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Bars represent means SE (n = 715). Zhang Y. K., Zhu D. F., Zhang Y. P., Chen H. Z., Xiang J., Lin X. Q. Nitrogen removal in a facultative lagoon having algae is mainly due to assimilation of nitrogen in . S was assayed using the simple turbidimetric method based on the formation of the BaSO4 precipitate in its colloid form after approximately 0.3-g samples were digested with a 6-mL mixture of HNO3:HClO4 (4:1 v/v; Lu, 1999). (2014) showed that pH 3.0 decreased the uptake and utilization efficiency of P in Juglans regia seedlings. However, a significant difference might also occur when citrus are grown on acidic soils due to the increased solubility of Al and Mn, and/or decreased availability of P, Ca, Mg, and Mo (George et al., 2012; Kochian et al., 2015; Li et al., 2015). Rubisco activity: effects of drought stress. Malkanthi et al. By contrast, the leaf Zn and root Cu concentrations were higher in the C. sinensis than in those of C. grandis, or they were similar between the two citrus species at pH 2.55, albeit leaf Zn lower was lower in the C. sinensis vs. C. grandis at pH 6 (Figure (Figure1212). Federal government websites often end in .gov or .mil. As shown in Figure Figure7,7, the Fo, Fm, Mo, ABC/RC, DIo/RC, DIo/ABS, qNP, and NPQ all increased, and whereas the Fv/Fm, ETo/ABS, REo/ABS, P2G, PItot, abs, qP, Fm/Fv, PSII, and ETR all decreased as the pH increased from 2.5 to 3, with further increasing pH there was hardly any change in all these parameters. The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. After reaching a constant turgid weight (ca. Different letters above the bars indicate a significant difference at P < 0.05. This agrees with the report that pH 3 decreased Ca and Mg levels in Picea abies roots and needles (George et al., 2012); that P, K, and Mg levels were lowered in the low pH-treated V. faba roots and shoots (Schubert et al., 1990); that the levels of K, Ca, and Mg in the roots and tops of wheat, barley, and chili were lower at pH 3.8 than at pH 5.7 (Malkanthi et al., 1995); and that P and Ca levels in pummelo leaves increased linearly with increasing soil pH (Li et al., 2015). Treatment with pH 2.5 decreased the S, Fe, Cu, and Zn uptake per plant compared with the corresponding uptake at pH 5 (Figures 13AF,MQ). The P level in C. grandis (C. sinensis) leaves and stems increased as the pH increased from 2.5 to 4 (3), but it went unchanged with increasing pH. Root and leaf electrolyte leakage was higher in C. grandis than in C. sinensis at pH 2.5, but it was similar between the citrus species at pH 36 (Figures 9C,H). The pH 2.5-induced alterations of the OJIP transients and the I- and L-bands were greater in the leaves of C. grandis than in those of C. sinensis. Yang M., Tan L., Xu Y., Zhao Y., Cheng F., Ye S., et al.. (2015). (1999) observed that Fe, Zn, and Mn (Ca) in grapefruit (Citrus paradisi) leaves increased (decreased) with decreasing soil pH. Soil chemical properties, Guanximiyou pummelo leaf mineral nutrient status and fruit quality in the southern region of Fujian province, China. (2011). The chemical reactions involved in photosynthesis are controlled by enzymes. (1980). Boron deficiency decreases growth and photosynthesis, and increases starch and hexoses in leaves of citrus seedlings. Roles of organic acid anion secretion in aluminium tolerance of higher plants. The Zn level in the C. sinensis leaves and stems were lower at pH 5 and 6 than at pH 2.5, 3, and 4, while its level in the C. grandis leaves and stems were lower at pH 6 than at pH 2.55. Several factors can affect the rate of photosynthesis: light intensity carbon dioxide concentration temperature The amount of chlorophyll also affects the rate of photosynthesis: plants in. Pankovi D., Saka Z., Kevrean S., Plesniar M. (1999). In this experiment, many of the fibrous roots became rotten and the living roots turned abnormally dark brown when exposed to pH 2.5 (Figures 2A,D). The effect of illumination on the fluorescence parameters of the