Non-photosynthetic Tissues Research Articles

Carotenoid-carbohydrate crosstalk: evidence for genetic and physiological interactions in storage tissues across crop species.

Carotenoids play essential roles in photosynthesis, photoprotection, and human health. Efforts to increase carotenoid content in several staple crops have been successful through both conventional selection and genetic engineering methods. Interestingly, in some cases, altering carotenoid content has had unexpected effects on other aspects of plant metabolism, impacting traits like sugar content, dry matter percentage, fatty acid content, stress tolerance, and phytohormone concentrations. Studies across several diverse crop species have identified negative correlations between carotenoid and starch contents, as well as positive correlations between carotenoids and soluble sugars. Collectively, these reports suggest a metabolic interaction between carotenoids and carbohydrates. We synthesize evidence pointing to four hypothesized mechanisms: (1) direct competition for precursors; (2) physical interactions in plastids; (3) influences of sugar or apocarotenoid signaling networks; and (4) nonmechanistic population or statistical sources of correlations. Though the carotenoid biosynthesis pathway is well understood, the regulation and interactions of carotenoids, especially in nonphotosynthetic tissues, remain unclear. This topic represents an underexplored interplay between primary and secondary metabolism where further research is needed.

Research progress on differentiation and regulation of plant chromoplasts.

Carotenoids, natural tetraterpenoids found abundantly in plants, contribute to the diverse colors of plant non-photosynthetic tissues and provide fragrance through their cleavage products, which also play crucial roles in plant growth and development. Understanding the synthesis, degradation, and storage pathways of carotenoids and identifying regulatory factors represents a significant strategy for enhancing plant quality. Chromoplasts serve as the primary plastids responsible for carotenoid accumulation, and their differentiation is linked to the levels of carotenoids, rendering them a subject of substantial research interest. The differentiation of chromoplasts involves alterations in plastid structure and protein import machinery. Additionally, this process is influenced by factors such as the ORANGE (OR) gene, Clp proteases, xanthophyll esterification, and environmental factors. This review shows the relationship between chromoplast and carotenoid accumulation by presenting recent advances in chromoplast structure, the differentiation process, and key regulatory factors, which can also provide a reference for rational exploitation of chromoplasts to enhance plant quality.

Complementary environmental analysis and functional characterization of lower glycolysis-gluconeogenesis in the diatom plastid.

Organic carbon fixed in chloroplasts through the Calvin-Benson-Bassham Cycle can be diverted toward different metabolic fates, including cytoplasmic and mitochondrial respiration, gluconeogenesis, and synthesis of diverse plastid metabolites via the pyruvate hub. In plants, pyruvate is principally produced via cytoplasmic glycolysis, although a plastid-targeted lower glycolytic pathway is known to exist in non-photosynthetic tissue. Here, we characterized a lower plastid glycolysis-gluconeogenesis pathway enabling the direct interconversion of glyceraldehyde-3-phosphate and phospho-enol-pyruvate in diatoms, ecologically important marine algae distantly related to plants. We show that two reversible enzymes required to complete diatom plastid glycolysis-gluconeogenesis, Enolase and bis-phosphoglycerate mutase (PGAM), originated through duplications of mitochondria-targeted respiratory isoforms. Through CRISPR-Cas9 mutagenesis, integrative 'omic analyses, and measured kinetics of expressed enzymes in the diatom Phaeodactylum tricornutum, we present evidence that this pathway diverts plastid glyceraldehyde-3-phosphate into the pyruvate hub, and may also function in the gluconeogenic direction. Considering experimental data, we show that this pathway has different roles dependent in particular on day length and environmental temperature, and show that the cpEnolase and cpPGAM genes are expressed at elevated levels in high-latitude oceans where diatoms are abundant. Our data provide evolutionary, meta-genomic, and functional insights into a poorly understood yet evolutionarily recurrent plastid metabolic pathway.

