Non-photosynthetic Organs Research Articles

When the Mediterranean becomes harsh: Heat pulses strongly affect C allocation in plant-soil-atmosphere continuum in Eucalyptus camaldulensis

Mediterranean vegetation is frequently subjected to the heat waves with temperature peaks exceeding 45 °C. While the effects of drought on assimilates allocation are deeply studied, little information is available regarding the effects of the acute heat stress on this aspect of the plant physiology. We evaluated the effects of acute heat pulses on plant C internal allocation in Eucalyptus camaldulensis saplings and its transfer into the soil. Shoots of Eucalyptus were subjected to an increase of the air temperatures up to 45 °C. Allocation of C was inferred after pulse labelling of heated and non-heated shoots with 13CO2. Heat pulse lowered the photosynthesis and almost tripled the C losses from shoots through respiration. Assimilated 13C was preferentially allocated to leaf soluble sugar fractions rather than to starch. Heat pulse interrupted 13C allocation to the trunk, roots and soil. 13C allocation to new non labelled leaves was detected in some replicate plants, independently of treatment, suggesting that this process is sink-driven. Although, shoots subjected to the heat pulse modified allocation strategy, “saving” assimilated 13C at a leaf level, the rate of 13C involvement in shoot respiration was restricted by photosynthetic rates measured during the labelling. The results suggest that photosynthetic and non-photosynthetic organs may be constrained to rely on C reserves under elevated temperatures with further negative consequences for C balance in case the duration and frequency of the heat waves will increase as projected by climate change.

The distribution of vertical leaves and leaves biomass on ten mangrove species at Ngurah Rai Forest Park, Denpasar, Bali, Indonesia

Ardhana IPG, Rimbawan IMGS., Cahyo PN, Fitriani Y, Rohani S. 2018. The distribution of vertical leaves and leaves biomass on ten mangrove species at Ngurah Rai Forest Park, Denpasar, Bali, Indonesia. Biodiversitas 19: 918-926. The distribution of vertical leaves and leaves biomass which forms the mangrove crown divides into three parts, namely the top part, the middle part, and the bottom part. Total amount leaves on any parts are highly variable depending on height of each species. The relationship between the height of tree with the crown thickness depended on total leaves of each crown on each species as well as the relationship between the total leaves biomass of each crown on each species, with the height and volume of tree stems, also has variation on each mangrove species. Author examined the relationship between the photosynthetic organs with non-photosynthetic organs, especially on stems in each species. The high total amount of leaves and large total amount leaves biomass deeply is depended on diameter and growth height which are presented in the form of stem volumes. On the relationship between the photosynthetic organs and non-photosynthetic organs on each mangrove species, also has been examined. The higher total amount of leaves or the total amount of leaves biomass largely depended on the stem volume of mangroves. This indicates that the production of photosynthetic organ either in the form of the total amount of leaves or the total amount of leaves biomass serve to support growth of the stem mangrove trees (height of stem, diameter of stem and volume of stem).

Phosphatidylglycerophosphate phosphatase is required for root growth in Arabidopsis

Phosphatidylglycerol (PG) is an indispensable lipid class in photosynthetic activity. However, the importance of PG biosynthesis in non-photosynthetic organs remains elusive. We previously identified phosphatidylglycerophosphate phosphatase 1 (PGPP1), which catalyzes the last step of PG biosynthesis in Arabidopsis thaliana. In the present report, we noted considerably shorter roots of the pgpp1-1 mutant compared to the wild type. We observed defective order of columella cells in the root apices, which was complemented by introducing the wild-type PGPP1 gene. Although PGPP1 is chloroplast-localized in leaf mesophyll cells, we observed mitochondrial localization of PGPP1 in root cells, suggesting possible dual targeting of PGPP1. Moreover, we identified previously uncharacterized 2 protein tyrosine phosphatase-like proteins as functional PGPPs. These proteins, designated PTPMT1 and PTPMT2, complemented growth and lipid phenotypes of Δgep4, a Saccharomyces cerevisiae mutant of PGPP. The ptpmt1-1 ptpmt2-1 exhibited no visible phenotype; however, the pgpp1-1 ptpmt1-1 ptpmt2-1 significantly enhanced the root phenotype of pgpp1-1 without further affecting the photosynthesis, suggesting that these newly found PGPPs are involved in the root phenotype. Radiolabeling experiment of mutant roots showed that decreased PG biosynthesis is associated with the mutation of PGPP1. These results suggest that PG biosynthesis is required for the root growth.

