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1.
Sulfate assimilation and glutathione synthesis were traditionally believed to be differentially compartmentalised in C4 plants with the synthesis of cysteine and glutathione restricted to bundle sheath and mesophyll cells, respectively. Recent studies, however, showed that although ATP sulfurylase and adenosine 5′ phosphosulfate reductase, the key enzymes of sulfate assimilation, are localised exclusively in bundle sheath in maize and other C4 monocot species, this is not true for the dicot C4 species of Flaveria. On the other hand, enzymes of glutathione biosynthesis were demonstrated to be active in both types of maize cells. Therefore, in this review the recent findings on compartmentation of sulfate assimilation and glutathione metabolism in C4 plants will be summarised and the consequences for our understanding of sulfate metabolism and C4 photosynthesis will be discussed.  相似文献   

2.
R.Z. Wang 《Photosynthetica》2005,43(4):535-549
Of the total 570 species, 194 species in 116 genera and 52 families were found with C3 photosynthesis, 24 species in 17 genera and 6 families with C4 photosynthesis, and 2 species in 1 genera and 1 family with CAM photosynthesis. 90 % of the total species can be found in Changbai Mountain flora, more a half (69 %) in North China flora, and about 1/3 in Mongolian flora and Xinan flora, respectively. The occurrence of C4 species was not as common as that in adjacent grasslands and deserts, but relatively more than in the adjacent forests. Of the total 24 C4 species, 63 % C4 species (15 of 24) was found in Gramineae. Nine life form types can be found, reflecting the moist climate in the region, especially the occurrence of epiphyte and liana forms. Relatively more geophyte life form plants suggested the winter in the region was much colder than in grasslands. These indicated that both ecological studies and land management decisions must take into account plant photosynthetic pathway and life form patterns, for both of them are closely related to climatic changes and land use.  相似文献   

3.
Ogle K 《Oecologia》2003,136(4):532-542
The distance between veins has the potential to affect photosynthesis in C4 grasses because photon capture and photosynthetic carbon reduction are primarily restricted to vascular bundle sheath cells (BSC). For example, BSC density should increase as interveinal distance (IVD) decreases, and thus IVD may influence photon capture and photosynthesis in C4 grasses. The objective of this study is to evaluate the potential importance of IVD to the function of C4 grasses, and a literature survey is conducted to test the hypothesis that quantum yield of photosynthesis () increases with decreasing IVD. First, a meta-analysis of and IVD values obtained for 12 C4 grass species supports this hypothesis as and IVD are significantly negatively correlated (r=–0.61). Second, a regression of carbon isotope discrimination () versus IVD was conducted and the regression equation was used in a simple biochemical model that relates to and leakage of CO2 from the BSC. The modeling analysis also supports the hypothesis that decreases with increasing IVD in C4 grasses. C4 grasses are virtually absent from shaded habitats, and the biochemical model is employed to examine the implications of IVD for shade-tolerance in C4 grasses. The model predicts that only those species with uncommonly small IVD values are able to tolerate prolonged shade.  相似文献   

4.
Potato (Solanum tuberosum L. cv. Désirée) plants with an antisense reduction in the P-protein of the glycine decarboxylase complex (GDC) were used to study the interaction between respiration and photorespiration. Mitochondria isolated from transgenic plants had a decreased capacity for glycine oxidation and glycine accumulated in the leaves. Malate consumption increased in leaves of GDC deficient plants and the capacity for malate and NADH oxidation increased in isolated mitochondria. A lower level of alternative oxidase protein and decreased partitioning of electrons to the alternative pathway was found in these plants. The adenylate status was altered in protoplasts from transgenic plants, most notably the chloroplastic ATP/ADP ratio increased. The lower capacity for photorespiration in leaves of GDC deficient plants was compensated for by increased respiratory decarboxylations in the light. This is interpreted as a decreased light suppression of the tricarboxylic acid cycle in GDC deficient plants in comparison to wild-type plants. The results support the view that respiratory decarboxylations in the light are restricted at the level of the pyruvate dehydrogenase complex and/or isocitrate dehydrogenase and that this effect is likely to be mediated by mitochondrial photorespiratory products.  相似文献   

