Photosynthetic changes that occur during aging of cypress (Cupressus sempervirens L.) needles

2-years-old cypress needles (A2) were physiologically most active with regard to net photosynthetic (P _N) and electron transport rates. Variable to maximum fluorescence (F_v/F_m) ratios of dark-adapted needles were higher in A2 needles than in current year (A1) or senescent (A4) needles. Lower F_v/F_m values in these stages seemed to be caused not by photoinhibition but by a low photochemical capacity as suggested from the chlorophyll (Chl) a/b ratios. In isolated thylakoids, lower rates of whole chain and photosystem 2 (PS2) activities were observed in A4 needles, while higher rates were observed in A2 needles. A similar trend was noticed for contents of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPC) and total soluble proteins. The artificial exogenous electron donor Mn²⁺ failed to restore the loss of PS2 activity in 3-year-old (A3) and A4 needles, while diphenylcarbazide and NH₂OH significantly restored the loss of PS2 activity. The marked loss of PS2 activity in A4 needles was primarily the result of the loss of 33, 28–25, 23, and 17 kDa polypeptides. A marked loss of RuBPC activity in A4 needles is mainly due to the loss of 15 (SSU) and 55 (LSU) kDa polypeptides.     

Photosynthetic Characteristics of Non-Leaf Organs of Winter Wheat Cultivars Differing in Ear Type and their Relationship with Grain Mass Per Ear

Chlorophyll content, photosystem 2 functioning (F_v/F_m, F_v/F₀), activity of ribulose-1,5-bisphosphate carboxylase/oxygenase, and net photosynthetic rates (P _N) of flag leaf blade, sheath, peduncle, and ear organs were assessed in large-ear type (Pin 7) and small-ear type (ND93) wheat cultivars. Some differences were found in photosynthetic properties between different green plant parts, the values of all studied parameters in ear parts being higher in Pin7 than in ND93. Furthermore, ear surface areas and ear P _N in 26 wheat genotypes measured at anthesis showed highly significant positive correlation with grain mass per ear. Hence a greater capability of ear photosynthesis may result in a greater grain yield in large-ear type cultivars.     

Photosynthetic Characteristics of Two New Chlorophyll b-Less Rice Mutants

Two yellow rice mutants VG28-1 and VG30-5 were obtained during the tissue culture process from a rice plant (cv. Zhonghua No.11 japonica) with inserted maize Ds transposon element. Absorption spectra and pigment composition showed that two mutants had no chlorophyll (Chl) b and lower Chl a content in comparison to the wild type (WT). Net photosynthetic rate (P _N), total electron transport rate (J_F), photochemical quenching (q_p), quantum yield of PS2 dependent non-cyclic electron transport (_PS2), fraction of P_rate, and leaf area were lower but F_v/F_m and apparent quantum yield (AQY) remained at similar levels as in the WT plant. Xanthophyll cycle pool size (V+A+Z) on a Chl basis, and de-epoxidation state were enhanced in the mutants. The mutants had larger amounts of soluble protein and ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO), especially the small subunit of RuBPCO, than WT. The characteristics of two rice mutants differed somewhat from the other common Chl b-less mutants originating from mutagenic agent treatments.     

Effect of Iron Deficiency Induced Changes on Photosynthetic Pigments,Ribulose-1,5-Bisphosphate Carboxylase,and Photosystem Activities in Field Grown Grapevine (Vitis Vinifera L. cv. Pinot Noir) Leaves

The effect of iron deficiency on photosynthetic pigments, ribulose-1,5-bisphosphate carboxylase (RuBPC), and photosystem activities were investigated in field grown grapevine (Vitis vinifera L. cv. Pinot noir) leaves. The contents of chlorophyll (Chl) (a+b) and carotenoids per unit fresh mass showed a progressive decrease upon increase in iron deficiency. Similar results were also observed in content of total soluble proteins and RuBPC activity. The marked loss of large (55 kDa) and small (15 kDa) subunits of RuBPC was also observed in severely chlorotic leaves. However, when various photosynthetic electron transport activities were analysed in isolated thylakoids, a major decrease in the rate of whole chain (H₂O methyl viologen) electron transport was observed in iron deficient leaves. Such reduction was mainly due to the loss of photosystem 2 (PS2) activity. The same results were obtained when F_v/F_m was evaluated by Chl fluorescence measurements in leaves. Smaller inhibition of photosystem 1 (PS1) activity was also observed in both mild and severely chlorotic leaves. The artificial electron donors, diphenyl carbazide and NH₂OH, markedly restored the loss of PS2 activity in severely chlorotic leaves. The marked loss of PS2 activity was evidently due to the loss of 33, 23, 28-25, and 17 kDa polypeptides in iron deficient leaves.     

