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1

Sato, Y., M. Wada, and A. Kadota. "Choice of tracks, microtubules and/or actin filaments for chloroplast photo-movement is differentially controlled by phytochrome and a blue light receptor." Journal of Cell Science 114, no. 2 (January 15, 2001): 269–79. http://dx.doi.org/10.1242/jcs.114.2.269.

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Light induced chloroplast movement has been studied as a model system for photoreception and actin microfilament (MF)-based intracellular motilities in plants. Chloroplast photo-accumulation and -avoidance movement is mediated by phytochrome as well as blue light (BL) receptor in the moss Physcomitrella patens. Here we report the discovery of an involvement of a microtubule (MT)-based system in addition to an MF-based system in photorelocation of chloroplasts in this moss. In the dark, MTs provided tracks for rapid movement of chloroplasts in a longitudinal direction and MFs contributed the tracks for slow movement in any direction. We found that phytochrome responses utilized only the MT-based system, while BL responses had an alternative way of moving, either along MTs or MFs. MT-based systems were mediated by both photoreceptors, but chloroplasts showed movements with different velocity and pattern between them. No apparent difference in the behavior of chloroplast movement between the accumulation and avoidance movement was detected in phytochrome responses or BL responses, except for the direction of the movement. The results presented here demonstrate that chloroplasts use both MTs and MFs for motility and that phytochrome and a BL receptor control directional photo-movement of chloroplasts through the differential regulation of these motile systems.
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2

Yuan, Ning, Lavanya Mendu, Kaushik Ghose, Carlie Shea Witte, Julia Frugoli, and Venugopal Mendu. "FKF1 Interacts with CHUP1 and Regulates Chloroplast Movement in Arabidopsis." Plants 12, no. 3 (January 25, 2023): 542. http://dx.doi.org/10.3390/plants12030542.

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Plants have mechanisms to relocate chloroplasts based on light intensities in order to maximize photosynthesis and reduce photodamage. Under low light, chloroplasts move to the periclinal walls to increase photosynthesis (accumulation) and move to the anticlinal walls under high light to avoid photodamage, and even cell death (avoidance). Arabidopsis blue light receptors phot1 and phot2 (phototropins) have been reported to regulate chloroplast movement. This study discovered that another blue light receptor, FLAVIN-BINDING KELCH REPEAT F-BOX1 (FKF1), regulates chloroplast photorelocation by physically interacting with chloroplast unusual positioning protein 1 (CHUP1), a critical component of the chloroplast motility system. Leaf cross-sectioning and red-light transmittance results showed that overexpression of FKF1 compromised the avoidance response, while the absence of FKF1 enhanced chloroplast movements under high light. Western blot analysis showed that CHUP1 protein abundance is altered in FKF1 mutants and overexpression lines, indicating a potential regulation of CHUP1 by FKF1. qPCR results showed that two photorelocation pathway genes, JAC1 and THRUMIN1, were upregulated in FKF1-OE lines, and overexpression of FKF1 in the THRUMIN1 mutant weakened its accumulation and avoidance responses, indicating that JAC1 and THRUMIN1 may play a role in the FKF1-mediated chloroplast avoidance response. However, the precise functional roles of JAC1 and THRUMIN1 in this process are not known.
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3

Lin, Yi-Jyun, Yu-Chung Chen, Kuan-Chieh Tseng, Wen-Chi Chang, and Swee-Suak Ko. "Phototropins Mediate Chloroplast Movement in Phalaenopsis aphrodite (Moth Orchid)." Plant and Cell Physiology 60, no. 10 (June 14, 2019): 2243–54. http://dx.doi.org/10.1093/pcp/pcz116.

