Rongcheng Lin’s research while affiliated with Chinese Academy of Sciences and other places

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Publications (94)


Advances in photosynthesis research: Unlocking the potential for food security, renewable energy, and environmental sustainability
  • Article
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April 2025

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6 Reads

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Rongcheng Lin
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Spectral values of treatments, red LEDs peak at 660 nm, blue LEDs peak at 460 nm. (a) BC, blue LEDs constant irradiation. (b) B5/B15/B30/B45, blue LEDs on. (c) B5/B15/B30/B45, blue LEDs off.
Effects of intermittent supplemental irradiation with blue light on lettuce morphology. (a) Representative shapes of lettuce under five experimental lighting treatments. (b) Dynamic curves of numbers of lettuce heads (means of three replicates) per days of light treatment. (c) Number of treatment days for half of lettuce plants to reach the heading stage under different treatments. (d) Crown widths, (e) root lengths, (f) leaf indexes, and (h) petiole bending angles of lettuce under five experimental lighting treatments at harvest. (g) Lateral anatomy of a lettuce head, with red lines indicating the approximate position of the leaves used to collect leaf data. The values are the mean ± SD of three replicates. Means denoted by different letters are significantly different according to Duncan’s multiple range test at the 0.05 significance level.
Effects of intermittent supplemental irradiation with blue light on hormone levels in lettuce. (a) Heat map of hormone contents. BC.L, B5.L, B15.L, B30.L, and B45.L indicate hormone contents in leaves, and BC.P, B5.P, B15.P, B30.P, and B45.P indicate hormone contents in petioles. Data was standardized by unit variance scaling (UV), and means of three replicates are shown. (b) Correlation of different leaf hormones with leaf indexes. (c) Correlation of different petiole hormones with petiole bending angles.
Effects of intermittent supplemental irradiation with blue light on the lettuce transcriptome. (a) MA plot of expression differences between BC and B15, with each point in the Figure representing a different gene. (b) Heatmap of differentially expressed genes between BC and B15, with expression values represented as mean log2(TPM + 1) of three replicates. (c) Heatmap of differentially expressed cytochrome family genes between BC and B15, with expression values represented as mean log2(TPM + 1) of three replicates. KEGG enrichment plots for differentially expressed genes (d) upregulated and (e) downregulated between BC and B15.KOG enrichment plots for differentially expressed genes (f) upregulated and (g) downregulated between BC and B15. Rich factor refers to the ratio of the number of differentially expressed genes located under this term to the total number of genes located in this pathway among all annotated genes. The larger the rich factor, the greater the enrichment.
Effects of intermittent supplemental irradiation with blue light on relative gene expression levels of light response and hormone metabolism genes in lettuce leaves at a fixed time point, the expression levels are based on the reference of the expression at 14:00 in the BC treatment. Relative expression levels of LsCRY1 (a), LsPHOT1 (b), LsHSP70 3 (c), LsRUP2 (d), LsHY5 (e), LsYUC5 (f), LsNCED2 (g), LsGA3OX1 (h), and LsIPT1 (i). The values are the mean ± SD of three replicates. Means denoted by different letters are significantly different according to Duncan’s multiple range test at the 0.05 significance level.

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Intermittent Supplemental Irradiation With Blue Light Promotes Leafy Heads in Lettuce

April 2025

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28 Reads

Yanke Liu

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Yiqun Chen

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Rongcheng Lin

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Red and blue lights are the most effective spectral components absorbed by plants and are typically applied in a constant spectrum in plant factory agricultural practices. Research and application of non‐constant light modes are relatively rare. In this study, we examined the impacts of varying blue light photon intensity and durations at 5‐, 15‐, 30‐, and 45‐min intervals of intermittent exposure on the growth and development of lettuce (Lactuca sativa) in plant factories while maintaining a constant red light photon flux and daily light integral (DLI). Compared to the constant light condition, intermittent blue light irradiation treatments accelerated the emergence of the leafy head trait in lettuce without compromising photosynthetic capacity and biomass. Specifically, intermittent blue light treatment with 15‐min intervals led to a reduction of 8 days in the time needed for half of the lettuce plants to reach the heading stage. Furthermore, the petiole bending angle in treated lettuce was just 70.2% of that observed under constant light conditions, with strong correlations between multiple hormone levels and bending angle in petioles. Transcriptome sequencing analyses revealed significant differential expression of signaling‐related genes between constant and intermittent blue light treatments. The transient and dynamic expression of light‐responsive and hormone metabolism‐related genes indicated that 15‐min intermittent blue light exposure better maintained the rhythmic differential expression of response genes, leading to different hormone accumulations and consequently accelerating the development of leafy heads.


