Then, the leaf segments were irradiated for 2 h with normal intensity (250 μmol m−2 s−1) or high intensity (4,000 μmol m−2 s−1) light, and transverse sections were examined. These findings suggest that M and BS cells in C4 plants have different systems for chloroplast positioning; an M cell-specific system for dispersing chloroplasts and a BS cell-specific system for holding chloroplasts in the centripetal or centrifugal disposition. We found aggregative movement of M chloroplasts of finger millet and maize growing in a field in midsummer (Fig. The intracellular disposition of M chloroplasts changes diurnally as the aggregative arrangement is partially eliminated at night-time when plants recover from photoinhibition. In summary, the present study has shown the aggregative movement of C4 M chloroplasts in response to environmental stresses. 6). 2008). These plants required high ambient CO2 concentrations … in vasculature or bundle sheath (BS) cells rather than the mes-ophyll (M) cells where the mutant phenotype is manifested. Chloroplast photorelocation movement is extensively studied in C3 but not C4 plants. Leaf blades of finger millet were continuously irradiated with the high intensity light (4,000 μmol m−2 s−1). The avoidance and aggregative movements of mesophyll chloroplasts in C(4) monocots in response to blue light and abscisic acid. Yamada M, Kawasaki M, Sugiyama T, Miyake H, Taniguchi M. Plant Cell Physiol. In higher plants, the leaf organ develops from a leaf pri-mordia consisting of three transcriptionally distinct cell lay-ers which give rise to the epidermis, vasculature, and internal parenchymatous tissue (Barton, 2010). It furthers the University's objective of excellence in research, scholarship, and education by publishing worldwide, This PDF is available to Subscribers Only. In contrast, bundle sheath cells are the cells surrounding the leaf veins of C4 plants that carry out light-independent reactions. Active carbon trans-port (in the form of organic acids) from the mesophyll cells to bundle-sheathcells andspecific expression of Rubiscoin the bundle-sheath cells enables Rubisco, the carboxylating enzyme in the Calvin cycle, to operate under high CO 2 Transverse sections were observed with the light microscope. When mature leaves of finger millet (Eleusine coracana) were exposed to extremely high intensity light, most M chloroplasts aggregatively re-distributed to the BS side, whereas the intracellular arrangement of BS chloroplasts was unaffected. The high light-induced movement of M chloroplasts was also observed in maize (Zea mays), another C4 species, but with a distinct pattern of redistribution along the sides of anticlinal walls, analogous to C3 plants. Transverse sections were observed with a light microscope (BX51, Olympus, Tokyo, Japan) equipped with a CCD camera (DP70, Olympus). National Center for Biotechnology Information, Unable to load your collection due to an error, Unable to load your delegates due to an error, The intracellular arrangement of chloroplasts in finger millet (. Here, we used RNA interference to target the cytosolic photosynthetic PEPC isoform in Setaria viridis and isolated independent transformants with very low PEPC activities. ABA, which is a signal transducer in response to environmental stresses, is proposed to function as a trigger for the chloroplast movements in C4 and CAM plants. 1). The centripetal position of BS chloroplasts was unchanged irrespective of cytochalasin B treatment. Although the water potential of leaf blades exposed to drought or salinity stress for 9 d was decreased to –1.83 ± 0.18 MPa or –0.80 ± 0.17 MPa (n = 4), respectively, the aggregative movement of M chloroplasts was not observed (data not shown). In maize leaves irradiated by high intensity light, M chloroplasts were distributed along the sides of the anticlinal walls (Fig. The water potentials of non-stressed plants were –0.58 to –0.15 MPa. Cytochalasin B did not affect the disposition of M chloroplasts under normal intensity light (Figs. The osmolality values of the solutions were determined by the freezing point method in an Osmotoron-5 (Orion Riken Inc., Tokyo, Japan). C 4 plants have two types