Presence of chloroplasts in mestome sheath cells of the C3 Pooid grass Elymus tsukushiensis

ABSTRACT

The successful introduction of the C₄ pathway into C₃ crops would increase photosynthetic rates and crop productivity. However, our poor understanding of how Kranz leaf anatomy develops poses a great obstacle. In particular, the origin, development, and genetics of bundle sheath (BS) cells in C₄ plants are key points to elucidate. Here we report that Elymus tsukushiensis, a common C₃ grass of the subfamily Pooideae, contains chloroplasts in the mestome sheath (MS) cells of the leaf, unlike most MS cells of C₃ grasses. The chloroplasts are smaller than those of mesophyll cells. Immunogold localization showed that the chloroplasts and mitochondria of MS cells, respectively, accumulate ribulose 1,5-bisphosphate carboxylase/oxygenase and a photorespiratory enzyme, glycine decarboxylase, as in mesophyll cells. Thus, we suggest that the MS cells have weak photosynthetic and photorespiratory functions. This finding provides an insight into the development and evolution of C₄-type BS cells in leaves of C₃ grasses.     

Variations in structural,biochemical, and physiological traits of photosynthesis and resource use efficiency in Amaranthus species (NAD-ME-type C4)

C₄ plants show higher photosynthetic capacity and productivity than C₃ plants owing to a CO₂-concentrating mechanism in leaves, which reduces photorespiration. However, which traits regulate the photosynthetic capacity of C₄ plants remains unclear. We investigated structural, biochemical, and physiological traits associated with photosynthesis and resource use efficiency in 20 accessions of 12 species of Amaranthus, NAD-malic enzyme-type C₄ dicots. Net photosynthetic rate (P_N) ranged from 19.7 to 40.5 μmol m^?2 s^?1. P_N was positively correlated with stomatal conductance and nitrogen and chlorophyll contents of leaves and was weakly positively correlated with specific leaf weight. P_N was also positively correlated with the activity of the C₃ enzyme ribulose-1,5-bisphoshate carboxylase/oxygenase, but not with the activities of the C₄ enzymes phosphoenolpyruvate carboxylase and NAD-malic enzyme. Structural traits of leaves (stomatal density, guard cell length, leaf thickness, interveinal distance, sizes of mesophyll and bundle sheath cells and the area ratio between these cells) were not significantly correlated with P_N. These data suggest that some of the biochemical and physiological traits are involved in interspecific P_N variation, whereas structural traits are not directly involved. Photosynthetic nitrogen use efficiency ranged between 260 and 458 μmol mol^?1 N s^?1. Photosynthetic water use efficiency ranged between 5.6 and 10.4 mmol mol^?1. When these data were compared with previously published data of C₄ grasses, it is suggested that common mechanisms may determine the variations in resource use efficiency in grasses and this dicot group.     

Variations in physiological,biochemical, and structural traits of photosynthesis and resource use efficiency in maize and teosintes (NADP-ME-type C4)

C₄ plants show higher photosynthetic capacity and resource use efficiency than C₃ plants. However, the genetic variations of these traits and their regulatory factors in C₄ plants still remain to be resolved. We investigated physiological, biochemical, and structural traits involved in photosynthesis and photosynthetic water and nitrogen use efficiencies (PWUE and PNUE) in 22 maize lines and four teosinte lines from various regions of the world. Net photosynthetic rate (P_N) ranged from 32.1 to 46.5 μmol m^?2 s^?1. P_N was positively correlated with stomatal conductance, transpiration rate, and chlorophyll, nitrogen and soluble protein contents of leaves, but not with specific leaf weight. P_N was positively correlated with the activities of ribulose-1,5-bisphosphate carboxylase/oxygenase and the C₄-acid decarboxylases, NADP-malic enzyme and phosphoenolpyruvate carboxykinase, but not with the activity of phosphoenolpyruvate carboxylase. Leaf structural traits (stomatal parameters, leaf thickness, and interveinal distance) were not correlated with P_N. These data suggest that physiological and biochemical traits are involved in the genetic variation of P_N, but structural traits are not directly involved. PWUE is in the lower class of values reported for C₄ plants, whereas PNUE is in the highest class of values reported for C₄ plants. PNUE was negatively correlated with leaf nitrogen content but not significantly correlated with P_N. PWUE was not correlated with δ¹³C values of leaves, indicating difficulty in using δ¹³C values as an indicator of PWUE of maize. In general, teosinte lines showed lower P_N but higher PWUE than maize lines.     