Carbohydrate dynamics of three dominant species in a Chinese savanna under precipitation exclusion.

The potential impact of drought on the carbon balance in plants has gained great attention. Non-structural carbohydrate (NSC) dynamics have been suggested as an important trait reflecting carbon balance under drought conditions. However, NSC dynamics under drought and the response mechanisms of NSC to drought remain unclear, especially in water-limited savanna ecosystems. A precipitation exclusion experiment was performed to simulate different drought intensities in a savanna ecosystem in Yuanjiang valley in southwestern China. Growth, total NSC concentration and diurnal change of NSC were determined for the leaves and non-photosynthetic organs of three dominant species (Lannea coromandelica, Polyalthia cerasoides and Heteropogon contortus) throughout the growing season. Drought significantly reduced the growth of all the three species. Total NSC concentration averaged ~8.1%, varying with species, organ and sampling period, and did not significantly decrease under drought stress. By contrast, the diurnal change of NSC in these three species increased under drought stress. These results indicate that these three dominant species did not undergo carbon limitation. Thus, relative change in NSC is a more sensitive and effective indicator than carbon reserves in evaluation of plant carbon balance. These findings provide new insights for the understanding of carbon balance and the mechanisms of carbon starvation.

Fluorescein Transport Assay to Assess Bulk Flow of Molecules Through the Hypocotyl in Arabidopsis thaliana.

The bulk transport of molecules through plant tissues underpins growth and development. The stem acts as a conduit between the upper and low domains of the plant, facilitating transport of solutes and water from the roots to the shoot system, and sugar plus other elaborated metabolites towards the non-photosynthetic organs. In order to perform this function efficiently, the stem needs to be optimized for transport. This is achieved through the formation of vasculature that connects the whole plant but also through connectivity signatures that reduce path length distributions outside the vascular system. This protocol was devised to characterize how cell connectivity affects the bulk flow of molecules traversing the stem. This is achieved by exposing young seedlings to fluorescein, for which no specific transporter is assumed to be present in A. thaliana, and assessing the relative concentration of this fluorescent compound in individual cells of the embryonic stem (hypocotyl) using confocal microscopy and quantitative 3D image analysis after a given exposure time.

Biomass Allocation Patterns Are Linked to Genotypic Differences in Whole-Plant Transpiration Efficiency in Sunflower.

Increased transpiration efficiency (the ratio of biomass to water transpired, TE) could lead to increased drought tolerance under some water deficit scenarios. Intrinsic (i.e., leaf-level) TE is usually considered as the primary source of variation in whole-plant TE, but empirical data usually contradict this assumption. Sunflower has a significant variability in TE, but a better knowledge of the effect of leaf and plant-level traits could be helpful to obtain more efficient genotypes for water use. The objective of this study was, therefore, to assess if genotypic variation in whole-plant TE is better related to leaf- or plant-level traits. Three experiments were conducted, aimed at verifying the existence of variability in whole-plant TE and whole-plant and leaf-level traits, and to assess their correlation. Sunflower public inbred lines and a segregating population of recombinant inbred lines were grown under controlled conditions and subjected to well-watered and water-deficit treatments. Significant genotypic variation was found for TE and related traits. These differences in whole-plant transpiration efficiency, both between genotypes and between plants within each genotype, showed no association to leaf-level traits, but were significantly and negatively correlated to biomass allocation to leaves and to the ratio of leaf area to total biomass. These associations are likely of a physiological origin, and not only a consequence of genetic linkage in the studied population. These results suggest that genotypic variation for biomass allocation could be potentially exploited as a source for increased transpiration efficiency in sunflower breeding programmes. It is also suggested that phenotyping for TE in this species should not be restricted to leaf-level measurements, but also include measurements of plant-level traits, especially those related to biomass allocation between photosynthetic and non-photosynthetic organs.