The phytochrome B signaling regulates salt-mediated seedling growth in the dark

March 2025

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5 Reads

Plant and Cell Physiology

Light is an essential environmental factor that facilitates the robust upward growth of post-germinative seedlings emerging from buried seeds that is partly mediated by the photoreceptors. Salinity stress hampers plant growth and development and reduces yield. However, the involvement and regulatory role of photoreceptors and light signaling factors to salt stress are largely unknown. Here, we report that mutants of the phytochrome B (phyB) photoreceptor showed reduced sensitivity to salt-inhibited hypocotyl elongation in darkness, and that PHYTOCHROME-INTERACTING FACTOR 3 (PIF3) acts downstream of phyB in regulating this process in Arabidopsis thaliana. We also show that SALT OVERLY SENSITIVE 2 (SOS2) regulates phyB protein accumulation under salt stress in darkness. Surprisingly, salt treatment induces phyB nuclear body formation in darkness. Moreover, we found that the phosphorylation at residue Ser-86 of phyB is essential for its function, and the scaffold protein 14-3-3κ is involved in the regulation of phyB under salt stress in darkness. Taken together, our study reveals a regulatory role of the phyB–PIF3 module in mediating post-germination growth in darkness in response to salt stress.


Light regulates seed dormancy through FHY3-mediated activation of ACC OXIDASE 1 in Arabidopsis

Plant Molecular Biology

Seed dormancy enables plants to delay germination until conditions are favorable for the survival of the next generation. Seed dormancy and germination are controlled by a combination of external and internal signals, in which light and ethylene act as critical regulators. However, how light and ethylene are interlinked to control these two processes remains to be investigated. Here, we show that ethylene and its precursor, 1-aminocyclopropane-1-carboxylic acid (ACC), promote seed germination under light. Light facilitates the conversion of ACC to ethylene, in which phytochrome B (phyB) and FAR-RED ELONGATED HYPOCOTYL3 (FHY3) are functionally required. ACC oxidases (ACOs) catalyze the conversion of ACC to ethylene, among which ACO1 is specifically and predominantly expressed in imbibed seeds. Ethylene induces FHY3 protein accumulation in imbibed seeds, whereby FHY3 directly binds to ACO1 promoter and specifically mediates light-promoted ACO1 expression. Light promotes ACO1 protein accumulation. Overexpression of ACO1 significantly promotes seed germination, and almost completely restores the dormant defect of fhy3 loss-of-function mutants. In summary, this study reveals an ethylene-responsive regulatory cascade of phyB-FHY3-ACO1 that integrates external light input with internal factors to regulate seed dormancy and germination.