Cd2+ Effect on Photosynthetic Apparatus in Synechococcus elongatus and Spinach (Spinacia oleracea L.)

Thylakoid membranes (TM) of the cyanobacterium Synechococcus elongatus were exposed for 30 min to the influence of 0, 10, 100, and 1 000 mM CdCl₂ (= Cd₀, Cd₁₀, Cd₁₀₀, and Cd₁₀₀₀). Cd₁₀ and Cd₁₀₀ caused some increase in activity of photosystem 2, PS2 (H₂O DCPIP), while distinct inhibition was observed with Cd₁₀₀₀. We also observed a similar effect when measuring oxygen evolution (H₂O PBQ + FeCy). Chloroplasts of spinach (Spinacia oleracea L.) were incubated for 30 min with 0, 15, 30, and 60 mM CdCl₂ (= Cd₀, Cd₁₅, Cd₃₀, and Cd₆₀). All concentrations studied inhibited the PS2 activity, the effect being stronger with increasing concentration of Cd²⁺. The photosynthetic oxygen evolution activity was also influenced most distinctly by the highest concentration employed, i.e. Cd₆₀. Electrophoretic analysis of the protein composition of cyanobacterium TM showed chief changes in the molecular mass regions of M_r 29 000 and 116 000, while with spinach chloroplasts the most distinct differences were observed in the regions of M_r 15 000 and 50 000. Ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) activity in cyanobacterial spheroplasts still remained on the 40 % level in the case of Cd₁₀₀₀, but it decreased down to approx. 2.5 % in the Cd₆₀ sample of spinach chloroplasts.     

Chloroplastic Carbon Dioxide Concentration in Norway Spruce (Picea Abies [L.] Karst.) needles relates to the position within the crown

Differences between sun (E) and shaded (S) foliage were studied in a Norway spruce (Picea abies [L.] Karst.) stand. Response curves describing the dependence of the CO2 assimilation rate (PN) on the CO2 concentration at the catalytic site of ribulose-1,5-bisphosphate carboxylase/oxygenase, RuBPCO (PN-Cc) were estimated using the simultaneous measurements of chlorophyll fluorescence and leaf gas exchange. Higher PN, higher electron transport (Ja), higher carboxylation capacity (Vc), and higher RuBPCO activity () for sun acclimated needles was found. The S-needles had higher portion of internal limitation and higher CO2 compensation concentration () than the E-needles. Because higher degree of limitation of photosynthesis by carboxylation was ascertained, it can be assumed that photosynthesis in shade foliage is limited mainly by lower carboxylation capacity and by low chloroplastic CO2 concentration     

Photosynthetic acclimation to elevated CO2 in wheat cultivars

Wheat (T. aestivum) cvs. Kalyansona and Kundan grown under atmospheric (CA) and elevated CO₂ concentrations (650±50 cm³ m^-3 - CE) in open top chambers were examined for net photosynthetic rate (PN), stomatal limitation (l _s) of P _N, ribulose-1,5-bisphosphate carboxylase (RuBPC) activity, and saccharide content of the leaves. The P _N values of both CA- and CE-grown plants compared at the same CO₂ concentration showed a down regulation under CE at the post-anthesis stage. The negative acclimation of P _N appeared to be due to both stomatal and mesophyll components, and the RuBPC activity got also adjusted. There was a decrease in activation state of RuBPC under CE. In connection with this, an increased accumulation of saccharides in wheat leaf under CE was observed. Kalyansona, owing to its larger sink potential in terms of the number of grains, showed a greater enhancement under CE in both post-ear emergence dry matter production and grain yield. Under CE, this cultivar also showed a lower down regulation of P _N than Kundan. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of water deficit on photosynthetic and other physiological responses in grapevine (Vitis vinifera L. cv. Riesling) plants

The grapevine (Vitis vinifera L. cv. Riesling) plants subjected to water deficit were studied for changes in relative water content (RWC), leaf dry mass, contents of chlorophyll (Chl), total leaf proteins, free amino acids, and proline, and activities of ribulose-1,5-bisphosphate carboxylase (RuBPC), nitrate reductase (NR), and protease. In water-stressed plants RWC, leaf dry matter, Chl content, net photosynthetic rate (P _N), and RuBPC and NR activities were significantly decreased. The total leaf protein content also declined with increase in the accumulation of free amino acids. Concurrently, the protease activity in the tissues was also increased. A significant two-fold increase in proline content was recorded.     