Changes in carbon metabolism and dissolved organic carbon fluxes on seagrass patches (Halodule wrightii) with different ages in Southern Gulf of California

Seagrass meadows are highly dynamic, particularly in sites where extreme climatological events may produce a mosaic of patches with different ages. This study evaluates the community carbon metabolism, dissolved organic carbon (DOC) fluxes and associated fauna in patches of Halodule wrightii with different ages since establishment. Net community production declined as patch age increased, probably due to the increase in non-photosynthetic tissues, higher respiration rates of the community assemblage and a likely increase in self-shading of the canopy. The export of DOC was significantly higher in the youngest patches, mainly as a consequence of the lower seagrass net production recorded in older meadows. We concluded that ‘colonizers’ seagrass species may show higher production rates and DOC release during the first stages of colonization, which suggest that, the production, organic carbon exportation and their role as relevant blue carbon communities may be higher than expected.

Photosynthesis and whole-plant carbon balances of warm affinity Halophila nipponica and cold affinity Zostera marina in relation to water temperature rise: implication for future geographic distribution

Climate change associated sea surface temperature rise could significantly affect the physiology, and consequently geographic distribution of seagrasses. Photosynthetic and respiratory responses of warm affinity Halophila nipponica and cold affinity Zostera marina to a change in water temperature (7–26°C) were investigated to estimate whole-plant carbon balance for prediction of the future geographic distributions of these seagrasses in the northwestern Pacific region. It was hypothesized that H. nipponica and Z. marina showed distinctly different patterns of carbon balance with changes in water temperature. Photosynthetic and respiratory rates of H. nipponica and Z. marina generally increased with increasing water temperature, except for Ic and Ik of H. nipponica and α of Z. marina. H. nipponica showed high Pmax and α values, which could support a greater respiratory demand related to the higher proportion of non-photosynthetic tissues (~67%), compared with Z. marina (~34%). The whole-plant carbon balance of Z. marina remained positive throughout the experiment, although it decreased to nearly zero during the high water-temperature period. H. nipponica exhibited a negative carbon balance during winter and early spring, which corresponded with severely limited growth at water temperatures< 15°C. These results suggest that increases in water temperature, particularly during winter, in relation to continuous climate change, could induce substantial changes in the seagrass ecosystem structure and corresponding changes in coastal ecosystem services in the temperate coastal waters of the northwestern Pacific region. The results of this study will provide valuable information on the effective management and conservation of coastal and estuarine ecosystems under ongoing climate change.

18 O enrichment of sucrose and photosynthetic and nonphotosynthetic leaf water in a C3 grass-atmospheric drivers and physiological relations.

The 18 O enrichment (Δ18 O) of leaf water affects the Δ18 O of photosynthetic products such as sucrose, generating an isotopic archive of plant function and past climate. However, uncertainty remains as to whether leaf water compartmentation between photosynthetic and nonphotosynthetic tissue affects the relationship between Δ18 O of bulk leaf water (Δ18 OLW ) and leaf sucrose (Δ18 OSucrose ). We grew Lolium perenne (a C3 grass) in mesocosm-scale, replicated experiments with daytime relative humidity (50% or 75%) and CO2 level (200, 400 or 800 μmol mol-1 ) as factors, and determined Δ18 OLW , Δ18 OSucrose and morphophysiological leaf parameters, including transpiration (Eleaf ), stomatal conductance (gs ) and mesophyll conductance to CO2 (gm ). The Δ18 O of photosynthetic medium water (Δ18 OSSW ) was estimated from Δ18 OSucrose and the equilibrium fractionation between water and carbonyl groups (εbio ). Δ18 OSSW was well predicted by theoretical estimates of leaf water at the evaporative site (Δ18 Oe ) with adjustments that correlated with gas exchange parameters (gs or total conductance to CO2 ). Isotopic mass balance and published work indicated that nonphotosynthetic tissue water was a large fraction (~0.53) of bulk leaf water. Δ18 OLW was a poor proxy for Δ18 OSucrose , mainly due to opposite Δ18 O responses of nonphotosynthetic tissue water (Δ18 Onon-SSW ) relative to Δ18 OSSW , driven by atmospheric conditions.

Beyond width and density: stable carbon and oxygen isotopes in cork-rings provide insights of physiological responses to water stress in Quercus suber L.