Molecular regulation of sucrose catabolism and sugar transport for development, defence and phloem function.

Sucrose (Suc) is the major end product of photosynthesis in mesophyll cells of most vascular plants. It is loaded into phloem of mature leaves for long-distance translocation to non-photosynthetic organs where it is unloaded for diverse uses. Clearly, Suc transport and metabolism is central to plant growth and development and the functionality of the entire vascular system. Despite vast information in the literature about the physiological roles of individual sugar metabolic enzymes and transporters, there is a lack of systematic evaluation about their molecular regulation from transcriptional to post-translational levels. Knowledge on this topic is essential for understanding and improving plant development, optimizing resource distribution and increasing crop productivity. We therefore focused our analyses on molecular control of key players in Suc metabolism and transport, including: (i) the identification of promoter elements responsive to sugars and hormones or targeted by transcription factors and microRNAs degrading transcripts of target genes; and (ii) modulation of enzyme and transporter activities through protein-protein interactions and other post-translational modifications. We have highlighted major remaining questions and discussed opportunities to exploit current understanding to gain new insights into molecular control of carbon partitioning for improving plant performance.

Transcriptomic and proteomic approach to identify differentially expressed genes and proteins in Arabidopsis thaliana mutants lacking chloroplastic 1 and cytosolic FBPases reveals several levels of metabolic regulation.

BackgroundDuring the photosynthesis, two isoforms of the fructose-1,6-bisphosphatase (FBPase), the chloroplastidial (cFBP1) and the cytosolic (cyFBP), catalyse the first irreversible step during the conversion of triose phosphates (TP) to starch or sucrose, respectively. Deficiency in cyFBP and cFBP1 isoforms provokes an imbalance of the starch/sucrose ratio, causing a dramatic effect on plant development when the plastidial enzyme is lacking.ResultsWe study the correlation between the transcriptome and proteome profile in rosettes and roots when cFBP1 or cyFBP genes are disrupted in Arabidopsis thaliana knock-out mutants. By using a 70-mer oligonucleotide microarray representing the genome of Arabidopsis we were able to identify 1067 and 1243 genes whose expressions are altered in the rosettes and roots of the cfbp1 mutant respectively; whilst in rosettes and roots of cyfbp mutant 1068 and 1079 genes are being up- or down-regulated respectively. Quantitative real-time PCR validated 100% of a set of 14 selected genes differentially expressed according to our microarray analysis. Two-dimensional (2-D) gel electrophoresis-based proteomic analysis revealed quantitative differences in 36 and 26 proteins regulated in rosettes and roots of cfbp1, respectively, whereas the 18 and 48 others were regulated in rosettes and roots of cyfbp mutant, respectively. The genes differentially expressed and the proteins more or less abundant revealed changes in protein metabolism, RNA regulation, cell signalling and organization, carbon metabolism, redox regulation, and transport together with biotic and abiotic stress. Notably, a significant set (25%) of the proteins identified were also found to be regulated at a transcriptional level.ConclusionThis transcriptomic and proteomic analysis is the first comprehensive and comparative study of the gene/protein re-adjustment that occurs in photosynthetic and non-photosynthetic organs of Arabidopsis mutants lacking FBPase isoforms.Electronic supplementary materialThe online version of this article (doi:10.1186/s12870-016-0945-7) contains supplementary material, which is available to authorized users.

Identification and expression analysis of non-photosynthetic Rubisco small subunit, OsRbcS1-like genes in plants

Rubisco is the key enzyme of photosynthetic CO2 fixation, whereas a Rubisco small subunit homolog (OsRbcS1) in rice, was exclusively expressed in non-photosynthetic organs. In this study, OsRbcS1-like genes were identified in several plant species including foxtail millet (SiRbcS1), tomato (SlRbcS1), Lotus japonicus (LjRbcS1), grape (VvRbcS1A, VvRbcS1B) and Selaginella moellendorffii (SmRbcS1A, SmRbcS1B). RT-PCR analysis revealed that SiRbcS1 was expressed in seeds of foxtail millet. In tomato, SlRbcS1 was expressed in the stamen, pistil and green fruit. In L. japonicus, LjRbcS1 was expressed in the root, nodule and seed. In grape, both VvRbcS1A and VvRbcS1B were expressed in mature leaves and the green berry. In S. moellendorffii, SmRbcS1A and SmRbcS1B displayed similar expression patterns and were expressed in the rhizome and root. Although these OsRbcS1-like genes were expressed in photosynthetic organs in S. moellendorffii, its expression levels were higher in non-photosynthetic organs than in photosynthetic organs. These results suggest that OsRbcS1-like RbcS functions in metabolic pathways other than photosynthetic CO2 fixation in various organs.