Shade effect alters leaf pigments and photosynthetic responses in Norway spruce (Picea abies L.) grown under field conditions

The contents of chlorophyll (Chl) and carotenoids (Car) per fresh mass were lower in shade needles than in sun needles. Ribulose-1,5-bisphosphate carboxylase (RuBPC) activity and contents of soluble proteins were also significantly lower in shade needles. In isolated thylakoids, a marked lower rate of whole chain and photosystem (PS) 2 activities were observed in shade needles. Smaller lower rate of PS1 activity was also observed in shade needles. The artificial exogenous electron donors, diphenyl carbazide (DPC) and NH₂OH, significantly restored the loss of PS2 activity in shade needles. Similar results were obtained when F_v/F_m was evaluated by Chl fluorescence measurements. The marked lower rate of PS2 activity in shade needles was due to the lower contents of 47, 33, 28–25, 23, and 17 kDa polypeptides. This conclusion was confirmed by immunological studies showing that the content of the 33 kDa protein of the watersplitting complex was diminished significantly in shade needles.     

Photosynthetic characteristics of C4 trait in chlorina mutant of rice (Oryza sativa L.)

Biao 810S is a chlorina mutant of the thermosensitive genic male sterile (TGMS) rice. We compared photosynthetic characteristics of these two lines. The contents of chlorophylls and carotenoids in Biao 810S were approximately half of those in 810S. However, the net photosynthetic rate (P _N) of Biao 810S was higher than that of 810S under high irradiance or low concentration of carbon dioxide, and the photon quantum efficiency was higher than that of 810S. The activity of ribulose-1,5-bisphosphate carboxylase/oxygenase in Biao 810S was only 69.80 % of that in 810S, but the activities of phosphoenolpyruvate carboxylase and NADP-malic enzyme were 79.50 and 69.06 % higher than those of 810S, respectively, suggesting that the efficiency of photon energy utilization in Biao 810S was enhanced by reduction of thermal dissipation and increase of electron transfer rate to generate sufficient assimilation power for the dark reactions. Consequently, the increased activities of C₄ photosynthetic enzymes lead to more effective fixation of CO₂ and the synergistic effect of light and dark reactions contributed to the higher P _N of Biao 810S.     

Plant Responses to Drought,Acclimation, and Stress Tolerance

At the whole plant level, the effect of stress is usually perceived as a decrease in photosynthesis and growth. That is why this review is focused mainly on the effect of drought on photosynthesis, its injury, and mechanisms of adaptation. The analysed literature shows that plants have evolved a number of adaptive mechanisms that allow the photochemical and biochemical systems to cope with negative changes in environment, including increased water deficit. In addition, the acquisition of tolerance to drought includes both phenotypic and genotypic changes. The approaches were made to identify those metabolic steps that are most sensitive to drought. Some studies also examined the mechanisms controlling gene expression and putative regulatory pathways.     

Changes of Photosynthetic Characteristics in Relation to Leaf Senescence in Two Maize Hybrids with Different Senescent Appearance

A field experiment was conducted to investigate the changes in chlorophyll (Chl) and nitrogen (N) contents, ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) and phosphoenolpyruvate carboxylase (PEPC) contents and PEPC activity, and the photon-saturated net photosynthetic rate (P _Nsat), and their relationships with leaf senescence in two maize hybrids with different senescent appearance. One stay-green (cv. P3845) and one earlier senescent (cv. Hokkou 55) hybrid were used in this study, and we found that Chl and N contents and the P _Nsat in individual leaves of P3845 were greater than those in corresponding leaves of Hokkou 55 at the successive growth stages. In addition, larger contents of RuBPCO and PEPC, and a greater activity of PEPC were observed in P3845. Due to the lower rates of decrease of Chl, RuBPCO, and PEPC amounts per unit of N, and the lower net C translocation rate per unit of N in the stay-green hybrid, leaf senescence was delayed in comparison to the earlier senescent hybrid.     