As climate change increasingly affects forest ecosystems, detailed understanding of major effects is important to anticipate their consequences under future climate scenarios. The Mediterranean region is a prominent climate change hotspot, and evergreen cork oak (Quercus suber L.) woodlands are particularly climatically sensitive due to cork (bark) harvesting. Cork oak's drought avoidance strategy is well-known and includes structural and physiological adaptations that maximise soil water uptake and transport and limit water use, potentially leading to reduced stem and cork growth. Trees' responses to cope with water-limited conditions have been extensively described based on cork-rings width and, more recently, on cork-rings density, in dendroecological studies. However, so far, tree functional attributes and physiological strategies, namely photosynthetic metabolism adjustments affecting cork formation, have never been addressed and/or integrated on these previous cork-rings-based studies. In this study, we address the relation between carbon and oxygen stable isotopes of cork rings and precipitation and temperature, in two distinct locations of southwestern Portugal-the (wetter) Tagus basin peneplain and the (drier) Grândola mountains. We aimed at assessing whether the two climatic factors affect cork-ring isotopic composition under contrasting conditions of water availability, and, therefore, if carbon and oxygen signatures in cork can reflect tree functional (physiological and structural) responses to stressful conditions, which might be aggravated by climate change. Our results indicate differences between the study areas. At the drier site, the stronger statistically significant negative cork δ 13C correlations were found with mean temperature, whereas strong positive cork δ 18O correlations were fewer and found only with precipitation. Moreover, at the wetter site, cork rings are enriched in 18O and depleted in 13C, indicating, respectively, shallow groundwater as the water source for physiological processes related with biosynthesis of non-photosynthetic secondary tissues, such as suberin, and a weak stomatal regulation under high water availability, consistent with non-existent water availability constrains. In contrast, at the drier site, trees use water from deeper ground layers, depleted in 18O, and strongly regulate stomatal conductance under water stress, thus reducing photosynthetic carbon uptake and probably relying on stored carbon reserves for cork ring formation. These results suggest that although stable isotopes signatures in cork rings are not proxies for net growth, they may be (fairly) robust indicators of trees' physiological and structural adjustments to climate and environmental changes in Mediterranean environments.

Identification and expression analysis of pineapple sugar transporters reveal their role in the development and environmental response.

In plants, sugars are required for several essential functions, including growth, storage, signaling, defense and reproduction. Sugar transporters carry out the controlled movement of sugars from source (leaves) to sink (fruits and roots) tissues and determine the overall development of the plant. Various types of sugar transporter families have been described in plants, including sucrose transporters (SUC/SUT), monosaccharide transporter (MST) and SWEET (from "Sugar Will Eventually be Exported Transporters"). However, the information about pineapple sugar transporters is minimal. This study systematically identified and classified 45 MST and 4 SUC/SUT genes in the pineapple genome. We found that the expression patterns of sugar transporter genes have a spatiotemporal expression in reproductive and vegetative tissues indicating their pivotal role in reproductive growth and development. Besides, different families of sugar transporters have a diel expression pattern in photosynthetic and non-photosynthetic tissues displaying circadian rhythm associated participation of sugar transporters in the CAM pathway. Moreover, regulation of the stress-related sugar transporters during cold stress indicates their contribution to cold tolerance in pineapple. Heterologous expression (yeast complementation assays) of sugar transporters in a mutant yeast strain suggested that SUT1/2 have the ability to transport sucrose, and STP13, STP26, pGlcT-L2 and TMT4 are able to transport glucose, whereas SWEET11/13 transport both sucrose and fructose. The information provided here would help researchers further explore the underlying molecular mechanism involved in the sugar metabolism of pineapple.

Source-To-Sink Transport of Sugar and Its Role in Male Reproductive Development.

Sucrose is produced in leaf mesophyll cells via photosynthesis and exported to non-photosynthetic sink tissues through the phloem. The molecular basis of source-to-sink long-distance transport in cereal crop plants is of importance due to its direct influence on grain yield—pollen grains, essential for male fertility, are filled with sugary starch, and rely on long-distance sugar transport from source leaves. Here, we overview sugar partitioning via phloem transport in rice, especially where relevant for male reproductive development. Phloem loading and unloading in source leaves and sink tissues uses a combination of the symplastic, apoplastic, and/or polymer trapping pathways. The symplastic and polymer trapping pathways are passive processes, correlated with source activity and sugar gradients. In contrast, apoplastic phloem loading/unloading involves active processes and several proteins, including SUcrose Transporters (SUTs), Sugars Will Eventually be Exported Transporters (SWEETs), Invertases (INVs), and MonoSaccharide Transporters (MSTs). Numerous transcription factors combine to create a complex network, such as DNA binding with One Finger 11 (DOF11), Carbon Starved Anther (CSA), and CSA2, which regulates sugar metabolism in normal male reproductive development and in response to changes in environmental signals, such as photoperiod.