Photosynthetic Characteristics of an Amphibious C4 Plant,Eleocharis retroflexa ssp. chaetaria

Abstract

Eleocharis retroflexa (Poir.) Urban ssp. chaetaria (Roem. & Schult.) T. Koyama, an amphibious leafless sedge, grows not only under terrestrial conditions but also under completely submerged aquatic conditions. We investigated the photosynthetic traits and structural features of the culms, which are the photosynthetic organs, in the terrestrial and submerged forms of this species and compared them with those of other amphibious species of Eleocharis which are known to change the photosynthetic modes. The culms of the terrestrial form had Kranz anatomy with well-developed Kranz (bundle sheath) cells and high levels of C₄ enzyme activity typical of the NAD-malic enzyme (NAD-ME) subtype of C₄ metabolism. They also had a δ ¹³C value typical of C₄ plants, indicating that the terrestrial form fixes carbon through the C₄ pathway. The culms of the submerged form had not only a Kranz-like anatomy but also revealed anatomical traits typical of leaves of submerged aquatic plants. The activities of the C₄ enzymes in the submerged form were lower than those in the terrestrial form, but were still in the range typical of G4 plants, ¹⁴C pulse-¹²C chase experiments with the submerged form indicated that almost all of the fixed ¹⁴C was incorporated into G4 compounds, and subsequently the raioactivity was transferred into C₃ compounds and sucrose. The submerged form showed no diurnal fluctuation in malate level. These data demonstrate that a C₄ metabolism is operative even in the submerged form. This unique amphibious C₄ plant provides an intriguing example of the physiological and ecological adaptability of C₄plants.     

Comparison of Susceptibility to Photoinhibition and Energy Partitioning of Absorbed Light in Photosystem II in Flag Leaves of Two Rice (Oryza sativa L.) Cultivars that Differ in Their Responses to Nitrogen-Deficiency

Abstract

Abstract: The energy partitioning in photosystem II (PSII) and the susceptibility to photoinhibition in PSII were investigated in flag leaves of two rice cultivars, Shirobeniya (a traditionaljaponica) and Akenohoshi (an improved japonica-indica intermediate) grown under standard-nitrogen (N) (SN) and low-N (LN) conditions. N-deficiency resulted in significant decreases in total dry weight, net photosynthetic rate (P_n), the energy flux via carboxylation (J_c), and content of ribulose-1,5-bisphosphate carboxylase /oxygenase (Rubisco) in flag leaves in the two cultivars, and these parameters of Shirobeniya were lower than those in Akenohoshi under the LN condition. In the two cultivars, the energy flux via alternative electron flow was significantly increased by N-deficiency, which was accompanied by enhanced activity of superoxide dismutase (SOD). Although under the LN condition no cultivar differences were found in J_a and SOD, ascorbate peroxidase activity in Shirobeniya was lower than that in Akenohoshi. N-deficiency resulted in more significant increases in the susceptibility to photoinhibition (the degree of decrease in maximum quantum yield of PSII), hydrogen peroxide (H₂O₂) content and malondialdehyde content after exposure to high irradiance in Shirobeniya than those in Akenohoshi. These results indicated that the increased susceptibility to photoinhibition in the LN plants of Shirobeniya was mainly due to oxidative damages to chloroplasts, resulting from lower carboxylation and H₂O₂-scavenging capacities. Therefore, both carboxylation and H₂O₂-scavenging capacities could be important factors in determining the cultivar difference in the productivity of rice under LN conditions.     

Intracellular position of mitochondria and chloroplasts in bundle sheath and mesophyll cells of C3 grasses in relation to photorespiratory CO2 loss

