Primary Carbon Assimilation Research Articles

Is Pteropyrum a pathway to C4 evolution in Polygonaceae? An integrative approach to the taxonomy and anatomy of Pteropyrum (C3), an immediate relative of Calligonum (C4)

AbstractPteropyrum is a small genus of Polygonaceae with four species from the arid regions of Iran and adjacent countries. Pteropyrum spp. are not precisely delimitated and are difficult to identify because of their high plasticity in morphological characters. Pteropyrum (C3) has a close affinity to Calligonum (C4) and is therefore a suitable case for C4 evolutionary studies. We investigated the morphology and micromorphology (including pollen morphology) of Pteropyrum and elucidated the phylogenetic relationships with Atraphaxis and Calligonum using nuclear ITS sequences. Characteristics of the photosynthetic tissues such as volume and number of layers of primary carbon assimilation tissues (PCA) and photosynthetic carbon reduction tissue (PCR) were studied. In addition, the leaf and cotyledon anatomical characters of Pteropyrum (C3), Atraphaxis (C3) and Calligonum (C4), and their δ 13C values were compared to look for evolutionary changes in assimilating organs. The molecular phylogenetic tree identifies two strongly supported clades in Pteropyrum and its close relationship with Calligonum, confirming previous studies. Some morphologically similar species belong to different clades, which is probably due to convergent evolution and homoplasy. Leaf anatomical studies show that Atraphaxis has a multilayered mesophyll tissue, whereas Calligonum has one-layered mesophyll cells. The volume and layer number of mesophyll tissue cells decreases, whereas water storage tissue area significantly increases from Atraphaxis to Pteropyrum and Calligonum. This phenomenon confirms previous studies in other lineages with C4 salsoloid anatomy that have evolved through increasing of water storage tissue and succulence of assimilating organs. In the taxonomic part of the paper, a key to identification of accepted taxa of Pteropyrum, description of species and distribution maps are presented based on numerous herbarium specimens and our own rich collections from the field. Four new species are described based on a combination of morphology of seedlings and mature plants, pollen morphology and molecular data. A subspecific classification is suggested to show morpho-geographical variation of Pteropyrum aucheri s.l.

Transcriptomic and proteomic comparison of two Miscanthus genotypes: high biomass correlates with investment in primary carbon assimilation and decreased secondary metabolism

Background and aims The plant genus Miscanthus combines high biomass production in temperate climates with low nutrient input requirements. Among the widely studied genotypes, the triploid hybrid Miscanthus x giganteus has attracted most interest because of its outstanding yields. Here, M. x giganteus was compared to its parent M. sinensis to identify traits associated with high biomass.

Responses of primary and secondary metabolism to sugar accumulation revealed by microarray expression analysis of the Arabidopsis mutant, pho3.

The Arabidopsis mutant pho3 accumulates sucrose and other carbohydrates to high levels, providing a means of investigating the genomic response to sucrose accumulation using microarray analysis. Wild-type and mutant plants were grown in soil to the mature rosette stage for the analysis of gene expression using the Affymetrix ATH1 chip, containing more than 22,500 probe sets. Small, but significant, decreases were observed in the expression of many genes encoding enzymes and regulatory proteins involved in primary carbon assimilation, suggesting that, in mature leaves of Arabidopsis, there is limited feedback regulation on gene expression by sugars. The study revealed a striking increase in the expression of the plastid glucose 6-phosphate/phosphate translocator, characteristically expressed only in heterotrophic tissues. This indicated a change in the nature of metabolite exchange between the plastid and the cytosol in the pho3 mutant. The expression of enzymes of starch synthesis also increased significantly. Very large increases were observed in the expression of transcription factors and enzymes involved in anthocyanin biosynthesis. This finding reinforces the emerging picture of an important role for primary metabolism in regulating secondary metabolism.