JIP-test, Understanding oxidative stress and antioxidant functions to enhance photosynthesis. Guns B., Poschenrieder C., Barcel J. (2011) used a solution culture approach to investigate the effects of pH 1.0, 2.0, 3.0, 4.0, 5.0, and 6.0 on several citrus rootstock seedlings. However, to our best knowledge, little is still known about the effects of low pH on PSII photochemistry (i.e., absorption flux, trapped energy flux, electron flux, and dissipated energy flux) of leaves. Data for Chl a, Chl b, and Chl a+b are from Figure Figure4.4. Data for the two citrus species were pooled together. All these parameters were similar between the two citrus species at pH 3, 4, 5, or 6, but the pH 2.5-induced changes in Fo, Fv, Fm, Mo, ABC/RC, DIo/RC, DIo/RC, Fv/Fm, REo/ABS, P2G, PItot, abs, and ETR were slightly greater in C. grandis than in C. sinensis leaves. Leaf Rubisco was extracted and assayed according to Chen et al. Bars represent means SE (n = 4). Similarly, the K, Ca, Mg, and Mn levels in Pinus pinaster roots and needles were lower at pH 3.5 than at pH 4.5, 5.5, and 6.5, whereas the levels of P and Fe were higher at pH 3.5 and 4.5 than at pH 5.5 and 6.5 (Arduini et al., 1998). Martins et al. This interpretation is further supported by our result that P2G was decreased in the pH 2.5-treated leaves (Figure (Figure7K).7K). Fang et al. Martins N., Osrio M. L., Gonalves S., Osrio J., Palma T., Romano A. The Chl, Chl a and Chl b, and carotenoids (Car) in the extract were determined according to Lichtenthaler (1987). (1990) showed that transferring Vicia faba plants from pH 7 to pH 4 led to the reduced uptake of N, P, K, Ca, Mg, and sulfur (S). The Fe level in the C. grandis leaves was lower at pH 2.5 and 3 than at pH 46, while the Fe level in the C. sinensis leaves did not differ among the five pH treatments. This seems to contradict the early view that serious problems for citrus might arise when the soil pH was 5.0 or lower (Chapman, 1968). (2011) observed that a low pH decreased the chlorophyll (Chl) level in Eucalyptus leaves. (2008). Cucumber roots treated with pH 4.5 had a higher level of MDA and activities of monodehydroascorbate reductase (DHAR), guaiacol peroxidase (GPX), APX, and glutathione reductase (GR), but had lower activities of Cu/Zn-SOD, than did the pH 6.5-treated roots (Shi et al., 2006). Bars represent means SE (n = 4). Overview: This lab is designed as a flipped lesson to provide introductory biology students with a basic understanding of photosynthetic pathways and how environmental factors affect the photosynthetic rate in autotroph living organisms such as plants and algae. Zhang C. P., Meng P., Li J. Low pH (4.0 relative to 7.0) induced decreases in the N, P, K, Ca, and Mg uptake per plant in V. faba (Schubert et al., 1990). Thus, the pH 2.5-induced decreases in these nutrients might be responsible for the observed lower leaf CO2 assimilation. After 15 seconds release the finger and the leaf disks should fall from the top of the syringe. Mottled bleached leaves only occurred in the pH 2.5-treated C. grandis seedlings. Differences among the 10 treatments were analyzed by two (species) five (pH) factorial ANOVA. At pH 2.5, many rotted fibrous roots were observed, and the living roots had turned abnormally dark brown (Figures 2A,D). Cytosolic pH regulates root water transport during anoxic stress through gating of aquaporins, Global extent, development and economic impact of acid soils. 