The Interplay between Carbon Availability and Growth in Different Zones of the Growing Maize Leaf.

Plants assimilate carbon in their photosynthetic tissues in the light. However, carbon is required during the night and in nonphotosynthetic organs. It is therefore essential that plants manage their carbon resources spatially and temporally and coordinate growth with carbon availability. In growing maize (Zea mays) leaf blades, a defined developmental gradient facilitates analyses in the cell division, elongation, and mature zones. We investigated the responses of the metabolome and transcriptome and polysome loading, as a qualitative proxy for protein synthesis, at dusk, dawn, and 6, 14, and 24 h into an extended night, and tracked whole-leaf elongation over this time course. Starch and sugars are depleted by dawn in the mature zone, but only after an extension of the night in the elongation and division zones. Sucrose (Suc) recovers partially between 14 and 24 h into the extended night in the growth zones, but not the mature zone. The global metabolome and transcriptome track these zone-specific changes in Suc. Leaf elongation and polysome loading in the growth zones also remain high at dawn, decrease between 6 and 14 h into the extended night, and then partially recover, indicating that growth processes are determined by local carbon status. The level of Suc-signaling metabolite trehalose-6-phosphate, and the trehalose-6-phosphate:Suc ratio are much higher in growth than mature zones at dusk and dawn but fall in the extended night. Candidate genes were identified by searching for transcripts that show characteristic temporal response patterns or contrasting responses to carbon starvation in growth and mature zones.

Chlorophyll fluorescence induction kinetics and yield responses in rainfed crops with variable potassium nutrition in K deficient semi-arid alfisols

Optimum potassium (K) nutrition in semi-arid regions may help crop plants to overcome constraints in their growth and development such as moisture stress, leading to higher productivity of rainfed crops, thus judicious K management is essential. A study was conducted to evaluate the importance of K nutrition on physiological processes like photosynthesis through chlorophyll a fluorescence and chlorophyll fluorescence induction kinetics (OJIP) of rainfed crops viz., maize (Zea mays L.), pearl millet (Pennisetum glaucum), groundnut (Arachis hypogaea), sunflower (Helianthus annuus), castor (Ricinus communis L.) and cotton (Gossypium hirsutum) under water stress conditions by studying their growth attributes, water relations, yield, K uptake and use efficiency under varied K levels. Highest chlorophyll content was observed under K60 in maize and pearl millet. Narrow and wide Chl a:b ratio was observed in castor and groundnut respectively. The fluorescence yield decreased in the crops as K dosage increased, evidenced by increasing of all points (O, J, I and P) of the OJIP curves. The fluorescence transient curve for K60 was lower than K0 and K40 for all the crops. Potassium levels altered the fluorescence induction and impaired photosynthetic systems in all the crops studied. There was no distinct trend observed in leaf water potential of crops under study. Uptake of K was high in sunflower with increased rate of K application. Quantitatively, K uptake by castor crop was lesser compared to all other crops. Our results indicate that the yield reduction under low K was due to the low capacity of the crops to translocate K from non-photosynthetic organs such as stems and petioles to upper leaves and harvested organs and this in turn influenced the capacity of the crops to produce a high economic yield per unit of K taken up thus reducing utilization efficiency of K.

Nutrient allocation among plant organs across 13 tree species in three Bornean rain forests with contrasting nutrient availabilities.