The aerobic chlorophyll biosynthesis pathway
Overall reactions from Protoporphyrin IX (PPIX) to Chlorophyll (Chl) a and other Chls. The chemical changes in the molecular structures are marked in red at each step. 2OG, 2‐oxoglutarate; ADP, adenosine diphosphate; ATP, adenosine triphosphate; NADP⁺, nicotinamide adenine dinucleotide phosphate; NADPH, the reduced form of NADP⁺; Pi, inorganic phosphate; SAH, S‐adenosyl‐l‐homocysteine; SAM, S‐adenosyl‐l‐methionine. *The enzymatic steps for which non‐homologous isozymes are known in different phototrophs. †ChlF‐containing PSII (super‐rogue PSII), instead of an isolated ChlF subunit, catalyzes the formation of Chl f (Trinugroho et al., 2020).
The transcriptional regulatory network that controls tetrapyrrole biosynthesis (TBS) levels in response to light and hormonal signals
Light and hormones co‐regulate TBS gene expression. GLK1/GLK2 (Golden‐Like 1/2), GNC (GATA nitrate‐inducible carbon metabolism‐involved) and GNL (cytokinin‐induced GATA1/GNC‐like) directly activate TBS genes in response to light and hormones. The relative levels of GLK1/2, GNC and GNL are strictly regulated at both the transcriptional and protein levels by light and hormonal signaling factors. These include negative interactions with phytochrome‐interacting factors (PIFs), RVE1 (Reveille1), BPG4 (BRZ‐insensitive‐pale green (BPG)), and BZR1 (Brassinazole Resistant 1) in the dark, and positive contacts with HY5 (Elongated Hypocotyl 5), ARFs (Auxin Response Factors), and ARR10/12 (B‐type Arabidopsis response regulators) in the light. Under dark conditions, COP1 and PIFs play key roles in inhibiting the transcription of TBS genes. Conversely, PIFs recruit EIN3 (Ethylene insensitive 3), DELLA (the negative regulator of GA), SCL27 (miR171‐targeted Scarecrow‐like protein) and some chromatin‐remodeling elements, such as HDA15 (Histone Deacetylase 15) and BRM (an SWI2/SNF2 chromatin‐remodeling ATPase), to inhibit the transcription of most TBS genes, but activate the expression of PORA and PORB. Conversely, COP1 inhibits the transcriptional activation of TBS genes by HY5 by promoting its degradation. Upon illumination, the photoreceptors phytochrome (PHY) and cryptochrome (CRY) trigger the activity of HY5 by inhibiting COP1. HY5 then activates the transcription of GLK1/GLK2, GNC, and GNL and most TBS genes; FHY3 (Far‐red elongated Hypocotyl 3) directly activates the transcription of HEMB1. In addition, the auxin factors (ARFs) and the CK factor ARR10/12 induce the transcription of GLK1/GLK2, GNC, and GNL to promote chlorophyll (Chl) synthesis. The direct targets of these factors are shown in Figure 2, and other indirect targets are listed in Table 1.
The post‐translational regulatory network that controls the functions of tetrapyrrole biosynthesis (TBS) enzymes
Post‐translational regulation of TBS enzymes encompasses several modifiers of specific amino‐acid residues, auxiliary factors and chaperones, all of which are involved in the control of protein stability and enzyme activity. The precursors of TBS enzymes are transported into chloroplasts by the TOC–TIC complex, with the aid of molecular chaperones that include the cytoplasmic chaperones Hsp90 and Hsp70 (Heat Shock Proteins 90 and 70) and the chloroplast chaperone cpHsp70 (chloroplast stromal Hsp70). Interactions of the chaperones ClpC1, ClpC2, and ClpD with GluTR (Glutamyl‐tRNA Reductase) or CAO (Chlorophyll a oxygenase) lead to their degradation by Clp protease. CPP1/CDF1 (Chaperone of POR protein 1/Cell growth deficient factor 1) enhances the stability of POR. cpSRP43 (chloroplast Signal Recognition Particle 43) interacts with GluTR, GUN4, and GUN5 and prevents them from aggregating during heat stress. Heat‐induced dissociation of cpSRP43 from cpSRP54 enables cpSRP43 to protect these TBS enzymes under heat stress. MORF2 and 9 (Multiple organellar RNA‐editing factors) function as chaperones to inhibit the aggregation of PORB, and promote MgCh activity through interaction with GUN4. FLU (Fluorescent in blue light) and GBP (GluTR‐binding protein) act as an inhibitor and a stabilizer of GluTR, respectively, through their interactions with GluTR. LCAA interacts with CHL27 and promotes its enzyme activity. BCM1 and 2 (Balance of Chlorophyll Metabolism) optimize chlorophyll (Chl) biosynthesis by stimulating MgCh activity via their interactions with GUN4, and attenuating Chl degradation by inducing the degradation of SGR1 (Stay‐green 1). FC2 (Ferrochelatase 2) interacts with POR and FLU to stabilize the FLU‐mediated GluTR‐inactivation complex and reduce the synthesis of heme for plastid‐localized heme‐dependent proteins. PCD8 (Programmed Cell Death 8) interacts with HEMC, CHLD, and PORC to stimulate their proteolysis. In addition, heme and Pchlide contribute to the inhibition of GluTR activity, and Chl b causes the feedback‐induced breakdown of CAO (Yamasato et al., 2005; Herbst et al., 2019). The modifications of amino acids of TBS enzymes by thiol‐based redox switches, S‐nitrosylation, and phosphorylation have been summarized in an earlier review (Herbst et al., 2019) and are not shown in this figure.
Tetrapyrrole signaling and transport
The maintenance of chloroplast function in photosynthetic eukaryotes requires coordinated regulation between the nuclear genome and the plastid genome. Tetrapyrroles, such as heme and bilin, play crucial roles in plastid‐to‐nucleus retrograde signaling in plants and algae. Heme regulates the expression of nuclear genes, including PhANGs, HSP70A, and others. In Chlamydomonas, bilin serves as a chloroplast retrograde signal that regulates the expression of nuclear genes involved in ROS detoxification and genes coding for O2‐dependent enzymes. In addition, singlet oxygen (¹O2) produced by tetrapyrrole intermediates mediates retrograde signaling through the EX1/2 pathways, and regulates the expression of ¹O2‐responsive genes. Specific membrane transporter proteins for tetrapyrroles have not yet been identified. Heme can be transported from the chloroplast to the cytosol by TSPO, GSTUs, or other unidentified transport proteins. Heme is also transported to other organelles and interacts with heme‐binding proteins in the nucleus, endoplasmic reticulum, Golgi apparatus, mitochondria, and peroxisomes. Heme released from mitochondrial hemoproteins may also be transported to the cytosol. During chlorophyll (Chl) degradation, Chl breakdown products, such as pFCC and Hydroxy‐pFCC, can be transported into vacuoles by the vacuolar ABC transport proteins AtMRP1–3.
Regulatory and retrograde signaling networks in the chlorophyll biosynthetic pathway