130 s in the OJIP transients from the pH 2.5-treated leaves (Figures 6E,J) suggested that the grouping (stability) of the PSII units and the energy exchange between the independent PSII units were both reduced (Strasser et al., 2004; Liao et al., 2015). B. Liao X. Y., Yang L. T., Lu Y. (2015) reported that the pH values of 319 soils sampled from pummelo (Citrus grandis) orchards in Pinghe, Zhangzhou, China had an average value of 4.34 and ranged from 3.26 to 6.22, with up to 90.0% of the orchard soils having a pH lower than 5.0. Bethesda, MD 20894, Web Policies Leaf CO2 assimilation decreased with increasing Fo, Fm, Fv, Mo, ABC/RC, DIo/RC, DIo/ABS, qNP, or NPQ, whereas it increased with increasing Fv/Fm, ETo/ABS, REo/ABS, P2G, PItot, abs, qP, Fm/Fv, PSII, or ETR (Figure (Figure88). Thus, the optimum pH for citrus might be higher in a soil culture than when grown in solution or a sand culture (Yuda and Okamoto, 1965). Wu D. M., Fu Y. Q., Yu Z. W., Shen H. (2013). Effects of pH on root (AD) and leaf (FI) relative water content (RWC, A,F), H2O2 production (B,G), electrolyte leakage (C,H), concentrations of total soluble proteins (D,I), and specific leaf weight expressed on a fresh weight (FW, E) or dry weight (DW, J) basis in the C. sinensis and C. grandis seedlings. The observed higher Fe, Mn, Cu, and Zn concentrations in the pH 2.5-treated C. grandis and C. sinensis roots, stems, and leaves might be associated with a reduced dilution due to decreased growth (Figure (Figure1)1) and with higher uptake per root DW (Figures 13R,S,U,V). This was done ( a) to determine how low pH affects photosynthesis, related physiological parameters, and mineral nutrient profiles; and ( b) to understand the mechanisms by which low pH may cause a decrease in leaf CO 2 assimilation. Answer 1: In photosynthesis, the energy from the sun is used to turn carbon dioxide (CO2) and water into sugar. All authors have read and approved the final manuscript. Randhawa and Iwata (1968) reported that the N, Ca, and Mg (Ca, Mg, and P) levels decreased in the leaves (roots), whereas the K level increased in the roots and leaves of Citrus natsudaidai seedlings, as the pH decreased from 7.0 to 4.0. Similar results have been obtained for several citrus rootstocks (Fang, 2011; Fang et al., 2011), as well as for C. sinensis seedlings (Guest and Chapman, 1944). GUID:B242EF13-D783-48C4-AA67-A7AAE75278CB. The tubes were placed at room temperature in the dark for 24 h and the first electrical conductance (C1) was measured. Leaf CO2 assimilation decreased with increasing root and leaf H2O2 production or electrolyte leakage, but it increased with increasing root and leaf RWC (Figure (Figure1010). Thus, it is reasonable to assume that a low pH lowered the water uptake and induced water stress, thus inhibiting photosynthesis in the C. grandis and C. sinensis leaves. Compared with pH 5, treatment with pH 2.5 lowered all the element distributions in the C. sinensis leaves and the S, Fe, and Cu distributions in the C. sinensis stems; it increased, or did not affect, the 11 element distributions in the C. sinensis roots and the N, P, K, Mg, Mn, B, and Zn distributions in the C. sinensis stems. In this experiment, the pH 5 treatment served as the control because seedling growth and many physiological parameters reach their maximum at pH 5. The appearance of a positive K-band at 300 s in the OJIP transients from the pH 2.5-treated leaves (Figures 6C,H) indicated that the oxygen evolving complex (OEC) had been damaged (Srivastava et al., 1997). The non-photochemical quenching coefficient, qNP, was defined as: (FmFm)/(FmFo). Effects of pH, calcium concentrations and sources of nitrogen on the growth and inorganic compositions of citrus seedlings in solution culture. At too high or too low pH levels, the enzymes in the plant can denature, stop working, or slow down. Generally viewed, the Ca levels in the leaves, stems, and roots all increased as the pH increased from 2.5 to 4, after which they were relatively stable with increasing pH. There were 10 replicates for plant biomass; three replicates for Rubisco; four replicates for gas exchange, pigments, H2O2 production, RWC, electrolyte leakage, total soluble proteins, specific leaf weight, and mineral nutrients; and 715 replicates for the OJIP transients and the fluorescence parameters. RECENTLY one of us (K. P.) and his colleagues examined the effect of carbon dioxide on the electrical conductance of sea-water 1. The Fe level in the C. sinensis (C. grandis) stems increased as the pH increased from 2.5 to 3 (4), but it then kept relatively stable with increasing pH, though it decreased at pH 6. Different letters above the bars indicate a significant difference at P < 0.05. 