5.
Barbehenn RV  Karowe DN  Chen Z 《Oecologia》2004,140(1):96-103
The increasing CO2 concentration in Earths atmosphere is expected to cause a greater decline in the nutritional quality of C3 than C4 plants. As a compensatory response, herbivorous insects may increase their feeding disproportionately on C3 plants. These hypotheses were tested by growing the grasses Lolium multiflorum C3) and Bouteloua curtipendula C4) at ambient (370 ppm) and elevated (740 ppm) CO2 levels in open top chambers in the field, and comparing the growth and digestive efficiencies of the generalist grasshopper Melanoplus sanguinipes on each of the four plant × CO2 treatment combinations. As expected, the nutritional quality of the C3 grass declined to a greater extent than did that of the C4 grass at elevated CO2; protein levels declined in the C3 grass, while levels of carbohydrates (sugar, fructan and starch) increased. However, M. sanguinipes did not significantly increase its consumption rate to compensate for the lower nutritional quality of the C3 grass grown under elevated CO2. Instead, these grasshoppers appear to use post-ingestive mechanisms to maintain their growth rates on the C3 grass under elevated CO2. Consumption rates of the C3 and C4 grasses were also similar, demonstrating a lack of compensatory feeding on the C4 grass. We also examined the relative efficiencies of nutrient utilization from a C3 and C4 grass by M. sanguinipes to test the basis for the C4 plant avoidance hypothesis. Contrary to this hypothesis, neither protein nor sugar was digested with a lower efficiency from the C4 grass than from the C3 grass. A novel finding of this study is that fructan, a potentially large carbohydrate source in C3 grasses, is utilized by grasshoppers. Based on the higher nutrient levels in the C3 grass and the better growth performance of M. sanguinipes on this grass at both CO2 levels, we conclude that C3 grasses are likely to remain better host plants than C4 grasses in future CO2 conditions.  相似文献   

6.
We combined measurements of short-term (during gas exchange) and long-term (from plant dry matter) carbon isotope discrimination to estimate CO2 leakiness from bundle sheath cells in six C4 species (three grasses and three dicots) as a function of leaf insertion level, growth temperature and short-term irradiance. The two methods for determining leakiness yielded similar results (P > 0.05) for all species except Setaria macrostachya, which may be explained by the leaf of this species not being accommodating to gas exchange. Leaf insertion level had no effect on leakiness. At the highest growth temperature (36°C) leakiness was lower than at the two lower growth temperatures (16°C and 26°C), between which no differences in leakiness were apparent. Higher irradiance decreased leakiness in three species, while it had no significant effect on the others (there was an opposite trend in two species). The inverse response to increasing irradiance was most marked in the two NAD-ME dicots (both Amaranthus species), which both showed almost 50% leakiness at low light (300 μmol quanta m−2 s−1) compared to about 30% at high light (1,600 μmol quanta m−2 s−1). NADP-ME subtype grasses had lower leakiness than NAD-ME dicots. Although there were exceptions, particularly in the effect of irradiance on leakiness in Sorghum and Boerhavia, we conclude that conditions favourable to C4 photosynthesis (high temperature and high light) lead to a reduction in leakiness.  相似文献   

7.
Kubien DS  Sage RF 《Oecologia》2003,137(3):330-337
C4 plants are rare in cool climates, an ecological pattern attributable to their poor photosynthetic performance at low temperatures relative to C3 species. However, some C4 species are able to persist at high latitudes and high elevations, possibly due to the characteristics of the particular microsites they inhabit in these otherwise unfavourable environments. One such species is Muhlenbergia glomerata, which occurs above 60°N in Canada and is found in the atypical C4 habitat of boreal fens. In this study, we evaluate how microsite features affect the success of M. glomerata in boreal fens. We surveyed 19 populations across northern Ontario during the summers of 1999 and 2000. The ground coverage by woody vegetation was the most important parameter affecting the presence or absence of M. glomerata. Woody plants covered over 50% of the ground area in plots where M. glomerata is absent, but less than 20% where it is present. The minimum light intensity threshold for the presence of the C4 species was about 32% of full-sunlight at plant height. Surprisingly, in boreal fens M. glomerata was largely restricted to the wetter moss hollows, rather than occurring on the dry hummocks where its greater water use efficiency might have been advantageous. Woody species dominated the hummocks, but were uncommon in the hollows. In these cool northern climates M. glomerata apparently persists because sufficient periods of temperatures favourable to C4 photosynthesis occur, but this persistence likely requires some factor that suppresses the woody vegetation.  相似文献   