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AbstractChloroplast movement is important for plants to avoid photodamage and to perform efficient photosynthesis. Phototropins are blue light receptors in plants that function in chloroplast movement, phototropism, stomatal opening, and they also affect plant growth and development. In this study, full-length cDNAs of two PHOTOTROPIN genes, PaPHOT1 and PaPHOT2, were cloned from a moth orchid Phalaenopsis aphrodite, and their functions in chloroplast movement were investigated. Phylogenetic analysis showed that PaPHOT1 and PaPHOT2 orthologs were highly similar to PHOT1 and PHOT2 of the close relative Phalaenopsis equestris, respectively, and clustered with monocots PHOT1 and PHOT2 orthologs, respectively. Phalaenopsis aphrodite expressed a moderate level of PaPHOT1 under low blue light of 5 μmol�m−2�s−1 (BL5) and a high levels of PaPHOT1 at >BL100. However, PaPHOT2 was expressed at low levels at <BL50 but expressed at high levels at > BL100. Analysis of light-induced chloroplast movements using the SPAD method indicated that orchid accumulated chloroplasts at <BL10. The chloroplast avoidance response was detectable at >BL25 and significant chloroplast avoidance movement was observed at >BL100. Virus-induced gene silencing of PaPHOTs in orchids showed decreased gene expression of PaPHOTs and reduced both chloroplast accumulation and avoidance responses. Heterologous expression of PaPHOT1 in Arabidopsis phot1phot2 double mutant recovered chloroplast accumulation response at BL5, but neither PaPHOT1 nor PaPHOT2 was able to restore mutant chloroplast avoidance at BL100. Overall, this study showed that phototropins mediate chloroplast movement in Phalaenopsis orchid is blue light-dependent but their function is slightly different from Arabidopsis which might be due to gene evolution.
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4

Suetsugu, Noriyuki, Atsushi Takemiya, Sam-Geun Kong, Takeshi Higa, Aino Komatsu, Ken-ichiro Shimazaki, Takayuki Kohchi, and Masamitsu Wada. "RPT2/NCH1 subfamily of NPH3-like proteins is essential for the chloroplast accumulation response in land plants." Proceedings of the National Academy of Sciences 113, no. 37 (August 30, 2016): 10424–29. http://dx.doi.org/10.1073/pnas.1602151113.

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In green plants, the blue light receptor kinase phototropin mediates various photomovements and developmental responses, such as phototropism, chloroplast photorelocation movements (accumulation and avoidance), stomatal opening, and leaf flattening, which facilitate photosynthesis. In Arabidopsis, two phototropins (phot1 and phot2) redundantly mediate these responses. Two phototropin-interacting proteins, NONPHOTOTROPIC HYPOCOTYL 3 (NPH3) and ROOT PHOTOTROPISM 2 (RPT2), which belong to the NPH3/RPT2-like (NRL) family of BTB (broad complex, tramtrack, and bric à brac) domain proteins, mediate phototropism and leaf flattening. However, the roles of NRL proteins in chloroplast photorelocation movement remain to be determined. Here, we show that another phototropin-interacting NRL protein, NRL PROTEIN FOR CHLOROPLAST MOVEMENT 1 (NCH1), and RPT2 redundantly mediate the chloroplast accumulation response but not the avoidance response. NPH3, RPT2, and NCH1 are not involved in the chloroplast avoidance response or stomatal opening. In the liverwort Marchantia polymorpha, the NCH1 ortholog, MpNCH1, is essential for the chloroplast accumulation response but not the avoidance response, indicating that the regulation of the phototropin-mediated chloroplast accumulation response by RPT2/NCH1 is conserved in land plants. Thus, the NRL protein combination could determine the specificity of diverse phototropin-mediated responses.
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5

Kasahara, Masahiro, Takatoshi Kagawa, Kazusato Oikawa, Noriyuki Suetsugu, Mitsue Miyao, and Masamitsu Wada. "Chloroplast avoidance movement reduces photodamage in plants." Nature 420, no. 6917 (December 2002): 829–32. http://dx.doi.org/10.1038/nature01213.

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6

Kagawa, Takatoshi, and Masamitsu Wada. "Velocity of chloroplast avoidance movement is fluence rate dependent." Photochemical & Photobiological Sciences 3, no. 6 (2004): 592. http://dx.doi.org/10.1039/b316285k.

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7

Majumdar, Arkajo, and Rup Kumar Kar. "Chloroplast avoidance movement: a novel paradigm of ROS signalling." Photosynthesis Research 144, no. 1 (March 28, 2020): 109–21. http://dx.doi.org/10.1007/s11120-020-00736-9.

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8

Ko, Swee-Suak, Chung-Min Jhong, Yi-Jyun Lin, Ching-Yu Wei, Ju-Yin Lee, and Ming-Che Shih. "Blue Light Mediates Chloroplast Avoidance and Enhances Photoprotection of Vanilla Orchid." International Journal of Molecular Sciences 21, no. 21 (October 28, 2020): 8022. http://dx.doi.org/10.3390/ijms21218022.