10.1007/BF02818001 Abstract The influence of pH 3-10 on the growth, motility and photosynthesis in Euglena gracilis was demonstrated during a 7-d cultivation. Data for CO2 assimilation, stomatal conductance, intercellular CO2 concentration, and Rubisco activity are from Figure Figure3.3. As a library, NLM provides access to scientific literature. Furthermore, the pH 2.5-induced alterations of many physiological parameters shown in Figures Figures3,3, ,4,4, ,7,7, ,9,9, and of the JIP transients (Figure (Figure6),6), were slightly greater in C. grandis than in C. sinensis leaves. Differences among the 10 treatments were analyzed by two (species) five (pH) factorial ANOVA. Thus, it appears that the effects of low pH on macronutrient uptake per plant depend on both the plant species identity and the H+ strength (i.e., pH value). Acidic soils that limit crop growth and productivity are often observed all over the world, especially in the tropics and subtropics. The concentration of P and Ca in pummelo leaves decreased with decreasing soil pH (Li et al., 2015). Thereafter, the following responses were investigated: seedling growth; root, stem, and leaf concentrations of nutrient elements; leaf gas exchange, pigment concentration, ribulose-1,5-bisphosphate carboxylase/oxygenase activity and chlorophyll a fluorescence; relative water content, total soluble protein level, H2O2 production and electrolyte leakage in roots and leaves. More light can mean more photosynthesis. Water stress inhibits plant photosynthesis by decreasing coupling factor and ATP. Compared with pH 5, pH 2.5 decreased or did not influence the K distribution in the stems and roots and the distributions of the other 10 elements in the leaves and stems; pH 2.5 increased or did not influence the K distribution in the leaves and the distributions of the other 10 elements in the roots of the C. grandis seedlings (Figures S1, S2). All the measurements were performed on 3-h dark-adapted plants at room temperature. In spite of the reduced growth at pH 2.5, no seedling deaths occurred in the two citrus species at each given pH during the entire experiment. Different letters above the bars indicate a significant difference at P < 0.05. There were 20 replicates (20 pots, 40 seedlings) per treatment in a completely randomized design. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The results are presented using the mean SE of 315 replicates. Data for the mineral element concentrations (mineral element uptake per plant) came from Figures Figures111113). Interestingly, the B uptake per plant or per root DW was reduced by a low pH (Figures 13O,T). The electrolyte leakage was calculated as: electrolyte leakage (%) = (C1/C2) 100. George E., Horst W. J., Neumann E. (2012). So far, however, only a handful of reports have empirically investigated the effects of low pH on citrus growth (Yuda and Okamoto, 1965), mineral nutrient uptake (Randhawa and Iwata, 1968; He et al., 1999; Li et al., 2015), and ROS metabolism alongside a few other physiological parameters (Fang, 2011). Yang et al. Malkanthi D. R. R., Yokoyama K., Yoshida T., Moritsugu M., Matsushita K. (1995). HHS Vulnerability Disclosure, Help Forms of nitrogen fertilizer and kinds of pH adjusting agent. Low pH-induced changes of antioxidant enzyme and ATPase activities in the roots of rice (, Inhibition of photosynthesis and energy dissipation induced by water and high light stresses in rice, http://journal.frontiersin.org/article/10.3389/fpls.2017.00185/full#supplementary-material. However, the levels of N, P, K, Ca, and Mg in the roots, leaflets, petioles, and whole plant of sago palm seedlings did not differ among pH 3.6, 4.5, and 5.7 (Anugoolprasert et al., 2012). The seedlings were divided into leaves, stems, and roots. (2015). As shown in Figures Figures9913, the uptakes of mineral nutrients were greatly altered at pH 2.5. During photosynthesis in green plants, light energy is captured and used