Fang Lin
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Research Subjects
Research Interests
Fang Lin has a total of 47 co-authors. Here are the top co-authors by number of shared publications:
- Hua Zhou
Laboratory of Plant and Environment Interaction Study, Tsientang Institute for Advanced Study , Hangzhou 310000
4 shared publications - Dongqing Xu
State Key Laboratory of Crop Genetics & Germplasm Enhancement and Utilization, National Center for Soybean Improvement, College of Agriculture, Nanjing Agricultural University , Nanjing 210095
3 shared publications Jiale Yuan
Ministry of Education Key Laboratory of Cell Activities and Stress Adaptations, School of Life Sciences, Lanzhou University
3 shared publications- Jing Cao
Ministry of Education Key Laboratory of Cell Activities and Stress Adaptations, School of Life Sciences, Lanzhou University , Lanzhou 730000
3 shared publications Beihong Zhang
Ministry of Education Key Laboratory of Cell Activities and Stress Adaptations, School of Life Sciences, Lanzhou University, Lanzhou, China
2 shared publications- Jia Li
Guangdong Provincial Key Laboratory of Plant Adaptation and Molecular Design, School of Life Sciences, Guangzhou University, Guangzhou, China
2 shared publications
Latest Publications
- Open Access
- 21 September 2026
- Published by Oxford University Press (OUP) in The Plant Cell
- Vol. 38 (10)
BR-ENHANCED EXPRESSION 1/2/3 fine-tune plant photomorphogenesis by regulating the phyA/B-SPA1 cascade
BR-ENHANCED EXPRESSION1 (BEE1), BEE2, and BEE3, key transcriptional components of brassinosteroid signaling, have been shown to play critical roles in regulating light signaling. However, the precise molecular mechanism by which BEE1/2/3 repress photomorphogenesis remains elusive. In this study, we found that light triggers the degradation of BEE1/2/3 proteins through the 26S proteasome system. BEE1/2/3 interact with PHYTOCHROME A (phyA) and phyB, and this interaction promotes the degradation of BEE3. Mutations in BEE1/2/3 exhibit short hypocotyls and partially suppress the long hypocotyl phenotype of the phyA phyB double mutant. BEE1/2/3 attenuate the interaction between phyB and PHYTOCHROME-INTERACTING FACTOR 3 (PIF3), leading to PIF3 stabilization and subsequent repression of photomorphogenesis. In addition, BEE1/2/3 interact with SUPPRESSORs OF PHYTOCHROME A (SPAs) to disrupt the phyA/B-SPA1 association, thereby further attenuating the exaggerated light signaling. The COP1–SPAs complexes stabilize BEE1/3 under both dark and light conditions. Mutations in BEE1/2/3 enhance the constitutively photomorphogenic phenotype of spa1/2/4, whereas overexpression of BEE1/3 partially rescue the short hypocotyls of cop1 and spa1/2/4. Collectively, these findings demonstrate that BEE1/2/3 fine-tune the phyA/B-COP1/SPA1 signaling hub to negatively regulate photomorphogenesis in response to excessive light signaling. Our study provides insights into the pivotal roles of BEE1/2/3 as photomorphogenic repressors, facilitating adaptation to complex light conditions.