An Updated Review on the Modulation of Carbon Partitioning and Allocation in Arbuscular Mycorrhizal Plants.

Arbuscular mycorrhizal fungi (AMF) are obligate biotrophs that supply mineral nutrients to the host plant in exchange for carbon derived from photosynthesis. Sucrose is the end-product of photosynthesis and the main compound used by plants to translocate photosynthates to non-photosynthetic tissues. AMF alter carbon distribution in plants by modifying the expression and activity of key enzymes of sucrose biosynthesis, transport, and/or catabolism. Since sucrose is essential for the maintenance of all metabolic and physiological processes, the modifications addressed by AMF can significantly affect plant development and stress responses. AMF also modulate plant lipid biosynthesis to acquire storage reserves, generate biomass, and fulfill its life cycle. In this review we address the most relevant aspects of the influence of AMF on sucrose and lipid metabolism in plants, including its effects on sucrose biosynthesis both in photosynthetic and heterotrophic tissues, and the influence of sucrose on lipid biosynthesis in the context of the symbiosis. We present a hypothetical model of carbon partitioning between plants and AMF in which the coordinated action of sucrose biosynthesis, transport, and catabolism plays a role in the generation of hexose gradients to supply carbon to AMF, and to control the amount of carbon assigned to the fungus.

Two evolutionarily duplicated domains individually and post-transcriptionally control SWEET expression for phloem transport.

Cell type-specific gene expression is critical for the specialized functions within multicellular organisms. In Arabidopsis,SWEET11andSWEET12sugar transporters are specifically expressed in phloem parenchyma (PP) cells and are responsible for sucrose efflux from the PP, the first step of a two-step apoplasmic phloem-loading strategy that initiates the long-distance transport of sugar from leaves to nonphotosynthetic sink tissues. However, we know nothing about what determines the PP cell-specific expression of theseSWEETs. Sequence deletions, histochemical β-glucuronidase (GUS) analysis, cross-sectioning, live-cell imaging, and evolutionary analysis were used to elucidate domains responsible for PP specificity, while a Förster resonance energy transfer (FRET) sensor-based transport assay was used to determine whether substrate specificity coevolved with PP specificity. We identified two domains in the ArabidopsisSWEET11coding sequence that, along with its promoter (including 5' UTR), regulate PP-specific expression at the post-transcriptional level, probably involving RNA-binding proteins. This mechanism is conserved among vascular plants but independent of transport substrate specificity. We conclude that two evolutionarily duplicated coding sequence domains are essential and individually sufficient for PP-specific expression of SWEET11. We also provide a crucial experimental tool to study PP physiology and development.

Hidden fractions: Another look at micronutrient and sodium partitioning in modern cotton cultivars

AbstractCotton (Gossypium hirsutum L.) productivity and nutritional status depend on the uptake and allocation of both macro‐ and micronutrients. Micronutrient deficiencies or toxicities reduce yields and must be corrected to achieve optimal yields. There is evidence that macronutrient uptake and partitioning have changed in modern cultivars, and it is therefore important to know whether a similar change has occurred with micronutrient uptake or partitioning. Total uptake and partitioning of essential micronutrients (boron, B; iron, Fe; manganese, Mn; zinc, Zn; and copper, Cu) and of sodium (Na) in different tissues of modern cultivars (FiberMax 958, FM 958; and Deltapine 1646, DP 1646) and a 1990s cultivar (Paymaster HS26, PM HS26) were determined in 2018 and 2019 at New Deal, TX. These were compared to results obtained in studies conducted 30 yr ago. The total seasonal uptake of Cu and Zn increased, whereas the total seasonal uptake of Fe and Mn decreased in modern cultivars. Newer cultivars FM 958 and DP 1646 partitioned greater percentage of Zn and Cu to the developing bolls compared to PM HS26. Iron and Mn were mostly retained in the leaves throughout the crop's life cycle. Sodium accumulated in nonphotosynthetic tissues likely as a salt‐tolerance mechanism of cotton. However, when Na availability is far more than required for normal plant growth, Na is redistributed and accumulated into transpiring leaves. This study offers additional insights for making effective fertilizer management decisions for newer cultivars and possible problems due to excess trace elements in the soil and irrigation supply.