In C₃ plants, photosynthetic efficiency is reduced by photorespiration. A part of CO₂ fixed during photosynthesis in chloroplasts is lost from mitochondria during photorespiration by decarboxylation of glycine by glycine decarboxylase (GDC). Thus, the intracellular position of mitochondria in photosynthetic cells is critical to the rate of photorespiratory CO₂ loss. We investigated the intracellular position of mitochondria in parenchyma sheath (PS) and mesophyll cells of 10 C₃ grasses from 3 subfamilies (Ehrhartoideae, Panicoideae, and Pooideae) by immunostaining for GDC and light and electron microscopic observation. Immunostaining suggested that many mitochondria were located in the inner half of PS cells and on the vacuole side of chloroplasts in mesophyll cells. Organelle quantification showed that 62–75% of PS mitochondria were located in the inner half of cells, and 62–78% of PS chloroplasts were in the outer half. In mesophyll cells, 61–92% of mitochondria were positioned on the vacuole side of chloroplasts and stromules. In PS cells, such location would reduce the loss of photorespiratory CO₂ by lengthening the path of CO₂ diffusion and allow more efficient fixation of CO₂ from intercellular spaces. In mesophyll cells, it would facilitate scavenging by chloroplasts of photorespiratory CO₂ released from mitochondria. Our data suggest that the PS cells of C₃ grasses have already acquired an initial structure leading to proto-Kranz and further C₃–C₄ intermediate anatomy. We also found that in the Pooideae, organelle positioning in PS cells on the phloem side resembles that in mesophyll cells.     

Screening of High kcat Rubisco among Poaceae for Improvement of Photosynthetic CO2 Assimilation in Rice

Abstract

The activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) is a major limitation of photosynthetic CO₂ assimilation in C₃ plants. In order to find useful Rubisco for improvement of photosynthesis in rice under elevated CO₂, we analyzed the catalytic turnover rate (k_cat) of Rubisco in Poaceae including C₃ alpine plants, C₃ cold-resistant plants and C₄ plants. Rubisco in these plants showed 1.1- to 2.8-fold higher k_cat than that in rice. However, the most of high k_cat Rubisco also showed a higher km for CO₂ (Kc) than that of rice, indicating that increase in k_cat led to decrease in the affinity for CO₂. Rubisco in Festuca ovina, Phleum pratense and Sorghum bicolor showed relatively high k_cat to Kc. Although the k_cat of Rubisco in F. ovina and P. pratense was not so high (1.5-1.6 fold relative to rice), the Kc was comparable to that in rice and the amino acid sequence of RbcL shared higher identity to that in rice than that in S. bicolor. By contrast, Rubisco of S. bicolor showed considerably high k_cat (2.5-fold relative to rice), which is considered to be the most important factor for improvement of photosynthesis. In our estimation, the expression of high k_cat Rubisco of F. ovina and S. bicolor in rice could significantly enhance CO₂ assimilation at Ci of 50 Pa, the level assumed to be reached by the middle of this century.     

Effect of nitrogen supply on leaf appearance,leaf growth,leaf nitrogen economy and photosynthetic capacity in maize (Zea mays L.)

Leaf area growth and nitrogen concentration per unit leaf area, N_a (g m⁻² N) are two options plants can use to adapt to nitrogen limitation. Previous work indicated that potato (Solanum tuberosum L.) adapts the size of leaves to maintain N_a and photosynthetic capacity per unit leaf area. This paper reports on the effect of N limitation on leaf area production and photosynthetic capacity in maize, a C4 cereal. Maize was grown in two experiments in pots in glasshouses with three (0.84–6.0 g N pot⁻¹) and five rates (0.5–6.0 g pot⁻¹) of N. Leaf tip and ligule appearance were monitored and final individual leaf area was determined. Changes with leaf age in leaf area, leaf N content and light-saturated photosynthetic capacity, P_max, were measured on two leaves per plant in each experiment. The final area of the largest leaf and total plant leaf area differed by 16 and 29% from the lowest to highest N supply, but leaf appearance rate and the duration of leaf expansion were unaffected. The N concentration of expanding leaves (N_a or %N in dry matter) differed by at least a factor 2 from the lowest to highest N supply. A hyperbolic function described the relation between P_max and N_a. The results confirm the ‘maize strategy’: leaf N content, photosynthetic capacity, and ultimately radiation use efficiency is more sensitive to nitrogen limitation than are leaf area expansion and light interception. The generality of the findings is discussed and it is suggested that at canopy level species showing the ‘potato strategy’ can be recognized from little effect of nitrogen supply on radiation use efficiency, while the reverse is true for species showing the ‘maize strategy’ for adaptation to N limitation.     