January 2025

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93 Reads

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2 Citations

Plants, algae and photosynthetic bacteria convert light into chemical energy by means of photosynthesis, thus providing food and energy for most organisms on Earth. Photosynthetic pigments, including chlorophylls (Chls) and carotenoids, are essential components that absorb the light energy necessary to drive electron transport in photosynthesis. The biosynthesis of Chl shares several steps in common with the biosynthesis of other tetrapyrroles, including siroheme, heme and phycobilins. Given that many tetrapyrrole precursors possess photo‐oxidative properties that are deleterious to macromolecules and can lead to cell death, tetrapyrrole biosynthesis (TBS) requires stringent regulation under various developmental and environmental conditions. Thanks to decades of research on model plants and algae, we now have a deeper understanding of the regulatory mechanisms that underlie Chl synthesis, including (i) the many factors that control the activity and stability of TBS enzymes, (ii) the transcriptional and post‐translational regulation of the TBS pathway, and (iii) the complex roles of tetrapyrrole‐mediated retrograde signaling from chloroplasts to the cytoplasm and the nucleus. Based on these new findings, Chls and their derivatives will find broad applications in synthetic biology and agriculture in the future.


Experimental lighting conditions.
Intermittent Supplementation with Far-Red Light Accelerates Leaf and Bud Development and Increases Yield in Lettuce

January 2025

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29 Reads

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1 Citation

Supplementation with far-red light in controlled environment agriculture production can enhance yield by triggering the shade avoidance syndrome. However, the effectiveness of this yield enhancement can be further improved through intermittent far-red light supplementation. In this study, the effects are explored of varying far-red light photon intensities and intermittent exposure durations—specifically at 5, 15, 30, and 45 min intervals—on the growth and development of lettuce (Lactuca sativa) in plant factories, while maintaining a constant red light photon flux and daily light integral. The results showed that compared to constant far-red light, 30 min intermittent far-red light increased yield by 11.7% and the number of leaves and buds by 2.66. Furthermore, the various metrics demonstrated that intermittent far-red light supplementation enhanced the overall effectiveness of the far-red light treatment. This was validated by analyzing phytohormone content and the expression of genes related to hormone metabolism and transport at the tip of the lettuce stems. Transcriptome analysis revealed that the differences in gene expression between treatments were primarily concentrated in genes related to signaling, hormone metabolism, and transport. Weighted Gene Co-expression Network Analysis identified the co-expression modules associated with yield and quality. Additionally, dynamic expression analysis showed genes involved to far-red photoreception, response, and hormone metabolism and transport exhibited optimal rhythmic responses only under 30 min intermittent far-red light supplementation. This suggests that intermittent far-red light irradiation at 30 min intervals is the most effective for activating far-red light signaling influencing hormone metabolism and transport, thereby accelerating the growth of lettuce leaves and buds and ultimately increasing yield.