8.
The amphibious leafless sedge Eleocharis retroflexa ssp. chaetaria expresses C4-like biochemical characteristics in both the terrestrial and submerged forms. Culms of the terrestrial form have Kranz anatomy, whereas those of the submerged form have Kranz-like anatomy combined with anatomical features of aquatic plant leaves. We examined the immunolocalization of C3 and C4 enzymes in culms of the two forms. In both forms, phosphoenolpyruvate carboxylase; pyruvate, Pi dikinase; and NAD-malic enzyme were compartmentalized between the mesophyll (M) and Kranz cells, but their levels were somewhat reduced in the submerged form. In the terrestrial form, ribulose-1,5-bisphosphate carboxylase/oxygenase (rubisco) occurred mainly in the Kranz cells, and weakly in the M chloroplasts. In the submerged form, the rubisco occurred at higher levels in the M cells than in the terrestrial form. In both forms, the C4 pattern of enzyme expression was clearer in the M cells adjacent to Kranz cells than in distant M cells. During the transition from terrestrial to submerged conditions, the enzyme expression pattern changed in submerged mature culms that had been formed in air before submergence, and matched that in culms newly developed underwater. It seems that effects of both environmental and developmental factors overlap in the C4 pattern expression in this plant.  相似文献   

9.
We describe the long-term effects of a CO2 exhalation, created more than 70 years ago, on a natural C4 dominated sub-tropical grassland in terms of ecosystem structure and functioning. We tested whether long-term CO2 enrichment changes the competitive balance between plants with C3 and C4 photosynthetic pathways and how CO2 enrichment has affected species composition, plant growth responses, leaf properties and soil nutrient, carbon and water dynamics. Long-term effects of elevated CO2 on plant community composition and system processes in this sub-tropical grassland indicate very subtle changes in ecosystem functioning and no changes in species composition and dominance which could be ascribed to elevated CO2 alone. Species compositional data and soil δ13C isotopic evidence suggest no detectable effect of CO2 enrichment on C3:C4 plant mixtures and individual species dominance. Contrary to many general predictions C3 grasses did not become more abundant and C3 shrubs and trees did not invade the site. No season length stimulation of plant growth was found even after 5 years of exposure to CO2 concentrations averaging 610 μmol mol−1. Leaf properties such as total N decreased in the C3 but not C4 grass under elevated CO2 while total non-structural carbohydrate accumulation was not affected. Elevated CO2 possibly lead to increased end-of-season soil water contents and this result agrees with earlier studies despite the topographic water gradient being a confounding problem at our research site. Long-term CO2 enrichment also had little effect on soil carbon storage with no detectable changes in soil organic matter found. There were indications that potential soil respiration and N mineralization rates could be higher in soils close to the CO2 source. The conservative response of this grassland suggests that many of the reported effects of elevated CO2 on similar ecosystems could be short duration experimental artefacts that disappear under long-term elevated CO2 conditions.  相似文献   

10.
Mesophyll mitochondria from green leaves of the C(4) plants Zea mays (NADP-ME-type), Panicum miliaceum (NAD-ME-type) and Panicum maximum (PEP-CK-type) oxidized NADH, malate and succinate at relatively high rates with respiratory control, but glycine was not oxidized. Among the mitochondrial proteins involved in glycine oxidation, the L, P and T proteins of glycine decarboxylase complex (GDC) and serine hydroxymethyltransferase (SHMT) were present, while the H protein of GDC was undetectable. In contrast, mesophyll mitochondria from etiolated leaves of Z. mays oxidized glycine at a slow rate and with no respiratory control, and contained the H protein as well as the other GDC proteins and SHMT. The T and P proteins and SHMT were present in the mitochondria from etiolated leaves at significantly higher levels than in those from green leaves of Z. mays. The content of the L protein was almost identical in all three C(4) plants examined and close to the value obtained for mesophyll mitochondria from the C(3) plant Pisum sativum, whereas the other GDC proteins and SHMT were less abundant than the L protein. We discuss possible reasons for the H protein's absence in mesophyll mitochondria of C(4) plants, as well as the role(s) the other GDC components could play in its absence.  相似文献   