Production of Raphanus sativus (C3)-Moricandia arvensis (C3-C4 intermediate) Monosomic and Disomic Addition Lines with Each Parentl Cytoplasmic Background and their Photorespiratory Characteristics

Abstract

We are maintaining five Moricandia arvensis monosomic addition lines of Raphanus sativus carrying R. sativus cytoplasm (autoplasmic MALs) and twelve M. arvensis MALs of R. sativus carrying M. arvensis cytoplasm (alloplasmic MALs) from BC₆ to BC₈ generation, and newly produced five M. arvensis disomic addition lines of R. sativus (autoplasmic DALs) and seven M. arvensis DALs of R. sativus carrying M. arvensis cytoplasm (alloplasmic DALs) from selfing and sib-crossing of the MALs and DALs in S₃BC₅ and S₂BC₆ generations. The structural, biochemical and physiological characteristics related to photorespiration of these MALs and DALs were compared to study the genetic mechanisms of the C₃-C₄ intermediate photosynthesis in the individual chromosomes of M. arvensis. The CO₂ compensation point of the autoplasmic and alloplasmic DALs (RMa-b and MaR-b DALs) with one pair of M. arvensis ‘b’ chromosome were 29.4 and 30.1 μmol mol-¹, respectively, which were significantly lower than that of other DALs and MALs as well as R. sativus (34.5 𰂼mol mol-¹). An immunogold electron microscopic study of the P-protein of glycine decarboxylase (GDC) in photosynthetic cells of the RMa-b DAL revealed that the bundle sheath cell (BSC) mitochondria were more intensively labeled for the protein than the mesophyll cell (MC) mitochondria. The ratio of the labeling density of the BSC mitochondria to that of the MC mitochondria was 1.13, which lies between values in M. arvensis (2.66) and R. sativus (0.76). These data suggest that the ‘b’ chromosome of M. arvensis genome controls the expression of C₃-C₄ intermediate characteristics.     

Effects of Salinity Stress on the Structure of Bundle Sheath and Mesophyll Chloroplasts in NAD-Malic Enzyme and PCK Type C4 Plants

Abstract

The effect of NaCl stress on the structure of leaf chloroplasts was investigated in several NAD-Malic enzyme (NAD-ME) and phosphoenolpyruvate carboxykinase (PCK) type C₄ plant species. Seedlings of the monocot species, except Zoysia japonica, grown in 300 mL pots were subjected to salt stress by adding 50 mL of 3% NaCl solution per day to the soil for 5 d after the fourth leaf blades were fully developed. Z. japonica and the dicot species, Amaranthus tricolor, were also treated with 3% NaCl in a similar manner from 5 wk after germination. Salt stress negatively affected the growth, chlorophyll content and chloroplast structure in all the species. At the ultrastructure level, swelling of thylakoids and disruption of envelopes were more or less observed in mesophyll cell (MC) chloroplasts after salt treatment. The structure of bundle sheath cell (BSC) chloroplasts, on the other hand, was hardly damaged under salt condition although stromal and starch areas were considerably decreased. Furthermore, salinity induced granal development in BSC chloroplasts in most species; the number of thylakoids per granum, granal indices and appressed thylakoid density in salt-treated plants were generally higher than those in control. Since the similar responses have also been reported in all NADP-ME type C₄ species investigated in our previous study, the high sensitivity to salt stress in MC chloroplasts and the granal development in BSC chloroplasts by salinity were considered to be common phenomena in all three C₄ subtypes.     

高产水平下水肥耦合对小麦旗叶光合特性及产量的影响

为了探讨黄淮麦区冬小麦的高产机理,在大田条件下设置不同梯度的水肥耦合模式,分析了高产水平下旗叶光合特性与籽粒产量的效应.结果表明,不同模式的水肥耦合对旗叶光合特性和籽粒产量的影响不同,总体表现为土壤水分的处理效应大于施氮量的效应.旗叶光合速率(Pn)和气孔导度(G5)均随着土壤水分的增加而升高,表现为W3(充足水分)＞W2(适宜水分)＞W1(自然降水),差异达显著水平.胞间CO2浓度(Ci)随着土壤水分的增加而降低;不同的氮肥处理,旗叶Pn和Gs随着施氮量的增加而升高,表现为N3(270 kg·hm-2)＞N2(195 kg·hm-2)＞N1(CK),Ci在不同水分条件下与施氮量的关系不尽相同.不同水肥耦合模式对水分利用效率的影响表现为W2＞W3＞W1,并以W2N2为最高,不同处理组合的籽粒产量亦以W2N2为最高.表明高产条件下W2N2水肥组合应是高产高效的运筹模式.     