JASMONATE ZIM-DOMAIN PROTEIN 3 regulates photo- and thermo-morphogenesis through inhibiting PIF4 in Arabidopsis

March 2024

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25 Reads

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6 Citations

Plant Physiology

Light and temperature are two major environmental factors that affect growth and development of plants during their life cycle. Plants have evolved complex mechanisms to adapt to varying external environments. Here, we show that JASMONATE ZIM-domain protein 3 (JAZ3), a jasmonic acid signaling component, acts as a factor to integrate light and temperature in regulating seedling morphogenesis. JAZ3 overexpression transgenic lines display short hypocotyls under red, far-red, and blue light and warm temperature (28 °C) conditions compared to the wild type in Arabidopsis (Arabidopsis thaliana). We show that JAZ3 interacts with the transcription factor PHYTOCHROME-INTERACTING FACTOR4 (PIF4). Interestingly, JAZ3 spontaneously undergoes liquid-liquid phase separation (LLPS) in vitro and in vivo and promotes LLPS formation of PIF4. Moreover, transcriptomic analyses indicate that JAZ3 regulates the expression of genes involved in many biological processes, such as response to auxin, auxin-activated signaling pathway, regulation of growth, and response to red light. Finally, JAZ3 inhibits the transcriptional activation activity and binding ability of PIF4. Collectively, our study reveals a function and molecular mechanism of JAZ3 in regulating plant growth in response to environmental light and temperature.


Ubiquitin-mediated stabilization of SlPsbS regulates low night temperature tolerance in tomatoes

February 2024

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17 Reads

Non-photochemical quenching (NPQ) plays a fundamental role in safely operating photosynthesis under low night temperatures (LNT). PsbS protein is essential for the rapid induction of NPQ, and its stability is often affected by adverse environmental conditions. However, the regulatory mechanism for the stability of PsbS or chloroplast proteins remains to be fully characterized. We showed that LNT decreased NPQ levels and SlPsbS protein abundance in tomato leaves. COP9 signalosome subunit 5A (SlCSN5A) facilitated SlPsbS ubiquitination and degradation in the cytosol. Further, tomato chloroplast vesiculation (SlCV) was activated by LNT. Under LNT, SlCV targeted the chloroplasts and induced the formation of CV-containing vesicles (CCVs) containing SlPsbS, which were exported from the chloroplasts. Subsequently, SlCV and SlPsbS contact SlCSN5A in the cytosol and are ubiquitinated and degraded. Genetic evidence demonstrated that overexpression of SlCV aggravated SlPsbS protein degradation, whereas silencing of SlCSN5 and SlCV delayed LNT-induced NPQ reduction and SlPsbS protein turnover. This study provides evidence that CSN5A is associated with chloroplast proteins, and reveals a ubiquitin-dependent degradation pathway of chloroplast proteins co-mediated by CV and CSN5A, thereby providing new insights into the regulation of chloroplast protein stability under stress conditions.


Citations (74)


... Chlorophyll plays a crucial role in photoelectric conversion. Li et al. (2025) comprehensively reviewed the regulation of chlorophyll biosynthesis pathways and retrograde signaling networks, highlighting chlorophyll's central role in photosynthesis and its ...

Reference:

Advances in photosynthesis research: Unlocking the potential for food security, renewable energy, and environmental sustainability
Regulatory and retrograde signaling networks in the chlorophyll biosynthetic pathway

... The assumption and methodology that the appropriate supplemental frequency of signaling light can induce time-differentiated changes in relevant signals and thus enhance production value is validated here. In our other study on far-red light, intermittent far-red light exposure amplified the effects of far-red light, thereby accelerating leaf development and increasing yield (Liu et al. 2025). However, much more detail, such as which pathways respond to this changing environment and whether all types of signaling light can bring about benefits in this way, still needs to be investigated in depth. ...

Intermittent Supplementation with Far-Red Light Accelerates Leaf and Bud Development and Increases Yield in Lettuce

... Lhc subfamily primarily functions in light harvesting [63], while PsbS subfamily plays a crucial role in the initiation non-photochemical quenching (NPQ) and the protection of PSII [66]. Recent report has shown that SlPsbS regulated the stability of chloroplast proteins under stress conditions and a decrease in SlPsbS protein abundance under low night temperature stress led to low NPQ values, which exacerbated PS photodamage and reduced plant stress tolerance [67]. ...