Evolutionary loss of thermal acclimation accompanied by periodic monocarpic mass flowering in Strobilanthes flexicaulis

While life history, physiology and molecular phylogeny in plants have been widely studied, understanding how physiology changes with the evolution of life history change remains largely unknown. In two closely related understory Strobilanthes plants, the molecular phylogeny has previously shown that the monocarpic 6-year masting S. flexicaulis have evolved from a polycarpic perennial, represented by the basal clade S. tashiroi. The polycarpic S. tashiroi exhibited seasonal thermal acclimation with increased leaf respiratory and photosynthetic metabolism in winter, whereas the monocarpic S. flexicaulis showed no thermal acclimation. The monocarpic S. flexicaulis required rapid height growth after germination under high intraspecific competition, and the respiration and N allocation were biased toward nonphotosynthetic tissues. By contrast, in the long-lived polycarpic S. tashiroi, these allocations were biased toward photosynthetic tissues. The life-history differences between the monocarpic S. flexicaulis and the polycarpic S. tashiroi are represented by the “height growth” and “assimilation” paradigms, respectively, which are controlled by different patterns of respiration and nitrogen regulation in leaves. The obtained data indicate that the monocarpic S. flexicaulis with the evolutionary loss of thermal acclimation may exhibit increased vulnerability to global warming.

Differential effects of nitrogen vs. phosphorus limitation on terrestrial carbon storage in two subtropical forests: A Bayesian approach

Nitrogen (N) and phosphorus (P) have been demonstrated to limit terrestrial carbon (C) storage in terrestrial ecosystems. However, the reliable indicator to infer N and P limitation are still lacking, especially in subtropical forests. Here we used a terrestrial ecosystem (TECO) model framework in combination with a Bayesian approach to evaluate effects of nutrient limitation from added N/P processes and data sets on C storage capacities in two subtropical forests (Tiantong and Qianyanzhou [QYZ]). Three of the six simulation experiments were developed with assimilating data (TECO C model with C data [C-C], TECO C-N coupling model with C and N data [CN-CN], and TECO C-N-P model with C, N, and P data [CNP-CNP]), and the other three ones were simulated without assimilating data (C-only, CN-only, and CNP-only). We found that P dominantly constrained C storage capacities in Tiantong (42%) whereas N limitation decreased C storage projections in QYZ (44%). Our analysis indicated that the stoichiometry of wood biomass and soil microbe (e.g., N:P ratio) were more sensitive indicators of N or P limitation than that of other pools. Furthermore, effects of P-induced limitation were mainly on root biomass by additional P data and on both metabolic litter and soil organic carbon (SOC) by added P processes. N-induced effects were mainly from added N data that limited plant non-photosynthetic tissues (e.g., woody biomass and litter). The different effects of N and P modules on C storage projections reflected the diverse nutrient acquisition strategies associated with stand ages and plant species under nutrient stressed environment. These findings suggest that the interaction between plants and microorganisms regulate effects of nutrient availability on ecosystem C storage, and stoichiometric flexibility of N and P in plant and soil C pools could improve the representation of N and P limitation in terrestrial ecosystem models.

Tissue-type specific accumulation of the plastoglobular proteome, transcriptional networks, and plastoglobular functions.