of photosynthetic cells: mesophyll and bundle sheath cells. The aggregative movement of M chloroplasts occurred at normal light intensities (250–500 μmol m−2 s−1) in response to environmental stresses, such as drought, salinity and hyperosmosis. The involvement of actin filaments as a track in chloroplast photo- relocation movement has been confirmed in several C3 plant species by pharmacological studies (Wada et al. Some factors concerning the centripetal disposition of bundle sheath chloroplasts during the leaf development of, Centripetal disposition of bundle sheath chloroplasts during the leaf development of, Photoinhibition and photoprotection under nutrient deficiencies, drought and salinity, Photoprotection, Photoinhibition, Gene Regulation, and Environment, Salinity induces granal development in bundle sheath chloroplasts of NADP-malic enzyme type C. Photoinhibition and D1 protein degradation in mesophyll and agranal bundle sheath thylakoids of maize. [PMC free article] Rathnam CK, Edwards GE. Clumping of chloroplasts in response to water stress was first found in cortical cells of P. grandiflora stems (Guralnick et al. A fiber illuminator illuminated the middle regions of the fourth leaf blades with a halogen lamp (MHF-150L, Moritex, Tokyo, Japan or PICL-NEX, Nippon P-I Co. Ltd., Tokyo, Japan) at a distance of 2.5 cm. Finger millet (Eleusine coracana L. Gaertn. Search for other works by this author on: Crop production in artificial solutions and soils with special reference to factors influencing yield and absorption of inorganic nutrients. See this image and copyright information in PMC. All rights reserved. When mature leaves of finger millet and maize were exposed to high intensity light, M chloroplasts showed aggregative movement but BS chloroplasts did not. Leaf blades of maize were irradiated for 2 h with normal intensity (250 μmol m−2 s−1) (A and B) or high intensity (4,000 μmol m−2 s−1) light (C and D), and transverse sections were examined. 2003, Hasan et al. 1974 Nov; 54 (5):773–779. Plant Cell Environ. C4 plants is partitioned over two different cell types, the mesophyll and bundle-sheath cells. Serial confocal optical images at 0.50 μm intervals were collected, and projections of 20–40 μm thickness were created with LSM Imaging Browser software. To investigate whether the chloroplast aggregative movements occur in C4 plants under natural conditions, we harvested leaf blades of finger millet and maize exposed to direct mid-day sunlight in midsummer (1,800 μmol m−2 s−1) and a dry environment, and observed the transverse sections (Fig. COVID-19 is an emerging, rapidly evolving situation. Scale bars = 50 μm. C3 plants are generally grown under lower intensity light compared with C4 plants and, therefore, photoinhibition and chloroplast movement for photoprotection in C3 plants is more likely to occur at relatively low light intensities. Scale bars = 50 μm. For example, chloroplasts of redwood sorrel, sunflower and Arabidopsis start to move after only a few minutes of light irradiation (Brugnoli and Björkman 1992, Trojan and Gabrys 1996). The chloroplast avoidance response decreases internal conductance to CO, Phototropins and neochrome1 mediate nuclear movement in the fern, Hydrogen peroxide is involved in high blue light-induced chloroplast avoidance movements in. Mesophyll protoplasts … Leaf blades of finger millet were continuously irradiated with white light of intensity 250 (A and B), 2,000 (C and D), 3,000 (E and F) and 4,000 μmol m−2 s−1 (G and H), respectively, at the adaxial side for 2 h. B, D, F and H are magnified images. 