Ultrastructure of Fusion Product between Protoplasts from C3 and C4 Species of Amaranthaceae

Summary

This paper describes the ultrastructure of the electric field-induced fusion products of C₃ and C₄ species of Amaranthaceae at the early developmental stage. Protoplasts of C₃ species were isolated from a Ceiosia cristata L. cell suspension and, those of C₄ species were isolated from an Amaranthus tricolor L. cotyledon. Incompatibility occurred in the C₃/C₄ hybrid. The incompatibility reactions were detected in the newly formed hybrid cells accompanied with significant changes in the nucleolus (segregation of nucleolar components) and plastids (cup-like shape or amoeboid plastid enclosing cytoplasmic materials) of C₃ species parent. The structural changes in the organelles of the C₄ partner were less marked. After 5 days of culture, most organelles showed high cellular activity, and a normal dedifferentiation process of mesophyll chloroplasts was observed. At this stage nucleolar segregation was not detected and the C₃ species plastids were difficult to distinguish from the proplastids formed from mesophyll chloroplasts. In addition, some mitochondria showed bursting-like structure. However, under the culture condition used these somatic incompatibility did not seem to impair further growth of fusion products since they were still proliferating well resulting in callus formation.     

Culm sucrose accumulation promotes physiological decline of mature leaves in ripening sugarcane

Photosynthetic activity in C₄ sugarcane has been suggested to be regulated by the demand for photoassimilate from sink tissues, including culm storage of sucrose and other components of the plant (e.g. roots and shoot primordia). This study examined the extent to which sink-demand influences leaf photosynthetic activity and controls leaf turnover in field-grown sugarcane. To increase sink-demand on selected leaves, plants were defoliated apart from the immature leaf before the first fully expanded leaf (2nd leaf) and the mature 8th leaf. Changes in leaf gas exchange and fluorescence characteristics were recorded for both leaves over a 28 d period. Variations in leaf and culm sucrose and hexose concentrations and allocation of ¹⁴C-labelled photosynthate were also measured. A decrease in culm internodal sucrose concentrations in partially defoliated plants was associated with significant increases in assimilation (A) and electron transport rates (ETR) for both leaves 2 and 8. Conversely, accumulation of sucrose in the culms of control plants was related to a decline in photosynthetic rates in leaf 8 during the treatment period. Leaves 2 and 8 of defoliated plants (27 d) were characterised by an increase in partitioning of ¹⁴C to mature and immature internodes, respectively, compared to control plants. In addition, hexose levels in leaves of defoliated plants decreased significantly (36% and 48% decrease in leaves 2 and 8, respectively) compared to corresponding leaves of untreated controls over the duration of the experiment, indicating that the signaling mechanisms regulating the decline in leaf photosynthetic activity are likely hexose-mediated. It was concluded that leaf physiological ageing in sugarcane was promoted by sucrose accumulation during culm maturation as a consequence of decreased sink-demand for photosynthate.     

Correlation of Chlorophyll Meter Readings with Gas exchange and Chlorophyll Fluorescence in Flag Leaves of Rice (Oryza sativa L.) Plants

Abstract

The objective of this study was to establish the correlation of the chlorophyll meter (SPAD) readings with the contents of chlorophyll (Chl) and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), the gross photosynthetic rate (P_g), and the maximum quantum yield of photosystem II (PSII) (F_v/F_m) in flag leaves of rice (Oryza sativa L.) in ripening stage. The SPAD readings significantly correlated with the Chl content, the Rubisco content, P_g and F_v/F_m (R² = 0.848, 0.648, 0.671 and 0.712, respectively), which suggests that the SPAD meter has the potential to estimate the photosynthetic capacity of the flag leaves. However, both P_g and F_v/F_m had a stronger relationship with the Rubisco content than the SPAD readings, indicating that the PSII photochemical and CO₂ assimilation capacities are strongly influenced by the Rubisco content. Therefore, accurate calibration would be indispensable to obtain the physiological information from the SPAD readings of flag leaves.     