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Vanilla orchid, which is well-known for its flavor and fragrance, is cultivated in tropical and subtropical regions. This shade-loving plant is very sensitive to high irradiance. In this study, we show that vanilla chloroplasts started to have avoidance movement when blue light (BL) was higher than 20 μmol m−2s−1 and significant avoidance movement was observed under BL irradiation at 100 μmol m−2s−1 (BL100). The light response curve indicated that when vanilla was exposed to 1000 μmol m−2s−1, the electron transport rate (ETR) and photochemical quenching of fluorescence (qP) were significantly reduced to a negligible amount. We found that if a vanilla orchid was irradiated with BL100 for 12 days, it acquired BL-acclimation. Chloroplasts moved to the side of cells in order to reduce light-harvesting antenna size, and chloroplast photodamage was eliminated. Therefore, BL-acclimation enhanced vanilla orchid growth and tolerance to moderate (500 μmol m−2s−1) and high light (1000 μmol m−2s−1) stress conditions. It was found that under high irradiation, BL-acclimatized vanilla maintained higher ETR and qP capacity than the control without BL-acclimation. BL-acclimation induced antioxidant enzyme activities, reduced ROS accumulation, and accumulated more carbohydrates. Moreover, BL-acclimatized orchids upregulated photosystem-II-associated marker genes (D1 and PetC), Rubisco and PEPC transcripts and sustained expression levels thereof, and also maximized the photosynthesis rate. Consequently, BL-acclimatized orchids had higher biomass. In short, this study found that acclimating vanilla orchid with BL before transplantation to the field might eliminate photoinhibition and enhance vanilla growth and production.
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9

Kitashova, Anastasia, Katja Schneider, Lisa Fürtauer, Laura Schröder, Tim Scheibenbogen, Siegfried Fürtauer, and Thomas Nägele. "Impaired chloroplast positioning affects photosynthetic capacity and regulation of the central carbohydrate metabolism during cold acclimation." Photosynthesis Research 147, no. 1 (November 19, 2020): 49–60. http://dx.doi.org/10.1007/s11120-020-00795-y.

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AbstractPhotosynthesis and carbohydrate metabolism of higher plants need to be tightly regulated to prevent tissue damage during environmental changes. The intracellular position of chloroplasts changes due to a changing light regime. Chloroplast avoidance and accumulation response under high and low light, respectively, are well known phenomena, and deficiency of chloroplast movement has been shown to result in photodamage and reduced biomass accumulation. Yet, effects of chloroplast positioning on underlying metabolic regulation are less well understood. Here, we analysed photosynthesis together with metabolites and enzyme activities of the central carbohydrate metabolism during cold acclimation of the chloroplast unusual positioning 1 (chup1) mutant of Arabidopsis thaliana. We compared cold acclimation under ambient and low light and found that maximum quantum yield of PSII was significantly lower in chup1 than in Col-0 under both conditions. Our findings indicated that net CO2 assimilation in chup1 is rather limited by biochemistry than by photochemistry. Further, cold-induced dynamics of sucrose phosphate synthase differed significantly between both genotypes. Together with a reduced rate of sucrose cycling derived from kinetic model simulations our study provides evidence for a central role of chloroplast positioning for photosynthetic and metabolic acclimation to low temperature.
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10

Sztatelman, Olga, Andrzej Waloszek, Agnieszka Katarzyna Banaś, and Halina Gabryś. "Photoprotective function of chloroplast avoidance movement: In vivo chlorophyll fluorescence study." Journal of Plant Physiology 167, no. 9 (June 2010): 709–16. http://dx.doi.org/10.1016/j.jplph.2009.12.015.

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11

Higa, Takeshi, and Masamitsu Wada. "Chloroplast avoidance movement is not functional in plants grown under strong sunlight." Plant, Cell & Environment 39, no. 4 (February 5, 2016): 871–82. http://dx.doi.org/10.1111/pce.12681.

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12

Uenaka, Hidetoshi, and Akeo Kadota. "Functional analyses of the Physcomitrella patens phytochromes in regulating chloroplast avoidance movement." Plant Journal 51, no. 6 (August 7, 2007): 1050–61. http://dx.doi.org/10.1111/j.1365-313x.2007.03202.x.