Plastoglobules are dynamic protein-lipid microcompartments in plastids enriched for isoprenoid-derived metabolites. Chloroplast plastoglobules support formation, remodeling, and controlled dismantling of thylakoids during developmental transitions and environmental responses. However, the specific molecular functions of most plastoglobule proteins are still poorly understood. This review harnesses recent co-mRNA expression data from combined microarray and RNA-seq information in ATTED-II on an updated inventory of 34 PG proteins, as well as proteomics data across 30 Arabidopsis tissue types from ATHENA. Hierarchical clustering based on relative abundance for the plastoglobule proteins across non-photosynthetic and photosynthetic tissue types showed their coordinated protein accumulation across Arabidopsis parts, tissue types, development, and senescence. Evaluation of mRNA-based forced networks at different coefficient thresholds identified a central hub with seven plastoglobule proteins and four peripheral modules. Enrichment of specific nuclear transcription factors (e.g. Golden2-like) and support for crosstalk between plastoglobules and the plastid gene expression was observed, and specific ABC1 kinases appear part of a light signaling network. Examples of other specific findings are that FBN7b is involved with upstream steps of tetrapyrrole biosynthesis and that ABC1K9 is involved in starch metabolism. This review provides new insights into the functions of plastoglobule proteins and an improved framework for experimental studies.

Sucrose-phosphate phosphatase from sugarcane reveals an ancestral tandem duplication

BackgroundSugarcane is capable to store large amounts of sucrose in the culm at maturity hence it became a major source of sucrose for the food and the renewable energy industries. Sucrose, the main disaccharide produced by photosynthesis, is mainly stored in the vacuole of the cells of non-photosynthetic tissues. Two pathways are known to release free sucrose in plant cells, one is de novo synthesis dependent on sucrose phosphate synthase (SPS) and sucrose phosphate phosphatase (S6PP) while the other is regulatory and dependent on sucrose synthase (SuSy) activity. The molecular understanding of genes that give rise to the expression of the enzyme sucrose phosphate phosphatase, responsible for the release of sucrose in the last synthetic step lag behind the regulatory SuSy gene.ResultsSugarcane genome sequencing effort disclosed the existence of a tandem duplication and the present work further support that both S6PP.1 and S6PP_2D isoforms are actively transcribed in young sugarcane plants but significantly less at maturity. Two commercial hybrids (SP80–3280 and R570) and both Saccharum spontaneum (IN84–58) and S.officinarum (BADILLA) exhibit transcriptional activity at three-month-old plants of the tandem S6PP_2D in leaves, culm, meristem and root system with a cultivar-specific distribution. Moreover, this tandem duplication is shared with other grasses and is ancestral in the group.ConclusionDetection of a new isoform of S6PP resulting from the translation of 14 exon-containing transcript (S6PP_2D) will contribute to the knowledge of sucrose metabolism in plants. In addition, expression varies along plant development and between sugarcane cultivars and parental species.

Integrated analysis of DNA methylome and transcriptome reveals epigenetic regulation of CAM photosynthesis in pineapple

BackgroundCrassulacean acid metabolism (CAM) photosynthesis is an important carbon fixation pathway especially in arid environments because it leads to higher water-use efficiency compared to C3 and C4 plants. However, the role of DNA methylation in regulation CAM photosynthesis is not fully understood.ResultsHere, we performed temporal DNA methylome and transcriptome analysis of non-photosynthetic (white base) and photosynthetic (green tip) tissues of pineapple leaf. The DNA methylation patterns and levels in these two tissues were generally similar for the CG and CHG cytosine sequence contexts. However, CHH methylation was reduced in white base leaf tissue compared with green tip tissue across diel time course in both gene and transposon regions. We identified thousands of local differentially methylated regions (DMRs) between green tip and white base at different diel periods. We also showed that thousands of genes that overlapped with DMRs were differentially expressed between white base and green tip leaf tissue across diel time course, including several important CAM pathway-related genes, such as beta-CA, PEPC, PPCK, and MDH.ConclusionsTogether, these detailed DNA methylome and transcriptome maps provide insight into DNA methylation changes and enhance our understanding of the relationships between DNA methylation and CAM photosynthesis.

The Phosphoglycerate Kinase (PGK) Gene Family of Maize (Zea mays var. B73).