Allocation of nitrogen (N) and phosphorus (P) among plant organs is an important factor regulating growth rate, which is a key ecological process associated with plant life-history strategies. However, few studies have explored how N and P investment in photosynthetic (leaves) and non-photosynthetic (stems and roots) organs changes in relation to depletion of each element. We investigated nutrient concentrations of plant organs in relation to whole-plant nutrient concentration (total nutrient weight per total biomass) as an index of nutrient status of each individual using the saplings of the 13 species in three tropical rain forests with contrasting N and P availabilities (tropical evergreen forests and tropical heath forests). We found a steeper decrease in foliar N concentration than foliar P concentration with decreasing whole-plant nutrient concentration. Moreover, the steeper decrease in foliar N concentration was associated with relatively stable N concentration in stems, and vice versa for P. We suggest that the depletion of N is associated with a rapid dilution of foliar N because the cell walls in non-photosynthetic organs function as an N sink. On the other hand, these species can maintain foliar P concentration by decreasing stem P concentrations despites the depletion of P. Our results emphasize the significance of non-photosynthetic organs as an N sink for understanding the variation of foliar nutrient concentrations for the tree species in the three Bornean rain forests with different N and P availabilities.

Divergent variations in concentrations of chemical elements among shrub organs in a temperate desert.

Desert shrubs, a dominant component of desert ecosystems, need to maintain sufficient levels of nutrients in their different organs to ensure operation of various physiological functions for the purpose of survival and reproduction. In the present study, we analyzed 10 elements in leaves, stems, and roots of 24 dominant shrub species from 52 sites across a temperate desert ecosystem in northwestern China. We found that concentrations of all 10 elements were higher in leaves than in stems and roots, that non-legumes had higher levels of leaf Na and Mg than did legumes, and that Na was more concentrated in C4 leaves than in C3 leaves. Scaling relationships of elements between the photosynthetic organ (leaf) and non-photosynthetic organs (stem and root) were allometric. Results of principal components analysis (PCA) highlighted the important role of the elements responsible for osmoregulation (K and Na) in water utilization of desert shrubs. Soil properties and taxonomy explained most variation of element concentrations in desert shrubs. Desert shrubs may not be particularly susceptible to future change in climate factors, because most elements (including N, P, K, Ca, Mn, Zn, and Cu) associated with photosynthesis, osmoregulation, enzyme activity, and water use efficiency primarily depend on soil conditions.

Nitrate metabolism in tobacco leaves overexpressing Arabidopsis nitrite reductase

Primary nitrogen assimilation in plants includes the reduction of nitrite to ammonium in the chloroplasts by the enzyme nitrite reductase (NiR EC:1.7.7.1) or in the plastids of non-photosynthetic organs. Here we report on a study overexpressing the Arabidopsis thaliana NiR (AtNiR) gene in tobacco plants under the control of a constitutive promoter (CERV – Carnation Etched Ring Virus). The aim was to overexpress AtNiR in an attempt to alter the level of residual nitrite in the leaf which can act as precursor to the formation of nitrosamines. The impact of increasing the activity of AtNiR produced an increase in leaf protein and a stay-green phenotype in the primary transformed AtNiR population. Investigation of the T1 homozygous population demonstrated elevated nitrate reductase (NR) activity, reductions in leaf nitrite and nitrate and the amino acids proline, glutamine and glutamate. Chlorophyl content of the transgenic lines was increased, as evidenced by the stay-green phenotype. This reveals the importance of NiR in primary nitrogen assimilation and how modification of this key enzyme affects both the nitrogen and carbon metabolism of tobacco plants.

Functions of OsDof25 in regulation of OsC4PPDK.

Relative little is known about the functions of the so-called Dof zinc factors in plants. Here we report on the analysis of OsDof25 and show a function in regulation of the important C4 photosynthesis gene, OsC4PPDK in rice. Over-expression of OsDof25 enhanced the expression of OsC4PPDK in transient expression experiments by binding in a specific way to a conserved Dof binding site which was confirmed by yeast and in vitro binding studies. Expression studies using promoter GUS plants as well as qPCR experiments showed that OsDof25 expressed in different tissues including both photosynthetic and non-photosynthetic organs and that expression of OsDof25 was partially overlapping with the OsC4PPDK gene. Conclusive evidence for a role of OsDof25 in regulation of C4PPDK came from loss-of-function and gain-of-function experiments with transgenic rice, which showed that down-regulation or over-expression of OsDof25 correlated with OsC4PPDK expression and that OsDof25 has functions as transcriptional activator.Electronic supplementary materialThe online version of this article (doi:10.1007/s11103-015-0357-3) contains supplementary material, which is available to authorized users.