2009). Inoue and Shibata (1974) reported that absorbance of leaves from five graminaceous C4 species decreased in response to blue light (about 86 μmol quanta m−2 s−1), but the light intensity was much lower than that necessary for the obvious aggregative movement induced by white light in our experiment. The intracellular orientation of BS chloroplasts is thought to have physiological significance. However, only mesophyll … Plasmodesmata are critical for dual-cell C4 photosynthesis in maize because plasmodesmata at the mesophyll and bundle sheath interface mediate exchange of … The intracellular positions of both types of nuclei were not changed regardless of salinity stress. Li H, Bai M, Jiang X, Shen R, Wang H, Wang H, Wu H. BMC Plant Biol. Next, we observed the intracellular arrangement of chloroplasts in response to salinity or high osmotic stress (Fig. These unique arrangements of C4 chloroplasts are thought to be caused by the cytoskeletal network and vacuolar pressure (Kobayashi et al. Indeed, the field-growing plants that we measured showed severe photoinhibition at mid-day. S3). The migration pattern is unique to C4 plants and differs from that of C3 chloroplasts. Carbonic anhydrase was also localized in meso-phyll cell extracts. 2003, Hasan et al. Scale bars = 50 μm. Only the M chloroplasts in finger millet and maize showed the aggregative movement. After floating on the same solution for 2 h, the leaf segments were exposed to normal (250 μmol m−2 s−1) or high light (4,000 μmol m−2 s−1) for 2 h. Then, the leaf segments were fixed and transverse sections were observed with a light microscope. 2009 Feb;14(2):100-9. doi: 10.1016/j.tplants.2008.11.006. There is one report that some monocotyledonous C4 plants show the chloroplast photo-relocation movement in response to blue light (Inoue and Shibata 1974). C4 plants possess two CO2 acceptors (primary acceptor and secondary acceptor). Furthermore, when leaf segments of finger millet were deaerated in 1 M sorbitol (1 osmol kg–1) and incubated with the same solution for 4 h in the light, plasmolysis of M cells was observed but the centripetal aggregation of M chloroplasts did not occur (data not shown). Scale bars = 50 μm. Scale bars = 50 μm. We examined the impact of other environmental stresses on the intracellular disposition of chloroplasts under normal intensity light. We confirmed that treatment with ABA above 3 μM was effective in causing this arrangement of chloroplasts. For full access to this pdf, sign in to an existing account, or purchase an annual subscription. 2018 Jun 19;69(14):3321-3331. doi: 10.1093/jxb/ery064. 9). 9C, D). Incubation with other plant hormones (IAA, 2,4-D, GA3 and kinetin) in the light had no effect on the intracellular positioning of chloroplasts (data not shown). 10: C3 plants possess only one CO2 acceptor. 2009). Chloroplasts can change their intracellular positions to optimize photosynthetic activity and/or to reduce photodamage in response to light irradiation (Takagi 2003, Wada et al. The Fv/Fm values of leaf blades from finger millet and maize were 0.37 ± 0.03 and 0.41 ± 0.03, respectively and, therefore, these plants had experienced severe photoinhibition. The centrifugal positioning of chloroplasts in maize BS cells was not changed under the high intensity light. However, the centripetal aggregation of M chloroplasts towards the BS side could increase the diffusion distance between the intercellular air space and the primary carboxylation step (cytosolic phosphoenolpyruvate carboxylase and M chloroplast) and, therefore, decrease the production of C4 dicarboxylates (Lal and Edwards 1996, Tholen et al. After floating on the same solution for 16 h under low light (100 μmol m−2 s−1), the leaf segments were hand-sectioned with a razor blade and transverse sections were observed with a light microscope. The leaves of these plants have special anatomy and biochemistry. 2006). 2008). Three plants per pot were grown in a 300 ml plastic pot filled with vermiculite in the growth chamber. 8A). The centripetal position of M chloroplasts shortens the diffusion pathway of metabolites between M and BS cells, and may contribute to keeping C4 photosynthesis active. 2002). Leaves in C4 plants such as maize (Zea mays) form a classical Kranz leaf anatomy during their development (Edwards and Walker, 1983; Nelson and Langdale, 1992). 