- 21 September 2026
- Published by Oxford University Press (OUP) in The Plant Cell
- Vol. 38 (10)
Open AccessBR-ENHANCED EXPRESSION 1/2/3 fine-tune plant photomorphogenesis by regulating the phyA/B-SPA1 cascade
BR-ENHANCED EXPRESSION1 (BEE1), BEE2, and BEE3, key transcriptional components of brassinosteroid signaling, have been shown to play critical roles in regulating light signaling. However, the precise molecular mechanism by which BEE1/2/3 repress photomorphogenesis remains elusive. In this study, we found that light triggers the degradation of BEE1/2/3 proteins through the 26S proteasome system. BEE1/2/3 interact with PHYTOCHROME A (phyA) and phyB, and this interaction promotes the degradation of BEE3. Mutations in BEE1/2/3 exhibit short hypocotyls and partially suppress the long hypocotyl phenotype of the phyA phyB double mutant. BEE1/2/3 attenuate the interaction between phyB and PHYTOCHROME-INTERACTING FACTOR 3 (PIF3), leading to PIF3 stabilization and subsequent repression of photomorphogenesis. In addition, BEE1/2/3 interact with SUPPRESSORs OF PHYTOCHROME A (SPAs) to disrupt the phyA/B-SPA1 association, thereby further attenuating the exaggerated light signaling. The COP1–SPAs complexes stabilize BEE1/3 under both dark and light conditions. Mutations in BEE1/2/3 enhance the constitutively photomorphogenic phenotype of spa1/2/4, whereas overexpression of BEE1/3 partially rescue the short hypocotyls of cop1 and spa1/2/4. Collectively, these findings demonstrate that BEE1/2/3 fine-tune the phyA/B-COP1/SPA1 signaling hub to negatively regulate photomorphogenesis in response to excessive light signaling. Our study provides insights into the pivotal roles of BEE1/2/3 as photomorphogenic repressors, facilitating adaptation to complex light conditions.
- Open Access
- 15 September 2026
- Published by Frontiers Media SA in Frontiers in Plant Science
- Vol. 17, 1930423
Beyond light signaling: the broad regulatory spectrum of AFPs in plant growth and development
AFPs (ABI5-BINDING PROTEINS) are a unique class of small, plant-specific proteins, originally identified through their interaction with ABI5 (ABSCISIC ACID INSENSITIVE 5), a pivotal transcription factor in ABA (ABSCISIC ACID) signaling pathway. Emerging evidence indicates that AFPs act as multifunctional integrators governing not only ABA signaling but also a broad spectrum of plant growth, development, biotic and abiotic stress adaptation. This review systematically summarizes the domain architecture, interactome networks, and evolutionary conservation of AFPs, and further highlights the pleiotropic functions of AFPs in seed dormancy and germination, photoperiodic flowering, plant architecture modulation, abiotic stress tolerance and disease resistance. Through diverse molecular mechanisms, including recruitment of corepressors, control of target protein stability, and modulation of chromatin states, AFPs operate as microprotein modulators at critical nodes of phytohormone networks, offering novel insights into the regulatory principles balancing plant growth and stress resilience.
- 15 September 2026
- Published by Frontiers Media SA in Frontiers in Plant Science
- Vol. 17, 1930423
Open AccessBeyond light signaling: the broad regulatory spectrum of AFPs in plant growth and development
AFPs (ABI5-BINDING PROTEINS) are a unique class of small, plant-specific proteins, originally identified through their interaction with ABI5 (ABSCISIC ACID INSENSITIVE 5), a pivotal transcription factor in ABA (ABSCISIC ACID) signaling pathway. Emerging evidence indicates that AFPs act as multifunctional integrators governing not only ABA signaling but also a broad spectrum of plant growth, development, biotic and abiotic stress adaptation. This review systematically summarizes the domain architecture, interactome networks, and evolutionary conservation of AFPs, and further highlights the pleiotropic functions of AFPs in seed dormancy and germination, photoperiodic flowering, plant architecture modulation, abiotic stress tolerance and disease resistance. Through diverse molecular mechanisms, including recruitment of corepressors, control of target protein stability, and modulation of chromatin states, AFPs operate as microprotein modulators at critical nodes of phytohormone networks, offering novel insights into the regulatory principles balancing plant growth and stress resilience.