Ubiquitin-mediated degradation of SlPsbS regulates low night temperature tolerance in tomatoes

Cell Reports

... In a recent controversy, van Wijk proposed that the vesiclemediated pathways may be involved in the ubiquitinated degradation of chloroplast proteins, 41 and the CELL DIVISION CY-CLE48 (CDC48) dependence of the processing of ubiquitinated chloroplast proteins does point to the existence of an export system. 42 Similarly, our results revealed that LNT stress induced high expression of SlCV and promoted SlCV translocation from the chloroplasts ( Figure S5). In Arabidopsis, CV was found to destabilize the chloroplasts and induce the formation of CCVs. ...

Reply: Does the polyubiquitination pathway operate inside intact chloroplasts to remove proteins?
  • Citing Article
  • May 2024

The Plant Cell

... LUC results showed that these three TFs all have positive regulation of BcJAZ2 expression (Fig. 9e). Interestingly, these three TFs all belong to the bHLH family, which regulate cell proliferation and elongation, and their transgenic Arabidopsis exhibits an early flowering phenotype [41,42] . Recent studies showed that JAZ3 transgenic Arabidopsis displayed shorter hypocotyls and larger cotyledons under specific light or temperature environments, compared with the wild-type Arabidopsis [43] . ...

JASMONATE ZIM-DOMAIN PROTEIN 3 regulates photo- and thermo-morphogenesis through inhibiting PIF4 in Arabidopsis
  • Citing Article
  • March 2024

Plant Physiology

... For dark-activated TFs such as PIF3, FHY3, ARF6, and BZR1, RNA-Rx also detected more light-induced genes and less light-repressed genes than standard RNA-seq. In the case of PIF3 target genes, RNA-Rx detected 240 instead of 83 upregulated genes and 145 instead of 265 downregulated genes after 24 hours of light ( Fig. 4A-B), in agreement with PIF3 acting both as a transcriptional activator and repressor in etiolated cotyledons [39][40][41] . ...

The PIF1/PIF3‐MED25‐HDA19 transcriptional repression complex regulates phytochrome signaling in Arabidopsis

... The concept of green innovation, otherwise known as sustainable innovation or ecoinnovation within the manufacturing industry has been operationalized as strategies and/ or practices including adoption of advanced technologies, energy efficient production practices, and environmental sensitive manufacturing policy interventions (e.g., waste recycling) that enterprises integrate in their production lines or processes (Guan, 2017). Simply put, the systematic development and sequential application of new ideas, practices, services, or production policies that significantly improve ecological health (Huai & Lin, 2023;Sun & Chen, 2023;Uyarra et al., 2016). ...

Green innovation for green light
  • Citing Article
  • April 2023

Science China. Life sciences

... The two genes with the highest Pi values in the coding region in the C. fruticosum chloroplast genome were accD and clpP. ClpP encodes a protease that is involved in chloroplast protein homeostasis and gene expression regulation [52]. AccD encodes the β-carboxyltransferase subunit of acetyl-CoA carboxylase [53]. ...

Regulation of chloroplast protein degradation

Journal of Genetics and Genomics

... Generally, transcription factors are located in the nucleus, where they regulate the expression of target genes. PIF3 is also located in nucleus in A. annua and A. thaliana 13,23 . According to previously reported methods, we analyzed the localization of UrPIF3 using heterologous expression in tobacco 13 . ...

Arabidopsis EXECUTER1 interacts with WRKY transcription factors to mediate plastid-to-nucleus singlet oxygen signaling
  • Citing Article
  • November 2022

The Plant Cell

... Biomolecular condensates have been recognized as an important organizer for compartmentalization of cellular components 17 . Many condensates, which exhibit liquid-like properties, are assembled through liquid-liquid phase separation (LLPS) 18 and are responsive to environmental stimuli [19][20][21][22] . However, the role of cold-induced biomolecular condensates in plants remains unclear. ...

Condensation of SEUSS promotes hyperosmotic stress tolerance in Arabidopsis

Nature Chemical Biology