Developmental and Photosynthetic Characteristics of a Photoautotrophic Chrysanthemum Culture

Chrysanthemum plantlets were cultivated in vitro on media with 2.0, 0.3, or 0 % sucrose, or photoautotrophically without an organic carbon source but with supplementation of the culture vessel atmosphere with 2 % CO₂. The photoautotrophically cultivated plantlets showed a better growth and multiplication, higher contents of chlorophyll (Chl) and carotenoids, higher Chl a/b ratio, net photosynthetic rate and ribulose-1,5-bisphosphate carboxylase/oxygenase and phosphoenolpyruvate carboxylase activities than plantlets grown on the medium with sucrose. This revised version was published online in June 2006 with corrections to the Cover Date.     

Responses of Two Olive Tree (Olea Europaea L.) Cultivars to Elevated CO2 Concentration in the Field

Five-year-old plants of two olive cultivars (Frantoio and Moraiolo) grown in large pots were exposed for 7 to 8 months to ambient (AC) or elevated (EC) CO₂ concentration in a free-air CO₂ enrichment (FACE) facility. Exposure to EC enhanced net photosynthetic rate (P _N) and decreased stomatal conductance, leading to greater instantaneous transpiration efficiency. Stomata density also decreased under EC, while the ratio of intercellular (C _i) to atmospheric CO₂ concentration and chlorophyll content did not differ, except for the cv. Moraiolo after seven months of exposure to EC. Analysis of the relationship between photosynthesis and C _i indicated no significant change in carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase after five months of exposure to EC. Based on estimates derived from the P _N-C _i relationship, there were no apparent treatment differences in daytime respiration, CO₂ compensation concentration, CO₂-saturated photosynthetic rate, or photosynthetic rate at the mean C _i, but there was a reduction in stomata limitation to P _N at EC. Thus 5-year-old olive trees did not exhibit down regulation of leaf-level photosynthesis in their response to EC, though some indication of adjustment was evident for the cv. Frantoio with respect to the cv. Moraiolo.     

Characterization of Photosynthetic Apparatus of Pea Chlorophyll Mutants and their Heterotic F1 Hybrids with Standard Genotype (cv. Torsdag)

In comparison to cv. Torsdag, in leaves of low-productive Pisum sativum L. chlorophyll mutants the decrease in chlorophyll content was caused by the decrease in cell number per unit volume. Qualitative changes in activities of photosystem (PS) 1 in mutant M2004, and quantitative changes of PS1 and PS2 in mutants M2004 and M2014 and in hybrids were also found. However, the activity of ribulose-1,5-bisphosphate carboxylase (RuBPC) in M2014, and those of RuBPC and glyceraldehyde phosphate dehydrogenase in M2004 and hybrids were higher than in cv. Torsdag. The hybrids inherited the normal structure of photosynthetic apparatus of standard genotype in parallel with the compensatory gene complex of M2004, which was expressed at many levels of organization. This may be the basis of hybrid vigour in this case.     

Photosynthesis in leaves,fruits, stem and petioles of greenhouse-grown tomato plants

Gross photosynthetic capacity (P _G ) of greenhouse-grown tomato plants (Lycopersicon esculentum Mill.) decreased as the leaf aged. The P _G of the 10th, 15th and 18th leaves from the top was only 76, 37, and 18 % of P _G of the 5th leaf, respectively. Quantum yield (Y _Q ) and dark respiration rate (R _D ) were also lower in older leaves than in the younger ones. Net photosynthetic rate (P _G ) was apparent in young fruits (about 10 g FM) or young petioles but no P _N was found in large fruits (40 g or more FM) and stems because of high R _D . Both P _G and R _D were lower in older fruits and petioles or in lower parts of the stem compared to the younger ones or upper parts of stem. A sharp decrease in chlorophyll (Chl) content was only measured in the senescing 18th leaf. The Chl content in petioles, stems and fruits was proportional to P _G . Decreases in P _G of older leaves were attributed to decreases in content rather than activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) since soluble protein content was lower in older leaves than in the younger ones but the specific activity (activity per unit of protein) of RuBPCO was not so. The estimated values of P _N of the 10th, 15th and 18th leaves inside the canopy were only 50, 21, and 7 % of that in the 5th leaf. Therefore, leaves below the 18th can be removed in order to ensure a good air circulation and prevent diseases. The significance of photosynthesis in fruit, stem and petioles is not negligible because photosynthesis re-fixes the respired CO₂. This revised version was published online in September 2006 with corrections to the Cover Date.     