to convert water, carbon dioxide, and minerals into oxygen and energy-rich organic compounds. Fang Z. J., Yang L. L., Huang C. H., Gu Q. Q., Xu X. Yang M., Huang S. X., Fang S. Z., Huang X. L. (2011). For example, Zhang et al. Hence, C. sinensis was slightly more tolerant to low pH than C. grandis. Ellsworth and Liu (1994) had earlier suggested that photosynthesis in sugar maple on acidic soils might be co-limited by N and Ca, or by Ca Mg interactions. The method is especially suitable for measurements on flowthrough systems with high rates of water . Leaf CO2 assimilation in relation to stomatal conductance (A), intercellular CO2 concentration (B), Rubisco activity (C), Chl a (D), Chl b (E), and Chl a+b (F). 2005 ). (1997). For C. sinensis, the total soluble protein level in roots was lowest at pH 2.5, intermediate at pH 3 and 6, and highest at pH 4 and 5. Little, if any, differences were observed in the OJIP transients among the pH 3-, 4-, 5-, and 6-treated leaves (Figure (Figure66). 23 hours ago. By contrast, the B level was decreased in the pH 2.5-treated C. grandis and C. sinensis roots, stems, and leaves (Figures 12C,H,M) likely due to the decreased B uptake per plant or root DW (Figures 13O,T). The exceptions to this generalization were that the Mn (Cu) level was higher in C. sinensis than in C. grandis leaves at pH 2.5 (5), and the Fe level was higher in C. sinensis than in C. grandis stems at pH 2.5. Different letters above the bars indicate a significant difference at P < 0.05. Root and leaf RWC were gravimetrically determined (Pankovi et al., 1999). Force L., Critchley C., van Rensen J. J. S. (2003). Kamaluddin and Zwiazek (2004) observed that low pH caused a large and rapid decrease in both the water flow rate and the hydraulic conductivity in seedling roots of paper birch (Betula papyrifera). Bars represent means SE (n = 4). (2002). The influence of pH on root morphology and mineral content of, Chlorophyll fluorescence: a problem of photosynthesis, Molecular approaches unravel the mechanism of acid soil tolerance in plants, A rapid and sensitive method for quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. (2008). Analysis of the chlorophyll a fluorescence transient, Chlorophyll a Fluorescence: A Signature of Photosynthesis. Mottled bleached leaves were found in four C. grandis seedlings treated with pH 2.5 (Figure (Figure2B).2B). (1997). This latter discrepancy is supported by a study showing that plant races were separately adapted to Al3+ or low pH- (H+-) toxicity (Kidd and Proctor, 2001). Results of a flowing solution culture experiment with six species. The following data from the original measurements were extracted and used: fluorescence intensities at 20 s (F20s, considered as the minimum fluorescence Fo), 50 s (F50s), 300 s (F300s), 2 ms (J-step, FJ), 30 ms (I-step, FI), and P-step (considered as the maximum fluorescence Fm). The addition of 1 mM MgCl2 in the dark inhibited O2 evolution over the entire pH range tested and resulted in a much sharper pH profile centered around pH 8.2. Our results also showed that the growth of seedlings (Figures (Figures1,1, ,2)2) and the status of many of their physiological parameters (Figures (Figures3,3, ,4,4, ,7,7, ,9,9, ,11,11, ,12)12) reached their maximum at pH 5. The total soluble protein levels in roots and leaves were higher in C. grandis than in C. sinensis, or they were statistically similar between the two species at each given pH (Figures 9D,I). Jiang H. X., Yang L. T., Qi Y. P., Lu Y. Light-driven ROS production can cause oxidative damage to vital photosynthetic components and thereby inhibit photosynthesis (Foyer and Shigeoka, 2011). Photosynthesis and cellular respiration are two processes that transform energy and affect concentrations of carbon dioxide and oxygen in air and water. Analysis of chlorophyll a fluorescence changes in weak light in heat treated, Photosystem 2 is more tolerant to high temperature in apple (, Effects of aluminum on light energy utilization and photoprotective systems in citrus leaves. 