- Open Access
- 16 July 2026
- Published by Wiley in Journal of Integrative Plant Biology
- Vol. 68 (8) , 3107-3132
14‐3‐3 proteins: Central hubs in plant development and stress adaptation
14‐3‐3 proteins are evolutionarily conserved regulators that orchestrate diverse physiological processes by binding to phosphorylated client proteins. These interactions modulate client activity, localization, stability, or interactome composition, thereby coordinating plant growth, development, metabolism, and stress responses. Mounting evidence underscores their direct involvement in phytohormone signaling and stress‐response pathways, in which 14‐3‐3 binding often constitutes a key regulatory event. This review highlights recent advances elucidating the pivotal role of 14‐3‐3 proteins in regulating phytohormone networks, biotic and abiotic stress tolerance, and evolutionary adaptations. We position 14‐3‐3 proteins as crucial integrators of internal signals and external cues, serving as central hubs that rewire cellular networks to enable adaptation to complex, changing environments.
- 16 July 2026
- Published by Wiley in Journal of Integrative Plant Biology
- Vol. 68 (8) , 3107-3132
Open Access14‐3‐3 proteins: Central hubs in plant development and stress adaptation
14‐3‐3 proteins are evolutionarily conserved regulators that orchestrate diverse physiological processes by binding to phosphorylated client proteins. These interactions modulate client activity, localization, stability, or interactome composition, thereby coordinating plant growth, development, metabolism, and stress responses. Mounting evidence underscores their direct involvement in phytohormone signaling and stress‐response pathways, in which 14‐3‐3 binding often constitutes a key regulatory event. This review highlights recent advances elucidating the pivotal role of 14‐3‐3 proteins in regulating phytohormone networks, biotic and abiotic stress tolerance, and evolutionary adaptations. We position 14‐3‐3 proteins as crucial integrators of internal signals and external cues, serving as central hubs that rewire cellular networks to enable adaptation to complex, changing environments.
- 25 February 2026
- Published by Elsevier in Journal of Plant Physiology
- Vol. 319, 154738
Phytochrome-interacting factors (PIFs): Integrating phytohormone signals at the nexus of development and stress adaptation
- 25 February 2026
- Published by Elsevier in Journal of Plant Physiology
- Vol. 319, 154738
Phytochrome-interacting factors (PIFs): Integrating phytohormone signals at the nexus of development and stress adaptation
- Open Access
- 15 June 2024
- Published by Springer Nature in Nature Communications
- Vol. 15 (1) , 1-14
Light regulates nuclear detainment of intron-retained transcripts through COP1-spliceosome to modulate photomorphogenesis
Intron retention (IR) is the most common alternative splicing event in Arabidopsis. An increasing number of studies have demonstrated the major role of IR in gene expression regulation. The impacts of IR on plant growth and development and response to environments remain underexplored. Here, we found that IR functions directly in gene expression regulation on a genome-wide scale through the detainment of intron-retained transcripts (IRTs) in the nucleus. Nuclear-retained IRTs can be kept away from translation through this mechanism. COP1-dependent light modulation of the IRTs of light signaling genes, such as PIF4, RVE1, and ABA3, contribute to seedling morphological development in response to changing light conditions. Furthermore, light-induced IR changes are under the control of the spliceosome, and in part through COP1-dependent ubiquitination and degradation of DCS1, a plant-specific spliceosomal component. Our data suggest that light regulates the activity of the spliceosome and the consequent IRT nucleus detainment to modulate photomorphogenesis through COP1.
- 15 June 2024
- Published by Springer Nature in Nature Communications
- Vol. 15 (1) , 1-14
Open AccessLight regulates nuclear detainment of intron-retained transcripts through COP1-spliceosome to modulate photomorphogenesis
Intron retention (IR) is the most common alternative splicing event in Arabidopsis. An increasing number of studies have demonstrated the major role of IR in gene expression regulation. The impacts of IR on plant growth and development and response to environments remain underexplored. Here, we found that IR functions directly in gene expression regulation on a genome-wide scale through the detainment of intron-retained transcripts (IRTs) in the nucleus. Nuclear-retained IRTs can be kept away from translation through this mechanism. COP1-dependent light modulation of the IRTs of light signaling genes, such as PIF4, RVE1, and ABA3, contribute to seedling morphological development in response to changing light conditions. Furthermore, light-induced IR changes are under the control of the spliceosome, and in part through COP1-dependent ubiquitination and degradation of DCS1, a plant-specific spliceosomal component. Our data suggest that light regulates the activity of the spliceosome and the consequent IRT nucleus detainment to modulate photomorphogenesis through COP1.