Partial decline of Arachis hypogaea L. photosynthesis triggered by drought stress

Photosynthetic capacity (PC) of three peanut cultivars (Arachis hypogaea L. cvs. 57-422, 73-30, and GC 8-35) decreased during drought stress (decline in relative water content from ca. 95 to 70 %) and recovered two days after rewatering. Mild water stress was not limiting for the total ribulose-1,5-bisphosphate carboxylase/oxygenase activity, since this enzyme activity increased under drought. Photosystem (PS) 2 and PS1 (the latter only in cv. GC 8-35) electron transport activities decreased under drought. The ratio of the variable to maximal chlorophyll fluorescence (Fv/Fm) decreased mainly in the cv. GC 8-35. All cultivars showed decreases in photochemical quenching (qP) and quantum yield of PS2 electron transport (Φe). Increase of basal fluorescence (F0) was observed in the cvs. 73-30 and GC 8-35, while the cv 57-422 showed a decrease. After rewatering a sharp increase was observed in the majority of the parameters. Thus under the present stress conditions, the cv GC 8-35 was the most affected for all the parameters under study. The cv. 57-422 showed a higher degree of tolerance being gradually affected in photosynthetic capacity (PC) in contrast to the two other cvs. which showed a sharp decrease in PC at the beginning of the drought cycle. This revised version was published online in September 2006 with corrections to the Cover Date.     

Composition and Characteristic Differences in Photosynthetic Membranes of Two Ecotypes of Reed (Phragmites communis L.) from Different Habitats

As compared with the swamp reed (SR) ecotype of Phragmites communis growing in the desert region of northwest China, plants of the dune reed (DR) ecotype from the same region possessed lower chlorophyll (Chl) content in leaves, and less thylakoids and grana stacks in chloroplasts. Tube gel electrophoresis without stain showed that the contents of Chl-protein (Pro) components related to photosystem 2 (PS2) were markedly lower in the DR thylakoid membranes than in the SR thylakoid membranes, while the contents of Chl-Pro components associated with PS1 were almost the same in both types. SDS-PAGE analysis indicated that the content of polypeptides of the light-harvesting Chl a/b complex of PS2 (LHC2) was lower in the DR thylakoids. Besides, the conformation of LHC2 within the DR thylakoid membranes was also altered as indicated by circular dichroism spectra. Hence in the DR, reduced energy harvesting by declining the size of LHC2 might be responsible for the down-regulated PS2 activity. Chl fluorescence parameters. F_v/F_m and quantum efficiency of PS2 (_PS2), were lower in the DR leaves than in the SR ones. However, non-photochemical quenching coefficient (q_N) was greater in DR than that in SR, implying other energy dissipation way exists in the DR photosynthetic membranes.     

Diurnal and Seasonal Changes in Photosynthetic Characteristics in Different Olive (Olea europaea L.) Cultivars

Diurnal and seasonal changes in photosynthetic characteristics, leaf area dry mass (ADM), and reducing sugar and total chlorophyll (Chl) contents of leaves of Frantoio, Leccino, and Maurino olive cultivars were investigated in Central Italy. Leaf net photosynthetic rate (P _N) per unit leaf area changed during the growing season and during the day, but the cultivar did not significantly influence the changes. In both young and one-year-old leaves the highest P _N values were observed in October, while the lowest values were recorded in August and December; during the day the highest P _N values were generally found in the morning. The pattern of photosynthetic response to photosynthetic photon flux density (PPFD) of leaves was similar in the three genotypes. Sub-stomatal CO₂ concentration (C _I) tended to increase when P _N decreased. The increase in C _I was accompanied by a stomatal conductance to water vapor (g _S) decrease. In general, P _N and dark respiration rate (R _D) were correlated. Transpiration rate (E), with no differences between the cultivars, increased from April to July, decreased greatly in August, then increased in October and finally decreased again in December. Leaf water content increased from April to June, remained high until mid July, decreased significantly in August, remaining constant until December with no differences associated with the cultivar. In both young and one-year-old leaves, the leaf water content per unit leaf area was slightly greater in Frantoio than in the other two cultivars. The one-year-old leaves had a higher Chl content than the young ones. The cultivar did not substantially influence the leaf reducing sugar content which decreased from April to August, when it reached the lowest level, then increased rapidly until October. During the day the reducing sugar content did not change significantly. The leaf ADM was slightly higher in Frantoio than in the other cultivars and one-year-old leaves had higher values than the young ones. Leaf ADM decreased from April to June and then tended to increase until December. During the day there were no substantial variations.