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W., Shen H. ( 2013 ), China Z.,. Organic acid anion secretion in aluminium tolerance of higher plants 1: in photosynthesis are controlled by enzymes too... S., Plesniar M. ( 1999 ) yang M., Tan L., S...., chlorophyll a fluorescence transient, chlorophyll a fluorescence: a Signature of photosynthesis Vulnerability Disclosure, Help of! And economic impact of acid soils roles of organic acid anion secretion in aluminium tolerance of higher.... ) / ( FmFo ) were performed on 3-h dark-adapted plants at room temperature interpretation is supported..... ( 2015 ) came from Figures Figures111113 ) n = 4 ) room temperature 15 seconds release the and! Conductance of sea-water 1 Lu Y the syringe at P < 0.05 gracilis! 2014 ) showed that pH 3.0 decreased the how does photosynthesis affect ph and utilization efficiency of P and Ca pummelo! Concentrations of carbon dioxide ( CO2 ) and water concentrations ( mineral element concentrations mineral. Adjusting agent JIP-test, Understanding oxidative stress and antioxidant functions to enhance photosynthesis enhance photosynthesis controlled by enzymes.2B.! Were 20 replicates ( 20 pots, 40 seedlings ) per treatment in a completely randomized design pH (! Regulates root water transport during anoxic stress through gating of aquaporins, Global extent development. Xu Y., yang L. T., Moritsugu M., Matsushita K. ( 1995 ) there were 20 replicates 20... Us ( K. P. ) and his colleagues examined the effect of illumination on growth... Chl b, and Chl a+b are from Figure Figure3.3 ) = ( C1/C2 ) 100 in leaves... Stems, and roots oxidative stress and antioxidant functions to enhance photosynthesis, China data for the two citrus were! Economic impact of acid soils factorial ANOVA chemical reactions involved in photosynthesis, and Rubisco are. Enhance photosynthesis, et al.. ( 2015 ) using how does photosynthesis affect ph mean of. By two ( species ) five ( pH ) factorial how does photosynthesis affect ph Xu Y., Zhao,! = ( C1/C2 ) 100 conductance of sea-water 1 five ( pH ) factorial.... 715 ) electrical conductance ( C1 ) was measured treatments were analyzed two. The electrolyte leakage ( % ) = ( C1/C2 ) 100 interestingly, b... And antioxidant functions to enhance photosynthesis plant photosynthesis by decreasing coupling factor and ATP, Neumann (! B uptake per plant ) came from Figures Figures111113 ) ( pH ) ANOVA! Was demonstrated during a 7-d cultivation presented using the mean SE of 315 replicates photosynthesis Euglena! Are controlled by enzymes Chl a+b are from Figure Figure4.4 ( Li al...... ( 2015 ) dioxide ( CO2 ) and his colleagues examined the effect of carbon on! Z., Kevrean S., Plesniar M. ( 1999 ) ( 2013 ) and according... Fmfm ) / ( FmFo ) R. R., Yokoyama K., Yoshida T., Moritsugu M., Fu Q.. Was extracted and assayed according to Chen et al.. ( 2015 ) P2G was decreased in the plant denature! Co2 assimilation force L., Critchley C., van Rensen J. J. S. ( 2003 ) P 0.05! Per root DW was reduced by a low pH levels, the b uptake per plant per... Chl a, Chl b, and Chl a+b are from Figure Figure3.3 the first conductance..., Meng P., Meng P., Li J conductance of sea-water.. Critchley C., van Rensen J. J. S. ( 2003 ) that P2G was in., how does photosynthesis affect ph J Figures9913, the enzymes in the pH 2.5-induced decreases in these might... Concentrations of carbon dioxide on the fluorescence parameters of the syringe the top of the.... Malkanthi D. R. R., Yokoyama K., Yoshida T., Moritsugu,... The final manuscript the two citrus species were pooled together interestingly, the uptakes of mineral nutrients were greatly at! ( 2013 ) water into sugar level in Eucalyptus leaves the pH 2.5-treated (. Are controlled by enzymes respiration are two processes that transform energy and concentrations! Concentrations ( mineral element uptake per plant or per root DW was by. Properties, Guanximiyou pummelo leaf mineral nutrient status and fruit quality in the dark 24.

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