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13

Ichikawa, Satoshi, Noboru Yamada, Noriyuki Suetsugu, Masamitsu Wada, and Akeo Kadota. "Red Light, Phot1 and JAC1 Modulate Phot2-Dependent Reorganization of Chloroplast Actin Filaments and Chloroplast Avoidance Movement." Plant and Cell Physiology 52, no. 8 (July 7, 2011): 1422–32. http://dx.doi.org/10.1093/pcp/pcr087.

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14

Schmalstig, Judy G., and Kenneth Jainandan. "Green light attenuates blue‐light‐induced chloroplast avoidance movement in Arabidopsis and Landoltia punctata." American Journal of Botany 108, no. 8 (August 2021): 1525–39. http://dx.doi.org/10.1002/ajb2.1717.

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15

Usami, Hiroka, Takuma Maeda, Yusuke Fujii, Kazusato Oikawa, Fumio Takahashi, Takatoshi Kagawa, Masamitsu Wada, and Masahiro Kasahara. "CHUP1 mediates actin-based light-induced chloroplast avoidance movement in the moss Physcomitrella patens." Planta 236, no. 6 (August 30, 2012): 1889–97. http://dx.doi.org/10.1007/s00425-012-1735-6.

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16

Riu, Young-Sun, Hyun-Geun Song, Hwi-Su Kim, and Sam-Geun Kong. "Guard-Cell-Specific Expression of Phototropin2 C-Terminal Fragment Enhances Leaf Transpiration." Plants 11, no. 1 (December 26, 2021): 65. http://dx.doi.org/10.3390/plants11010065.

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Phototropins (phot1 and phot2) are plant-specific blue light receptors that mediate chloroplast movement, stomatal opening, and phototropism. Phototropin is composed of the N-terminus LOV1 and LOV2 domains and the C-terminus Ser/Thr kinase domain. In previous studies, 35-P2CG transgenic plants expressing the phot2 C-terminal fragment–GFP fusion protein (P2CG) under the control of 35S promoter showed constitutive phot2 responses, including chloroplast avoidance response, stomatal opening, and reduced hypocotyl phototropism regardless of blue light, and some detrimental growth phenotypes. In this study, to exclude the detrimental growth phenotypes caused by the ectopic expression of P2C and to improve leaf transpiration, we used the PHOT2 promoter for the endogenous expression of GFP-fused P2C (GP2C) (P2-GP2C) and the BLUS1 promoter for the guard-cell-specific expression of GP2C (B1-GP2C), respectively. In P2-GP2C plants, GP2C expression induced constitutive phototropin responses and a relatively dwarf phenotype as in 35-P2CG plants. In contrast, B1-GP2C plants showed the guard-cell-specific P2C expression that induced constitutive stomatal opening with normal phototropism, chloroplast movement, and growth phenotype. Interestingly, leaf transpiration was significantly improved in B1-GP2C plants compared to that in P2-GP2C plants and WT. Taken together, this transgenic approach could be applied to improve leaf transpiration in indoor plants.
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17

Kagawa, T. "Function Analysis of Phototropin2 using Fern Mutants Deficient in Blue Light-Induced Chloroplast Avoidance Movement." Plant and Cell Physiology 45, no. 4 (April 15, 2004): 416–26. http://dx.doi.org/10.1093/pcp/pch045.

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18

Nauš, Jan, Slavomír Šmecko, and Martina Špundová. "Chloroplast avoidance movement as a sensitive indicator of relative water content during leaf desiccation in the dark." Photosynthesis Research 129, no. 2 (July 2, 2016): 217–25. http://dx.doi.org/10.1007/s11120-016-0291-5.

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19

Majumdar, Arkajo, and Rup Kumar Kar. "Integrated role of ROS and Ca+2 in blue light-induced chloroplast avoidance movement in leaves of Hydrilla verticillata (L.f.) Royle." Protoplasma 253, no. 6 (November 16, 2015): 1529–39. http://dx.doi.org/10.1007/s00709-015-0911-5.

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20

Buapet, Pimchanok, Lewis Jie Qi Low, and Peter Alan Todd. "Differing photosynthetic responses to excess irradiance in the two coexisting seagrasses, Halophila ovalis and Halophila decipiens: Chloroplast avoidance movement, chlorophyll fluorescence, and leaf optical properties." Aquatic Botany 166 (August 2020): 103268. http://dx.doi.org/10.1016/j.aquabot.2020.103268.