Ontogenetic derivation and cell differentiation in photosynthetic tissues of C3 and C4 Cyperaceae

Four variants of Kranz anatomy occur in the Cyperaceae. Three of these anatomical types (fimbristyloid, chlorocyperoid, and eleocharoid) are unique among taxa with C(4) photosynthesis in that the photosynthetic carbon reduction tissue (PCR, functional equivalent of bundle sheath) is located within the vascular strand and is separated from the primary carbon assimilation tissue (PCA, positional equivalent of mesophyll) by the mestome sheath layer. In the fourth anatomical type, rhynchosporoid, PCR tissue is located in the position of the mestome sheath. In this study, we compared two aspects of development of PCR and PCA tissues in representatives of the C(3) and C(4) types: (1) ontogenetic derivation and (2) cellular differentiation. Analysis of the planes of cell division associated with procambial strand formation indicated that PCR tissue is always derived from the procambium, while PCA tissue is derived from the ground meristem. These cell lineages remain distinct after the initial organization of vascular strands. Analysis of cell differentiation using accumulation of cell-type-specific photosynthetic enzymes as markers of differentiation indicated that, with one exception, a low level of non-cell-specific enzyme accumulation preceded abundant and cell-specific accumulation of photosynthetic enzymes at the distal end of the leaf elongation zone. Enzyme accumulation coincided spatially (and temporally) with structural aspects of cell differentiation. Previous cladistic analyses have indicated that these anatomical types represent separate evolutionary origins of the C(4) pathway, and the differences in developmental pathways observed here reflect these independent origins from C(3) ancestors.

Quantitative Leaf Anatomy of C3 and C4 Cyperaceae and Comparisons with the Poaceae

Relative amounts of photosynthetic tissues were quantified in C3 and C4 Cyperaceae species to evaluate generalizations about C4 leaf anatomy on the basis of the well-characterized Poaceae. Three of four C4 anatomical types of the Cyperaceae have an unusual version of Kranz anatomy: one or two cell layers intervene between primary carbon assimilation (PCA), equivalent to mesophyll and photosynthetic carbon reduction (PCR), equivalent to chlorenchymatous inner-border parenchyma of the vascular tissue, increasing the number of cell diameters along the diffusion pathway for C4 metabolites. The absolute distances between PCA and PCR tissues are approximately equivalent in the two families, indicating that the extra layers do not increase the mean length of the diffusion pathway required for efficient metabolite transfer in C4 photosynthesis (although minimum distances are greater in C4 Cyperaceae). The proportion of intercellular space in contact with PCA cells (including that of adjacent aerenchyma) is greate...

Mechanism of Oxidation of Dimethyl Disulphide by Thiobacillus thioparus Strain E6

Summary: A recently isolated organism, capable of chemolithoautotrophic growth on dimethyl disulphide, was characterized as a strain of Thiobacillus thioparus. It had DNA with a base composition of 60.5 ± 1.0 mol% G + C, and ubiquinone-8 (UQ-8) as its only respiratory quinone. Its growth in chemostat culture (at a growth rate of 0.07-0.08 h−1) showed yields of 14.4, 11.8 and 2.45 g cell-carbon per mol of dimethyl disulphide (DMDS), dimethyl sulphide (DMS) and thiosulphate, respectively. This is consistent with energy generation from the oxidation of the methyl and the sulphide moieties of DMDS, with oxidation of sulphide to sulphate contributing a yield of about 2.8 g cell-carbon mol−1. From whole organism and cell-free extract studies, DMDS oxidation was shown to proceed by its (NADH-stimulated) reduction to methanethiol (MT), which was oxidized via sulphide, formaldehyde and formate to CO2 and sulphate by MT oxidase, formaldehyde and formate dehydrogenases, and an uncharacterized sulphide-oxidizing system. The MT oxidase had a K m of 9.7 μM and showed substrate inhibition with a K i of about 8 μM. The essential role of catalase during growth on DMDS was shown by the sensitivity of growth on DMDS (but not on thiosulphate) to 3-amino-1,2,4-triazole. Catalase is believed to destroy the peroxide produced by the MT oxidase reaction. DMDS-grown organisms oxidized sulphide, thiosulphate and tetrathionate (with the latter indicated to be an intermediate in thiosulphate oxidation), suggesting the pathway of sulphide oxidation to be similar to that in some other thiobacilli. Carbon assimilation was by the Calvin cycle, with ribulose bisphosphate carboxylase being present in cell-free extracts at a specific activity of 80 nmol CO2 fixed min−1 (mg protein)−1. Hydroxypyruvate reductase (HPR) was not detected at levels sufficient to indicate any role in primary carbon assimilation.