Inheritance of C3―C4 Intermediate Photosynthesis in Reciprocal Hybrids Between Moricandia Arvensis (C3―C4) and Brassica Oleracea (C3) That Differ in Their Genome Constitution

Abstract

To Elucidate The Genetic Mechanisms Underlying C₃―C₄ intermediate Photosynthesis, We investigated The Structural and Photosynthetic Characteristics of Leaves of Reciprocal Hybrids Between The C₃―C₄ intermediate Species Moricandia Arvensis (L.) Dc. (Mama) and The C₃ Species Brassica Oleracea L. (Cabbage; Cc), Which Differ in Genome Constitution. Moricandia Arvensis Bundle Sheath (Bs) Cells included Many Centripetally Located Chloroplasts and Mitochondria, Whereas Those of Cabbage Had Few Organelles. Hybrid Leaves Were Structurally intermediate Between Those of The Parents and Showed Stronger intermediate C₃―C₄ Features As The Proportion of The Ma Genome increased. The P-Protein of Glycine Decarboxylase (Gdc) Was Confined Mainly To Bs Mitochondria in M. Arvensis, But Accumulated More in The Mesophyll (M) of Cabbage. in The Hybrids, The Accumulation of Gdc in Bs Cells increased With An increasing Ma:C Ratio. Hybrids Exhibited Gradients in Structural and Biochemical Features, Even in Reciprocal Crosses. The Co₂ Compensation Point of Reciprocal Hybrids With High Ma:C Ratios Was Lower Than That of Cabbage But Higher Than That of M. Arvensis. Thus, The Structural and Biochemical Features in Hybrid Leaves Reduced Photorespiration. Moricandia Arvensis Had A Higher Photosynthetic Rate Than Cabbage, But The Photosynthetic Rates of Hybrids Were intermediate Between Those of The Parents Or Comparable To That of M. Arvensis. Our Results Demonstrate That The C₃―C₄ intermediate Characteristics Are inherited Based On The Ratio of The Parent Genomes, and That There Is No Evidence of Cytoplasmic inheritance in These Characteristics.     

Photosynthesis, symbiotic N and C accumulation in leaves of 30 nodulated cowpea genotypes grown in the field at Wa in the Guinea savanna of Ghana

Cowpea is a major food legume in Africa with relatively little information on N and C nutrition under field conditions in the continent. The aim of this study was to assess the relationship between leaf symbiotic N nutrition and photosynthetic carbon assimilation in cowpea genotypes grown under field conditions in the Guinea savanna of Ghana. The experiments were conducted in a randomized complete block design in Ghana, using 30 nodulated cowpea genotypes. Data collected from gas-exchange studies showed that genotypes with high photosynthetic rates expectedly exhibited greater stomatal conductance, high transpiration rates and increased water-use efficiency, leading to greater dry matter yield. In contrast, those with low photosynthetic rates revealed low stomatal conductance, low transpiration rates and low water-use efficiency. Quantification of symbiotic N in photosynthetic leaves indicated greater N₂ fixation in genotypes with higher photosynthetic rates, leaf transpiration and better water-use efficiency. Furthermore, measurement of C in leaves also indicated greater accumulation in genotypes with higher symbiotic N yield and total N, clearly indicating a link between C and N metabolism in photosynthetic leaves. Additionally, measures of photosynthetic N-use efficiency showed that genotypes with high levels of C-fixed per unit leaf total N also exhibited greater C-fixed per unit leaf N-fixed. Data suggest that where symbiotic legumes derive a large proportion of their N from N₂ fixation, photosynthetic C yield is much enhanced.     

玉米C4光合叶不同部位“花环”结构及叶绿素含量的变化

以玉米进行C4光合的全展第5位叶片为材料,分析从叶基部到顶部的解剖结构和叶绿素含量变化,研究玉米C4光合叶片"花环"结构随叶片发育的变化规律。结果表明,玉米第5位叶从基部到顶部都具有完整的典型"花环"结构,维管束鞘细胞(BSC)和叶肉细胞(MC)的体积在叶片发育过程中具有渐变性,从叶基部到顶部BSC和MC均呈先增大后变小的趋势,而且叶绿素a、b和a+b含量呈相同的变化趋势,说明BSC和MC细胞体积与叶绿素含量的变化具有相关性。叶绿素a/b总体呈上升趋势,说明玉米第5叶基部到顶部的光合途径存在C3向C4转变的过程。玉米第5叶不同部位C4光合途径发育的渐变性比前3叶更为明显。     