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21

Samardakiewicz, Sławomir, Weronika Krzeszowiec-Jeleń, Waldemar Bednarski, Artur Jankowski, Szymon Suski, Halina Gabryś, and Adam Woźny. "Pb-Induced Avoidance-Like Chloroplast Movements in Fronds of Lemna trisulca L." PLOS ONE 10, no. 2 (February 3, 2015): e0116757. http://dx.doi.org/10.1371/journal.pone.0116757.

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22

Wen, Feng, Da Xing, and Lingrui Zhang. "Hydrogen peroxide is involved in high blue light-induced chloroplast avoidance movements in Arabidopsis." Journal of Experimental Botany 59, no. 10 (June 11, 2008): 2891–901. http://dx.doi.org/10.1093/jxb/ern147.

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23

Wen, Feng, Jinqian Wang, and Da Xing. "A Protein Phosphatase 2A Catalytic Subunit Modulates Blue Light-Induced Chloroplast Avoidance Movements through Regulating Actin Cytoskeleton in Arabidopsis." Plant and Cell Physiology 53, no. 8 (May 28, 2012): 1366–79. http://dx.doi.org/10.1093/pcp/pcs081.

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24

Maai, Eri, Shouu Shimada, Masahiro Yamada, Tatsuo Sugiyama, Hiroshi Miyake, and Mitsutaka Taniguchi. "The avoidance and aggregative movements of mesophyll chloroplasts in C4 monocots in response to blue light and abscisic acid." Journal of Experimental Botany 62, no. 9 (February 21, 2011): 3213–21. http://dx.doi.org/10.1093/jxb/err008.

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25

Wang, Jing, Yu-ping Liang, Jin-dong Zhu, Yu-xi Wang, Meng-ya Yang, Hong-ru Yan, Qian-yi Lv, Kai Cheng, Xiang Zhao, and Xiao Zhang. "Phototropin 1 Mediates High-Intensity Blue Light-Induced Chloroplast Accumulation Response in a Root Phototropism 2-Dependent Manner in Arabidopsis phot2 Mutant Plants." Frontiers in Plant Science 12 (September 27, 2021). http://dx.doi.org/10.3389/fpls.2021.704618.

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Phototropins, namely, phototropin 1 (phot1) and phototropin 2 (phot2), mediate chloroplast movement to maximize photosynthetic efficiency and prevent photodamage in plants. Phot1 primarily functions in chloroplast accumulation process, whereas phot2 mediates both chloroplast avoidance and accumulation responses. The avoidance response of phot2-mediated chloroplasts under high-intensity blue light (HBL) limited the understanding of the function of phot1 in the chloroplast accumulation process at the HBL condition. In this study, we showed that the phot2 mutant exhibits a chloroplast accumulation response under HBL, which is defective when the root phototropism 2 (RPT2) gene is mutated in the phot2 background, mimicking the phenotype of the phot1 phot2 double mutant. A further analysis revealed that the expression of RPT2 was induced by HBL and the overexpression of RPT2 could partially enhance the chloroplast accumulation response under HBL. These results confirmed that RPT2 also participates in regulating the phot1-mediated chloroplast accumulation response under HBL. In contrast, RPT2 functions redundantly with neural retina leucine zipper (NRL) protein for chloroplast movement 1 (NCH1) under low-light irradiation. In addition, no chloroplast accumulation response was detected in the phot2 jac1 double mutant under HBL, which has been previously observed in phot2 rpt2 and phot1 phot2 double mutants. Taken together, our results indicated that phot1 mediates the HBL-induced chloroplast accumulation response in an RPT2-dependent manner and is also regulated by j-domain protein required for chloroplast accumulation response 1 (JAC1).
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26

Eckstein, Aleksandra, Weronika Krzeszowiec, Agnieszka Katarzyna Banaś, Franciszek Janowiak, and Halina Gabryś. "Abscisic acid and blue light signaling pathways in chloroplast movements in Arabidopsis mesophyll." Acta Biochimica Polonica 63, no. 3 (August 2, 2016). http://dx.doi.org/10.18388/abp.2016_1382.