Radiosynthesis of 6'-Deoxy-6'[18F]Fluorosucrose via Automated Synthesis and Its Utility to Study In Vivo Sucrose Transport in Maize (Zea mays) Leaves.

Sugars produced from photosynthesis in leaves are transported through the phloem tissues within veins and delivered to non-photosynthetic organs, such as roots, stems, flowers, and seeds, to support their growth and/or storage of carbohydrates. However, because the phloem is located internally within the veins, it is difficult to access and to study the dynamics of sugar transport. Radioactive tracers have been extensively used to study vascular transport in plants and have provided great insights into transport dynamics. To better study sucrose partitioning in vivo, a novel radioactive analog of sucrose was synthesized through a completely chemical synthesis route by substituting fluorine-18 (half-life 110 min) at the 6’ position to generate 6’-deoxy-6’[18F]fluorosucrose (18FS). This radiotracer was then used to compare sucrose transport between wild-type maize plants and mutant plants lacking the Sucrose transporter1 (Sut1) gene, which has been shown to function in sucrose phloem loading. Our results demonstrate that 18FS is transported in vivo, with the wild-type plants showing a greater rate of transport down the leaf blade than the sut1 mutant plants. A similar transport pattern was also observed for universally labeled [U-14C]sucrose ([U-14C]suc). Our findings support the proposed sucrose phloem loading function of the Sut1 gene in maize, and additionally demonstrate that the 18FS analog is a valuable, new tool that offers imaging advantages over [U-14C]suc for studying phloem transport in plants.

Plant zonation patterns reflected by the differences in plant growth, biomass partitioning and root traits along a water level gradient among four common vascular plants in freshwater marshes of the Sanjiang Plain, Northeast China

Water level is important in determining plant zonation distribution patterns in freshwater wetlands. However, the role of whole-plants’ morphological traits in response to water level gradients in accounting for the distribution of marsh species is far from clear. Four typical dominant vascular plants with different distribution patterns (high-elevation species Deyeuxia angustifolia, mid-elevation species Carex lasiocarpa and Glyceria spiculosa, and low-elevation species Carex pseudocuraica) in freshwater marshes in the Sanjiang Plain of Northeast China were treated with four static water level conditions (−5, 0, +5, and +15cm relative to the soil surface), and each plant’s growth characteristics, biomass allocation and root traits were investigated. With the rising water level, the growth of D. angustifolia performed worse than other species, whereas the total biomass, plant height and relative growth rate (RGR) of C. pseudocuraica increased steadily. Compared with the other three species, C. pseudocuraica showed a distinct pattern with increasing water levels in the biomass allocation to photosynthetic and non-photosynthetic organs, as well as for coarse and fine roots. Flooding stress significantly reduced root diameter and C/N ratio but simultaneously increased the specific root length (SRL), specific root area (SRA) and root nitrogen concentration (RNC) for both coarse roots (1st order, >0.2mm) and fine roots (2nd and 3rd orders, <0.2mm). However, these root parameters of C. pseudocuraica exhibited an opposite response with the increase in the water level between such two different branching orders. Our results indicate that wetland plants could acclimate to flooding stress through resource trade-off among different tissues and trait-coordinated variation within the root system. Furthermore, species-specific shifts in above- and below-ground traits could help to better understand and predict the vegetation zonation patterns in temperate freshwater wetlands.

Carotenoid content and root color of cultivated carrot: a candidate-gene association study using an original broad unstructured population.