Phosphoglycerate kinase (PGK, E.C. 2.7.2.3) interconverts ADP + 1,3-bisphospho-glycerate (1,3-bPGA) to ATP + 3-phosphoglycerate (3PGA). While most bacteria have a single pgk gene and mammals possess two copies, plant genomes contain three or more PGK genes. In this study, we identified five Pgk genes in the Zea mays var. B73 genome, predicted to encode proteins targeted to different subcellular compartments: ZmPgk1, ZmPgk2, and ZmPgk4 (chloroplast), ZmPgk3 (cytosol), and ZmPgk5 (nucleus). The expression of ZmPgk3 was highest in non-photosynthetic tissues (roots and cobs), where PGK activity was also greatest, consistent with a function in glycolysis. Green tissues (leaf blade and husk leaf) showed intermediate levels of PGK activity, and predominantly expressed ZmPgk1 and ZmPgk2, suggesting involvement in photosynthetic metabolism. ZmPgk5 was weakly expressed and ZmPgk4 was not detected in any tissue. Phylogenetic analysis showed that the photosynthetic and glycolytic isozymes of plants clustered together, but were distinct from PGKs of animals, fungi, protozoa, and bacteria, indicating that photosynthetic and glycolytic isozymes of plants diversified after the divergence of the plant lineage from other groups. These results show the distinct role of each PGK in maize and provide the basis for future studies into the regulation and function of this key enzyme.

Unexpectedly low δ 13C in leaves, branches, stems and roots of three acacia species growing in hyper-arid environments

AbstractAimsIn plant eco-physiology, less negative (enriched) carbon 13 (13C) in the leaves indicates conditions of reducing leaf gas exchange through stomata, e.g. under drought. In addition, 13C is expected to be less negative in non-photosynthetic tissues as compared with leaves. However, these relationships in δ 13C from leaves (photosynthetic organs) to branches, stems and roots (non-photosynthetic organs) are rarely tested across multiple closely related tree species, multiple compartments, or in trees growing under extreme heat and drought.MethodsWe measured leaf-to-root 13C in three closely related desert acacia species (Acacia tortilis, A. raddiana and A. pachyceras). We measured δ 13C in leaf tissues from mature trees in southern Israel. In parallel, a 7-year irrigation experiment with 0.5, 1.0 or 4.0 L day−1 was conducted in an experimental orchard. At the end of the experiment, growth parameters and δ 13C were measured in leaves, branches, stems and roots.Important FindingsThe δ 13C in leaf tissues sampled from mature trees was ca. −27‰, far more depleted than expected from a desert tree growing in one of the Earth’s driest and hottest environments. Across acacia species and compartments, δ 13C was not enriched at all irrigation levels (−28‰ to ca. −27‰), confirming our measurements in the mature trees. Among compartments, leaf δ 13C was unexpectedly similar to branch and root δ 13C, and surprisingly, even less negative than stem δ 13C. The highly depleted leaf δ 13C suggests that these trees have high stomatal gas exchange, despite growing in extremely dry habitats. The lack of δ 13C enrichment in non-photosynthetic tissues might be related to the seasonal coupling of growth of leaves and heterotrophic tissues.

Altering Tetrapyrrole Biosynthesis by Overexpressing Ferrochelatases (Fc1 and Fc2) Improves Photosynthetic Efficiency in Transgenic Barley

Ferrochelatase (FC) is the terminal enzyme of heme biosynthesis. In photosynthetic organisms studied so far, there is evidence for two FC isoforms, which are encoded by two genes (FC1 and FC2). Previous studies suggest that these two genes are required for the production of two physiologically distinct heme pools with only FC2-derived heme involved in photosynthesis. We characterised two FCs in barley (Hordeum vulgare L.). The two HvFC isoforms share a common catalytic domain, but HvFC2 additionally contains a C-terminal chlorophyll a/b binding (CAB) domain. Both HvFCs are highly expressed in photosynthetic tissues, with HvFC1 transcripts also being abundant in non-photosynthetic tissues. To determine whether these isoforms differentially affect photosynthesis, transgenic barley ectopically overexpressing HvFC1 and HvFC2 were generated and evaluated for photosynthetic performance. In each case, transgenics exhibited improved photosynthetic rate (Asat), stomatal conductance (gs) and carboxylation efficiency (CE), showing that both FC1 and FC2 play important roles in photosynthesis. Our finding that modified FC expression can improve photosynthesis up to ~13% under controlled growth conditions now requires further research to determine if this can be translated to improved yield performance under field conditions.