外源C4二羧酸对转玉米PEPC基因水稻C4光合途径的促进作用

以转PEPC基因水稻和野生型水稻Kitaake为材料, 研究了不同基因型水稻叶片中的C4光合微循环及其功能。通过测定与光合C4途径有关的关键酶,如磷酸烯醇式丙酮酸羧化酶(PEPC)、NADP苹果酸酶(NADP-ME)、NADP苹果酸脱氢酶(NADP-MDH)和丙酮酸磷酸双激酶(PPDK),说明野生型水稻叶片中具有完整的C4光合酶体系;用外源草酰乙酸(OAA)或苹果酸(MA)饲喂叶切片或叶绿体后明显增加光合速率,证明野生型水稻中具有一个有限的光合C4微循环。将玉米的PEPC基因导入原种水稻后,可大幅度提高光合C4微循环的速率。测定不同基因型的CO2交换速率,看出水稻中C4光合微循环的增强有提高净光合速率(Pn)和降低Pr/Pn比值的作用。叶绿素荧光特性分析表明,C4光合微循环的增强伴随着PSⅡ电子传递效率(Fv/Fm)和光化学猝灭(qp)的增加,以及非光化学猝灭(qN)的降低。      

Mechanism of High Photosynthetic Capacity in BC2F4 LinesDerived from a Cross between Oryza sativa and Wild Relatives O. rufipogon

Abstract

We found that several BC₂F₄ lines had high leaf photosynthetic rates under light-saturated and ambient CO₂ conditions. These lines are progenies of BC₂F₁ plants with high photosynthetic capacities which were generated by backcrossing between Oryza rufipogon (W630) and O. sativa cv. Nipponbare, as a recurrent parent. Some photosynthetic characteristics of the BC₂F₄ lines were investigated to identify the factors increasing photosynthetic rates. Photosynthetic rates of these lines under light-saturated conditions at 50 to 700 ppm CO₂ concentrations were higher than those in Nipponbare. The estimated-maximum photosynthetic rates under light-saturated and CO₂-saturated conditions in BC₂F₄ lines were also higher than that in Nipponbare. The photosynthetic rate under light-saturated and ambient CO₂ conditions was positively correlated with the carboxylation efficiency as an indicator of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) activity in vivo rather than stomatal conductance. Initial and total Rubisco activities in vitro tended to be higher in the BC₂F₄ lines than in Nipponbare. The content of active Rubisco calculated from the activation state of Rubisco was also higher in the BC₂F₄ lines than in Nipponbare. These results suggest that high photosynthetic capacities of BC₂F₁ plants can be maintained high in their progenies and high photosynthetic rates under light-saturated and ambient CO₂ conditions in the BC₂F₄ lines are achieved mainly by the high activity of Rubisco due to the high active Rubisco content.     

Differential Sensitivity of Chloroplasts in Mesophyll and Bundle Sheath Cells in Maize,an NADP-Malic Enzyme-Type C4 Plant,to Salinity Stress

Abstract

The changes in chloroplast ultrastructure and the contents of chlorophyll, Na and K in response to salinity stress were investigated in leaves of maize, an NADP-malic enzyme-type C₄ plant species possessing dimorphic chloroplasts. The seedlings were treated with 0, 1, 2 or 3% NaCl for three or five days under a light or dark condition. In both light and dark conditions, the dry weight of salt-treated plants decreased as NaCl concentration increased. Chlorophyll and K contents of the second leaf blade decreased as NaCl concentration increased under the light condition but not under the dark condition. Na content of the second leaf blade was significantly higher at high NaCl concentrations under both light and dark conditions. However, Na content was much lower under the dark condition than light condition. Higher concentrations (2 and 3%) of NaCl significantly increased the size of plastoglobules, decreased the number and size of starch granules and altered the chloroplast ultrastructure. Under the light condition, mesophyll cell (MC) chloroplasts appeared more sensitive to the damaging effect of salinity than the bundle sheath cell (BSC) chloroplasts. MC chloroplasts became more globular in shape and showed swollen and disorganized thylakoids and reduced thickness of grana by salinity. BSC chloroplasts were less affected by salinity than MC chloroplasts. Although chloroplast size and number and size of starch granules were reduced, there was no structural distortion in the thylakoids of BSC chloroplasts. However, the thickness of grana was increased by salinity. Under the dark condition, the chloroplast structure was less affected by salinity. Though the envelope of BSC chloroplasts was occasionally damaged, the thylakoids in both MC and BSC chloroplasts were preserved under salinity stress. The present study suggests that the chloroplast damage caused by salinity is light-dependent and MC chloroplasts are more sensitive to salinity than BSC chloroplasts.