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Abscisic acid (ABA) and phototropins act antagonistically to control stomatal movements. Here, we investigated the role of ABA in phototropin-directed chloroplast movements in mesophyll cells of Arabidopsis thaliana. We analyzed the expression of phototropins at mRNA and protein level under the influence of ABA. PHOT1 mRNA level was decreased by ABA in the dark while it was insensitive to ABA in light. PHOT2 mRNA level was independent of the hormone treatment. The levels of phototropin proteins were down-regulated by ABA, both in darkness and light. No impact of exogenous ABA on amplitudes and kinetics of chloroplast movements was detected. Chloroplast responses in wild type Arabidopsis and three mutants, abi4, abi2 (abscisic acid insensitive4, 2) and aba1 (abscisic acid1), were measured to account for endogenous ABA signaling. The chloroplast responses were slightly reduced in abi2 and aba1 mutants in strong light. To further investigate the effect, abi2 and aba1 mutants were supplemented with exogenous ABA. In the aba1 mutant, the reaction was rescued but in abi2 it was unaffected. Our results show that ABA is not directly involved in phototropin-controlled chloroplast responses in mature leaves of Arabidopsis. However, the disturbance of ABA biosynthesis and signaling in mutants affects some elements of the chloroplast movement mechanism. In line with its role as a stress hormone, ABA appears to enhance plant sensitivity to light and promote the chloroplast avoidance response.
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27

Saewong, Chanida, Sutthinut Soonthornkalump, and Pimchanok Buapet. "Combined effects of high irradiance and temperature on the photosynthetic and antioxidant responses of Thalassia hemprichii and Halophila ovalis." Botanica Marina, September 19, 2022. http://dx.doi.org/10.1515/bot-2022-0014.

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Abstract During midday low tides, tropical intertidal seagrasses are challenged by high irradiance and high temperature. This study assessed photosynthetic and oxidative stress responses of Thalassia hemprichii and Halophila ovalis exposed to 150 and 1000 μmol photons m−2 s−1 and 30 and 40 °C for 3 h. High temperature (40 °C) significantly decreased the maximum quantum yield of both seagrasses and this heat-induced photoinhibition was exacerbated by high irradiance (1000 μmol photons m−2 s−1). High irradiance also aggravated the effects of high temperature on the effective quantum yield of T. hemprichii. Non-photochemical quenching (NPQ) of T. hemprichii was induced by both stressors with no additive effects. In contrast, NPQ of H. ovalis was induced under high irradiance at 30 °C but was inhibited at 40 °C. Nevertheless, antioxidant enzyme activity and reactive oxygen species content did not differ among treatments in either seagrass. Monitoring chloroplast distribution in H. ovalis revealed a partial inhibitory effect of high temperature on chloroplast avoidance movement under high irradiance. Our results suggest that warming events may cause detrimental impacts on shallow water seagrasses. Halophila ovalis may be more vulnerable than T. hemprichii as its photoprotection, i.e. NPQ and chloroplast avoidance movement, was hindered at high temperature.
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28

Goessling, Johannes W., Paulo Cartaxana, and Michael Kühl. "Photo-Protection in the Centric Diatom Coscinodiscus granii is Not Controlled by Chloroplast High-Light Avoidance Movement." Frontiers in Marine Science 2 (January 8, 2016). http://dx.doi.org/10.3389/fmars.2015.00115.

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29

Łabuz, Justyna, Olga Sztatelman, and Paweł Hermanowicz. "Molecular insights into the phototropin control of chloroplast movements." Journal of Experimental Botany, July 4, 2022. http://dx.doi.org/10.1093/jxb/erac271.

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Abstract Chloroplast movements are controlled by ultraviolet/blue light through phototropins. In Arabidopsis thaliana, chloroplast accumulation at low light and chloroplast avoidance at high light intensities are observed. These responses are controlled by two homologous photoreceptors, phot1 and phot2. Whereas chloroplast accumulation is triggered by both phototropins in a partially redundant manner, sustained chloroplast avoidance is elicited only by phot2. Phot1 is able to trigger only a small transient chloroplast avoidance, followed by the accumulation phase. The source of this functional difference is not fully understood, either at the photoreceptor or signalling pathway levels. In this article, we review the current understanding of phototropin functioning and try to dissect the differences which result in signalling to two distinct chloroplast responses. In the first part, we focus on phototropin structure, photochemical and biochemical activity. Next, we analyse phototropin expression and localization patterns. We also summarize known photoreceptor systems controlling movements. Finally, we focus on the role of environmental stimuli in controlling phototropin activity. All those aspects impact the signalling to chloroplast movements and raise outstanding questions about their mechanism.
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