11.
The natural occurrence and altitudinal pattern of species with C4 photosynthesis were investigated on Qinghai Plateau, Qinghai province by using stable carbon isotopes in plant leaves and using additional data from references. A total of 58 species belonging to 10 families and 34 genera were identified using C4 photosynthetic pathway, which is only 1.66 % of total 3 500 plant species in Qinghai province. The leading two families, i.e. Gramineae (23 species) and Chenopodiaceae (22 species) contain 77.6 % of all C4 plants in the studied area. The number of C4 species increased from 1 600 to 2 400 m a.s.l. and then decreases quickly till 4 400 m a.s.l. with one half of C4 species distributing from 2 200 to 2 800 m a.s.l. (48 %). Eight plant species were found above 4 000 m a.s.l., but the distribution of these species is limited to the south of Qinghai province (low latitude area) where annual mean temperature is above 0 °C, suggesting that low temperature may generally limit the distribution of C4 plants.  相似文献   

12.
An important role in cell metabolism is played by transport of C4-dicarboxylates (C4-DCB). Specifically, they are intermediates of the citrate cycle. Transport of succinate across the mitochondrial membrane provides correlation between metabolism in peroxysomes and in mitochondria. There is known transport of C4-DCB across all kinds of energy-transforming membranes of the animal, plant, fungal, and bacterial cells. This review summarizes molecular characteristics of the C4-DCB transporters. Of particular interest are primary structures for the transporters with the known kinetic mechanism and kinetic transport parameters. For each studied group of organisms, the number of transmembrane segments in the carried molecule or the character of specificity does not correlate with the certain transport mechanism—antiport, symport with proton or symport with cation. The review describes perspective methodical approaches allowing association of peculiarities of structure with transport mechanism for individual transporters, preparation of functional hybrid transporters—“protein chimeras,” scanning of transporter transmembrane segments with aid of essential acids, probing of the transporter active center with aid of alkyl and acyl substrate derivatives used to obtain the “lipophilic profile” of the channel of the C4-DCB transporter. It is recommended to use these approaches to one transporter that has small sizes and large substrate specificity.  相似文献   

13.
The photosynthetic pathway composition (C3:C4 mixture) of an ecosystem is an important controller of carbon exchanges and surface energy flux partitioning, and therefore represents a fundamental ecophysiological distinction. To assess photosynthetic mixtures at a tallgrass prairie pasture in Oklahoma, we collected nighttime above-canopy air samples along concentration and isotopic gradients throughout the 1999 and 2000 growing seasons. We analyzed these samples for their CO2 concentration and carbon isotopic composition and calculated C3:C4 proportions with a two-source mixing model. In 1999, the C4 percentage increased from 38% in spring (late April) to 86% in early fall (mid-September). The C4 percentages inferred from ecosystem respiration measurements in 2000 indicate a smaller shift, from 67% in spring (early May) to 77% in mid-summer (late July). We also sampled daytime CO2 concentration and carbon isotope gradients above the canopy to determine ecosystem discrimination against 13CO2 during net uptake. These discrimination values were always lower than corresponding nighttime ecosystem respiration isotopic signatures would suggest. After accounting for the isotopic disequilibria between respiration and photosynthesis resulting from seasonal variations in the C3:C4 mixture, we estimated canopy photosynthetic discrimination. The C4 percentage calculated from this approach agrees with the percentage determined from nighttime respiration for sampling periods in both growing seasons. Isotopic imbalances between photosynthesis and respiration are likely to be common in mixed C3:C4 ecosystems and must be considered when using daytime isotopic measurements to constrain ecosystem physiology. Given the global extent of such ecosystems, isotopic imbalances likely contribute to global variations in the carbon isotopic composition of atmospheric CO2.  相似文献   