Accumulated in large amounts in carrot, carotenoids are an important product quality attribute and therefore a major breeding trait. However, the knowledge of carotenoid accumulation genetic control in this root vegetable is still limited. In order to identify the genetic variants linked to this character, we performed an association mapping study with a candidate gene approach. We developed an original unstructured population with a broad genetic basis to avoid the pitfall of false positive detection due to population stratification. We genotyped 109 SNPs located in 17 candidate genes – mostly carotenoid biosynthesis genes – on 380 individuals, and tested the association with carotenoid contents and color components. Total carotenoids and β-carotene contents were significantly associated with genes zeaxanthin epoxydase (ZEP), phytoene desaturase (PDS) and carotenoid isomerase (CRTISO) while α-carotene was associated with CRTISO and plastid terminal oxidase (PTOX) genes. Color components were associated most significantly with ZEP. Our results suggest the involvement of the couple PDS/PTOX and ZEP in carotenoid accumulation, as the result of the metabolic and catabolic activities respectively. This study brings new insights in the understanding of the carotenoid pathway in non-photosynthetic organs.

Divisions of labor in the thiamin biosynthetic pathway among organs of maize.

The B vitamin thiamin is essential for central metabolism in all cellular organisms including plants. While plants synthesize thiamin de novo, organs vary widely in their capacities for thiamin synthesis. We use a transcriptomics approach to appraise the distribution of de novo synthesis and thiamin salvage pathways among organs of maize. We identify at least six developmental contexts in which metabolically active, non-photosynthetic organs exhibit low expression of one or both branches of the de novo thiamin biosynthetic pathway indicating a dependence on inter-cellular transport of thiamin and/or thiamin precursors. Neither the thiazole (THI4) nor pyrimidine (THIC) branches of the pathway are expressed in developing pollen implying a dependence on import of thiamin from surrounding floral and inflorescence organs. Consistent with that hypothesis, organs of the male inflorescence and flowers are shown to have high relative expression of the thiamin biosynthetic pathway and comparatively high thiamin contents. By contrast, divergent patterns of THIC and THI4 expression occur in the shoot apical meristem, embyro sac, embryo, endosperm, and root-tips suggesting that these sink organs acquire significant amounts of thiamin via salvage pathways. In the root and shoot meristems, expression of THIC in the absence of THI4 indicates a capacity for thiamin synthesis via salvage of thiazole, whereas the opposite pattern obtains in embryo and endosperm implying that seed storage organs are poised for pyrimidine salvage. Finally, stable isotope labeling experiments set an upper limit on the rate of de novo thiamin biosynthesis in maize leaf explants. Overall, the observed patterns of thiamin biosynthetic gene expression mirror the strategies for thiamin acquisition that have evolved in bacteria.

Reserves Accumulated in Non-Photosynthetic Organs during the Previous Growing Season Drive Plant Defenses and Growth in Aspen in the Subsequent Growing Season

Plants store non-structural carbohydrates (NSC), nitrogen (N), as well as other macro and micronutrients, in their stems and roots; the role of these stored reserves in plant growth and defense under herbivory pressure is poorly understood, particularly in trees. Trembling aspen (Populus tremuloides) seedlings with different NSC and N reserves accumulated during the previous growing season were generated in the greenhouse. Based on NSC and N contents, seedlings were assigned to one of three reserve statuses: Low N-Low NSC, High N-Medium NSC, or High N-High NSC. In the subsequent growing season, half of the seedlings in each reserve status was subjected to defoliation by forest tent caterpillar (Malacosoma disstria) while the other half was left untreated. Following defoliation, the effect of reserves was measured on foliar chemistry (N, NSC) and caterpillar performance (larval development). Due to their importance in herbivore feeding, we also quantified concentrations of phenolic glycoside compounds in foliage. Seedlings in Low N-Low NSC reserve status contained higher amounts of induced phenolic glycosides, grew little, and supported fewer caterpillars. In contrast, aspen seedlings in High N-Medium or High NSC reserve statuses contained lower amounts of induced phenolic glycosides, grew faster, and some of the caterpillars which fed on these seedlings developed up to their fourth instar. Furthermore, multiple regression analysis indicated that foliar phenolic glycoside concentration was related to reserve chemistry (NSC, N). Overall, these results demonstrate that reserves accumulated during the previous growing season can influence tree defense and growth in the subsequent growing season. Additionally, our study concluded that the NSC/N ratio of reserves in the previous growing season represents a better measure of resources available for use in defense and growth than the foliar NSC/N ratios.