1 is a nuclear‐resident protein that physically interacts with GLK transcription factors within the nucleus”
e1“rice showed decreased photosynthesis”
Claim · model-written summary
The PEL-GLK module suppressed activation of core photosynthetic genes in rice.
- Taxon
- rice
Oryza sativae1
- Gene
- OsPEL1e0
- Outcome
- transcriptional activation of core photosynthetic genese0
- Direction
- represse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
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”
e1“rice showed decreased photosynthesis”
Claim · model-written summary
Rice relied on passive cyclic electron flow under fluctuating light.
- Taxon
- Rice
Oryza sativae0
- Outcome
- passive cyclic electron flow (CEF)e0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Rice also relied on passive cyclic electron flow (CEF)”
Claim · model-written summary
Rice dark respiration increased under fluctuating light.
- Taxon
- Rice
Oryza sativae0
- Treatment
- long‐term FLe1
- Outcome
- dark respiratione0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Rice also relied on passive cyclic electron flow (CEF) and exhibited increased dark respiration”
e1“responses to long‐term fluctuating light (FL) versus steady light (SL)”
Claim · model-written summary
Enhanced dark respiration impaired rice carbon assimilation.
- Taxon
- Rice
Oryza sativae1
- Outcome
- carbon assimilatione0
- Direction
- impairede0
- Qualifier
- increased dark respiratione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“exhibited increased dark respiration, forming a negative feedback loop that impaired carbon assimilation”
e1“Rice also relied on passive cyclic electron flow”
Claim · model-written summary
Maize had stronger long-term fluctuating-light adaptability than rice.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“C 4 crop maize exhibits stronger adaptability to long‐term FL than C 3 crop rice.”
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