14.
Park J  Knoblauch M  Okita TW  Edwards GE 《Planta》2009,229(2):369-382
Bienertia sinuspersici Akhani has an unusual mechanism of C4 photosynthesis which occurs within individual chlorenchyma cells. To perform C4, the mature cells have two cytoplasmic compartments consisting of a central (CCC) and a peripheral (PCC) domain containing dimorphic chloroplasts which are interconnected by cytoplasmic channels. Based on leaf development studies, young chlorenchyma cells have not developed the two cytoplasmic compartments and dimorphic chloroplasts. Fluorescent dyes which are targeted to membranes or to specific organelles were used to follow changes in cell structure and organelle distribution during formation of C4-type chlorenchyma. Chlorenchyma cell development was divided into four stages: 1—the nucleus and chloroplasts occupy much of the cytoplasmic space and only small vacuoles are formed; 2—development of larger vacuoles, formation of a pre-CCC with some scattered chloroplasts; 3—the vacuole expands, cells have directional growth; 4—mature stage, cells have become elongated, with a distinctive CCC and PCC joined by interconnecting cytoplasmic channels. By staining vacuoles with a fluorescent dye and constructing 3D images of chloroplasts, and by microinjecting a fluorescence dye into the vacuole of living cells, it was demonstrated that the mature cell has only one vacuole, which is traversed by cytoplasmic channels connecting the CCC with the PCC. Immunofluorescent studies on isolated chlorenchyma cells treated with cytoskeleton disrupting drugs suspended in different levels of osmoticum showed that both microtubules and actin filaments are important in maintaining the cytoplasmic domains. With prolonged exposure of plants to dim light, the cytoskeleton undergoes changes and there is a dramatic shift of the CCC from the center toward the distal end of the cell.  相似文献   

15.
Hydrilla verticillata has a facultative single-cell system that changes from C3 to C4 photosynthesis. A NADP+-dependent malic enzyme (NADP-ME) provides a high [CO2] for Rubisco fixation in the C4 leaf chloroplasts. Of three NADP-ME genes identified, only hvme1 was up-regulated in the C4 leaf, during the light period, and it possessed a putative transit peptide. Unlike obligate C4 species, H. verticillata exhibited only one plastidic isoform that may perform housekeeping functions, but is up-regulated as the photosynthetic decarboxylase. Of the two cytosolic forms, hvme2 and hvme3, the latter exhibited the greatest expression, but was not light-regulated. The mature isoform of hvme1 had a pI of 6.0 and a molecular mass of 64 kD, as did the recombinant rHVME1m, and it formed a tetramer in the chloroplast. The recombinant photosynthetic isoform showed intermediate characteristics between isoforms in terrestrial C3 and C4 species. The catalytic efficiency of rHVME1m was four-fold higher than the cytosolic rHVME3 and two-fold higher than recombinant cytosolic isoforms of rice, but lower than plastidic forms of maize. The K m (malate) of 0.6 mM for rHVME1 was higher than maize plastid isoforms, but four-fold lower than found with rice. A comprehensive phylogenetic analysis of 25 taxa suggested that chloroplastic NADP-ME isoforms arose from four duplication events, and hvme1 was derived from cytosolic hvme3. The chloroplastic eudicot sequences were a monophyletic group derived from a cytosolic clade after the eudicot and monocot lineages separated, while the monocots formed a polyphyletic group. The findings support the hypothesis that a NADP-ME isoform with specific and unusual regulatory properties facilitates the functioning of the single-cell C4 system in H. verticillata. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.  相似文献   

16.
The inhibitory effect of hydrated fullerene C60 and the sodium salt of the fullerene polycarboxylic derivative C60Cl(C6H4CH2COONa)5 on the formation of amyloid fibrils by X-protein in vitro has been studied by electron microscopy. It is shown that these compounds not only destroy mature amyloid fibrils but also prevent the formation of new fibrils. This property of fullerenes, which are nanoparticles, can be used to develop a novel medical nanotechnology in the therapy for amyloidoses.  相似文献   