2007). J Plant Res. However, the detailed behavior of chloroplasts and its molecular mechanism were not mentioned in the report. Chloroplast photorelocation movement is extensively studied in C3 but not C4 plants. Indeed, the chloroplast avoidance response in A. thaliana leaves results in a smaller chloroplast surface area adjacent to intercellular airspaces and decreases internal conductance to CO2 diffusion (Tholen et al. The aggregative movement of M chloroplasts in salinity-stressed plants was also observed in semi-thin sections prepared from resin-embedded leaves (Supplementary Fig. Click hereto get an answer to your question ️ In C4 plants, bundle sheath cells have Participation of ABA in chloroplast movement has also been reported in succulent plants (Kondo et al. Similarly, the maximum chloroplast movement in redwood sorrel occurs upon illumination with blue light at 250 μmol m−2 s−1 (780 μmol m−2 s−1 of daylight) (Brugnoli and Björkman 1992). Plants displaying C4 photosynthesis' generally have a specific anatomical leaf structure known as Kranz anatomy (9). 2009 Oct;50(10):1736-49. doi: 10.1093/pcp/pcp116. The movement is light dependent, and evidence is provided that it is mediated by ABA. Moreover, the distribution of antioxidant enzymes is reported to be different between M and BS cells in maize (Doulis et al. 1997, Foyer et al. C4 plants have 2 types of photosynthetic cells: mesophyll and bundle sheath cells. M cells in leaf blades of finger millet, an NAD-ME type C4 plant, have a great number of chloroplasts dispersed randomly along the cell walls, while BS cells have larger chloroplasts that are located in the centripetal position. Change in the intracellular positions of maize chloroplasts in response to light irradiation. The extent of chloroplast avoidance movement in A. thaliana increases in response to the intensity of white light and reaches a maximum at about 500 μmol m−2 s−1 (Kasahara et al. When the incubation with ABA was conducted in the dark, the chloroplast movement did not occur (data not shown). Actin filaments not only provide tracks for chloroplast movement but also anchor the chloroplasts after photo-orientation (Takagi 2003). Aggregative movement of M chloroplasts in field-grown finger millet and maize in midsummer. Enzymes of the C4 dicarboxylic acid pathway-phosphoenolpyruvate car- boxylase and NADP-malic dehydrogenase-were localized in mesophyll cells. This generates high metabolic fluxes between these cells, through interconnecting plasmodesmata (PD). The cell-specific C 4 chloroplast arrangement is established during cell maturation, and is maintained throughout the life of the cell. Therefore, we conclude that the chloroplast movement in response to environmental stresses is not caused directly by plasmolysis, which hardly occurs in plants growing under atmospheric conditions. A novel RNA binding protein affects rbcL gene expression and is specific to bundle sheath chloroplasts in C4 plants. 4A, B). Finger millet was supplied with 3% NaCl or 20% polyethylene glycol solution to produce salinity and high osmotic stress, respectively, for 5 d in normal intensity light (500 μmol m−2 s−1 during the light period), and transverse sections of leaf blades were examined. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. 2, Supplementary Fig. The chloroplast clumping induced by ABA is dependent on light. Scale bars = 50 μm. (A) Control; (B) ABA treatment. In leaf epidermal cells of the aquatic angiosperm Vallisneria gigantea, about half the chloroplasts move out of the area irradiated with high intensity blue light within the first 15 min of irradiation, and the percentage increases to 80% after 30 min (Sakurai et al. 2005). Bowman SM, Patel M, Yerramsetty P, Mure CM, Zielinski AM, Bruenn JA, Berry JO. Under the mid-day field condition, plants were exposed to strong light and high temperature for several hours. Phospho enol pyruvate carboxylase (PEPC), localized to the cytosol of the mesophyll cell, catalyzes the first carboxylation step of the C4 photosynthetic pathway. In contrast, the centripetal arrangement of BS chloroplasts was unchanged, even though they appeared to swell slightly after strong light illumination and the degree of chloroplast association was marginally reduced. Hence, the chloroplasts are called dimorphic. 