- Open Access
- 6 January 2023
- Published by Springer Nature in Stress Biology
- Vol. 3 (1) , 1-19
Multi-layered roles of BBX proteins in plant growth and development
Light and phytohormone are external and internal cues that regulate plant growth and development throughout their life cycle. BBXs (B-box domain proteins) are a group of zinc finger proteins that not only directly govern the transcription of target genes but also associate with other factors to create a meticulous regulatory network to precisely regulate numerous aspects of growth and developmental processes in plants. Recent studies demonstrate that BBXs play pivotal roles in light-controlled plant growth and development. Besides, BBXs have been documented to regulate phytohormone-mediated physiological procedures. In this review, we summarize and highlight the multi-faced role of BBXs, with a focus in photomorphogenesis, photoperiodic flowering, shade avoidance, abiotic stress, and phytohormone-mediated growth and development in plant.
- 6 January 2023
- Published by Springer Nature in Stress Biology
- Vol. 3 (1) , 1-19
Open AccessMulti-layered roles of BBX proteins in plant growth and development
Light and phytohormone are external and internal cues that regulate plant growth and development throughout their life cycle. BBXs (B-box domain proteins) are a group of zinc finger proteins that not only directly govern the transcription of target genes but also associate with other factors to create a meticulous regulatory network to precisely regulate numerous aspects of growth and developmental processes in plants. Recent studies demonstrate that BBXs play pivotal roles in light-controlled plant growth and development. Besides, BBXs have been documented to regulate phytohormone-mediated physiological procedures. In this review, we summarize and highlight the multi-faced role of BBXs, with a focus in photomorphogenesis, photoperiodic flowering, shade avoidance, abiotic stress, and phytohormone-mediated growth and development in plant.
- Open Access
- 16 March 2022
- Published by Oxford University Press (OUP) in The Plant Cell
- Vol. 34 (6) , 2266-2285
The photomorphogenic repressors BBX28 and BBX29 integrate light and brassinosteroid signaling to inhibit seedling development in Arabidopsis
B-box containing proteins (BBXs) integrate light and various hormonal signals to regulate plant growth and development. Here, we demonstrate that the photomorphogenic repressors BBX28 and BBX29 positively regulate brassinosteroid (BR) signaling in Arabidopsis thaliana seedlings. Treatment with the BR brassinolide stabilized BBX28 and BBX29, which partially depended on BR INSENSITIVE1 (BRI1) and BIN2. bbx28 bbx29 seedlings exhibited larger cotyledon aperture than the wild-type when treated with brassinazole in the dark, which partially suppressed the closed cotyledons of brassinazole resistant 1-1D (bzr1-1D). Consistently, overexpressing BBX28 and BBX29 partially rescued the short hypocotyls of bri1-5 and bin2-1 in both the dark and light, while the loss-of-function of BBX28 and BBX29 partially suppressed the long hypocotyls of bzr1-1D in the light. BBX28 and BBX29 physically interacted with BR-ENHANCED EXPRESSION1 (BEE1), BEE2, and BEE3 and enhanced their binding to and activation of their target genes. Moreover, BBX28 and BBX29 as well as BEE1, BEE2, and BEE3 increased BZR1 accumulation to promote the BR signaling pathway. Therefore, both BBX28 and BBX29 interact with BEE1, BEE2, and BEE3 to orchestrate light and BR signaling by facilitating the transcriptional activity of BEE target genes. Our study provides insights into the pivotal roles of BBX28 and BBX29 as signal integrators in ensuring normal seedling development.