17.
The cell-specific distribution of the four subunit proteins (P, L, T and H) of glycine decarboxylase (GDC) and of serine hydroxymethyltransferase (SHMT) has been studied in the leaves of C3-C4 intermediate and C4 species of three genera (Flaveria, Moricandia and Panicum) using immunogold localization. Antibodies raised against these proteins from pea leaf mitochondria were used to probe Western blots of total leaf proteins of F. linearis Lag., M. arvensis (L.) DC and P. milioides Nees ex Trin. (C3-C4), and F. trinervia (Spring.) Mohr and P. miliaceum (L.) (C4). For all species, each antibody recognised specifically a protein of similar molecular weight to that in pea leaves. In leaves of M. arvensis the P protein was present in the mitochondria of the bundle-sheath cells but was undetectable in those of the mesophyll, whereas the L, T and H proteins and SHMT were present in both cell types. The density of immunogold labelling of SHMT on the mitochondria of mesophyll cells was less than that on those of the bundle-sheath cells, which correlates with the relative activities of SHMT in these cell types. These data reveal that the lack of functional GDC in the mesophyll cells of M. arvensis, which is the principal biochemical reason for reduced photorespiration in this species, is due to the loss of a single subunit protein. This lack of coordinate expression of the subunit proteins of GDC within a photosynthetic cell represents a clear difference between M. arvensis and other C3 and C3-C4 species. None of the GDC proteins was detectable in the mesophyll cells of the C3-C4 and C4 Flaveria and Panicum species but all were present in the bundle-sheath cells. The differences in the distribution of the GDC proteins in leaves of the C3-C4 species studied are discussed in relation to the evolution of photosynthetic mechanisms.  相似文献   

18.
Physical, chemical, and regulatory properties of glycolate oxidase (GO) isolated from the leaves of C4 and C3 plants (Zea mays L., cv. Voronezhskaya 76 and Glycine max (L.) Merr., cv. Pripyat’, respectively) were studied. The homogenous preparations were obtained by multistage enzyme purification from soybean leaves and maize mesophyll and bundle sheath. The glycolate oxidase (GO) preparations obtained consisted of two types of subunits, 37 and 44 kD. The GO isolated from C3 plant leaves had many in common with that extracted from C4 plant bundle sheath as regards physical, chemical, and catalytic properties. The primary function of GO in both plant types is metabolism of glycolate, which is a product of ribulosebisphosphate oxalacetic acid oxidation and is used by plants for biosynthesis of hydrocarbons and amino acids.  相似文献   

19.
A mutant (LaPr 87/30) of barley (Hordeum vulgare L.) deficient in glycine decarboxylase (GDC; EC 2.1.2.10) was crossed with wild-type plants to generate heterozygous plants with reduced GDC activities. Plants of the F2 generation were grown in air and analysed for reductions in GDC proteins and GDC activity. The leaves of heterozygous plants contained reduced amounts of H-protein, and when the content of H-protein was lower than 60% of the wild-type, the P-protein was also reduced. The contents of the other two proteins of the GDC complex, T-protein and L-protein were not affected. Glycine decarboxylase activities, measured as the decarboxylation of [1-14C]glycine by intact mitochondria released from protoplasts, were between 47% and 63% of the wild-type activity in heterozygous plants and between 86% and 100% in plants with normal contents of H-protein. The enzyme activity was linearly correlated with the relative content of H-protein. Plants with reduced GDC activities developed normally and did not show major pleiotropic effects. In air, the reduction in GDC activity had no effect on the leaf metabolite content or photosynthesis, but under conditions of enhanced photorespiration (low CO2 and high light), glycine accumulated and the rates of photosynthesis decreased compared to the wild-type. The accumulation of glycine did not lead to a depletion of amino donors or to the accumulation of glyoxylate. The lower rates of photosynthesis were probably caused by an impaired recycling of carbon in the photorespiratory pathway. It is concluded that GDC has no control over CO2 assimilation under normal growth conditions, but appreciable control by GDC becomes apparent under conditions leading to higher rates of photorespiration. Received: 24 November 1996 / Accepted: 23 January 1997  相似文献   

20.
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