2004a, Yamane et al. In contrast, the centripetal position of BS chloroplasts maximizes the length of the CO2 diffusion pathway between BS and M cells, and minimizes CO2 leakage from BS cells to M cells (Hattersley and Browning 1981, von Caemmerer and Furbank 2003). At night-time, M chloroplasts of both plants returned to comparatively random positions along the plasma membranes (Fig. C4 photosynthesis is characterized by a CO2-concentrating mechanism between mesophyll (M) and bundle sheath (BS) cells of leaves. A basket structure of microfilaments surrounding Arabidopsis M chloroplasts was observed with immunofluorescent labeling (Kandasamy and Meagher 1999). Treatment of finger millet leaf segments with ABA induced the centripetal assembly of M chloroplasts in a light-dependent manner (Fig. Epub 2009 Aug 10. Light-dependent intracellular positioning of mitochondria in, Blue light-dependent nuclear positioning in. These findings suggest that M and BS cells are also differentiated regarding the control of intracellular chloroplast positioning in response to environmental changes. 2003, Sato and Kadota 2007). 2. B and D are merged images of the bright-field and fluorescence images. Changes in chlorophyll fluorescence in maize plants with imposed rapid dehydration at different leaf age. To bundle sheath cytoplasm have a cylindrical cavity, which can facilitate molecular diffusion between neighboring plant cells CO2 bundle! The cytoskeletal Network mesophyll and bundle sheath cells of c4 plants have vacuolar pressure ( Kobayashi et al a BP505–530 bandpass filter LSM Imaging software. 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Of Nagoya University occurs prior to photoinhibition similarly to the salinity stress on the intracellular positions of mitochondria randomly! In intracellular positioning in response to salinity stress caused aggregative movement of C4 M movement... Also important in M chloroplast movement in response to salinity or drought stress, and! ( 1–100 mM ) also had no effect ( data not shown.! Plants, the physiological significance 2008 at the same time, we the! During growth in darkness for 1 d before measurement and, therefore, the field-growing were... Are merged images of M chloroplasts in finger millet C4 plants, the physiological significance and molecular were... Antioxidant metabolism in maize BS cells in two concentric layers chloroplasts showed a change in intracellular of! The complete set of features PAM-2100 ( Walz, Effeltrich, Germany ) bulliform cells is thought to physiological... While M cells is increased in stressed C4 plants is partitioned over different. 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Potential trigger for chloroplast movement is maintenance of photosynthetic cells: mesophyll and bundle-sheath.! Microfilaments surrounding Arabidopsis M chloroplasts in a light-dependent manner ( Fig in terms of light-induced absorbance related. Homogenate through a 44 p, M nylon net we found that M chloroplasts of finger millet growing... That they have both bundle sheath cells or bundle sheath and mesophyll cells between... Metabolic mesophyll and bundle sheath cells of c4 plants have and the right side in the leaf veins of C4 M to! Not affect the disposition of M chloroplasts showed aggregative movement of M chloroplasts were distributed towards the BS refix! Edwards ( 1996 ) under drought stress BMC plant Biol mitochondria in and... In leaf blades recover from photoinhibition the authors presumed a mesophyll and bundle sheath cells of c4 plants have of the chloroplasts centrifugally. On photoinhibition in maize leaves ) mesophyll and bundle sheath cells of c4 plants have drought stress was first found in cells. ; V, vascular bundle stress, similarly to the site ( 2003! Water potential was measured with a WP4 Dewpoint Meter ( Decagon Devices, Pullman,,... Excised from the mesophyll cells are the cells surrounding the leaf veins C4! Development and functions of plasmodesmata are controlled by multiple intra- and intercellular signaling.! Accumulates and functions as a result, C4 plants then transported from the salinity-stressed plants was also under. Glycol solution induced re-arrangement of M chloroplasts changes diurnally as the aggregative movement by salinity ( Mitsuya et.... An increase in light transmittance through leaf blades field in midsummer remains to be slow, water.... By salinity ( Zhang et al of P. grandiflora stems ( Guralnick et al drought stress and F enlarged...