- 16 March 2022
- Published by Oxford University Press (OUP) in The Plant Cell
- Vol. 34 (6) , 2266-2285
Open AccessThe photomorphogenic repressors BBX28 and BBX29 integrate light and brassinosteroid signaling to inhibit seedling development in Arabidopsis
B-box containing proteins (BBXs) integrate light and various hormonal signals to regulate plant growth and development. Here, we demonstrate that the photomorphogenic repressors BBX28 and BBX29 positively regulate brassinosteroid (BR) signaling in Arabidopsis thaliana seedlings. Treatment with the BR brassinolide stabilized BBX28 and BBX29, which partially depended on BR INSENSITIVE1 (BRI1) and BIN2. bbx28 bbx29 seedlings exhibited larger cotyledon aperture than the wild-type when treated with brassinazole in the dark, which partially suppressed the closed cotyledons of brassinazole resistant 1-1D (bzr1-1D). Consistently, overexpressing BBX28 and BBX29 partially rescued the short hypocotyls of bri1-5 and bin2-1 in both the dark and light, while the loss-of-function of BBX28 and BBX29 partially suppressed the long hypocotyls of bzr1-1D in the light. BBX28 and BBX29 physically interacted with BR-ENHANCED EXPRESSION1 (BEE1), BEE2, and BEE3 and enhanced their binding to and activation of their target genes. Moreover, BBX28 and BBX29 as well as BEE1, BEE2, and BEE3 increased BZR1 accumulation to promote the BR signaling pathway. Therefore, both BBX28 and BBX29 interact with BEE1, BEE2, and BEE3 to orchestrate light and BR signaling by facilitating the transcriptional activity of BEE target genes. Our study provides insights into the pivotal roles of BBX28 and BBX29 as signal integrators in ensuring normal seedling development.
- Open Access
- 30 November 2021
- Published by MDPI AG in International Journal of Molecular Sciences
- Vol. 22 (23) , 12971
Integration of Light and Brassinosteroid Signaling during Seedling Establishment
Light and brassinosteroid (BR) are external stimuli and internal cue respectively, that both play critical roles in a wide range of developmental and physiological process. Seedlings grown in the light exhibit photomorphogenesis, while BR promotes seedling etiolation. Light and BR oppositely control the development switch from skotomorphogenesis in the dark to photomorphogenesis in the light. Recent progress report that substantial components have been identified as hubs to integrate light and BR signals. Photomorphogenic repressors including COP1, PIFs, and AGB1 have been reported to elevate BR response, while photomorphogenesis-promoting factors such as HY5, BZS1, and NF-YCs have been proven to repress BR signal. In addition, BR components also modulate light signal. Here, we review the current research on signaling network associated with light and brassinosteroids, with a focus on the integration of light and BR signals enabling plants to thrive in the changeable environment.
- 30 November 2021
- Published by MDPI AG in International Journal of Molecular Sciences
- Vol. 22 (23) , 12971
Open AccessIntegration of Light and Brassinosteroid Signaling during Seedling Establishment
Light and brassinosteroid (BR) are external stimuli and internal cue respectively, that both play critical roles in a wide range of developmental and physiological process. Seedlings grown in the light exhibit photomorphogenesis, while BR promotes seedling etiolation. Light and BR oppositely control the development switch from skotomorphogenesis in the dark to photomorphogenesis in the light. Recent progress report that substantial components have been identified as hubs to integrate light and BR signals. Photomorphogenic repressors including COP1, PIFs, and AGB1 have been reported to elevate BR response, while photomorphogenesis-promoting factors such as HY5, BZS1, and NF-YCs have been proven to repress BR signal. In addition, BR components also modulate light signal. Here, we review the current research on signaling network associated with light and brassinosteroids, with a focus on the integration of light and BR signals enabling plants to thrive in the changeable environment.