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Leaf hydraulic conductance in relation to anatomical and functional traits during Populus tremula leaf ontogeny.

Leaf hydraulic conductance (K(leaf)) and several characteristics of hydraulic architecture and physiology were measured during the first 10 weeks of leaf ontogeny in Populus tremula L. saplings growing under control, mild water deficit or elevated temperature conditions. During the initial 3 weeks of leaf ontogeny, most measured characteristics rapidly increased. Thereafter, a gradual decrease in K(leaf) was correlated with a decrease in leaf osmotic potential under all conditions, and with increases in leaf dry mass per area and bulk modulus of elasticity under mild water deficit and control conditions. From about Week 3 onward, K(leaf) was 33% lower in trees subjected to mild water deficit and 33% higher in trees held at an elevated temperature relative to control trees. Mild water deficit and elevated temperature treatment had significant and opposite effects on most of the other characteristics measured. The ontogenetic maximum in K(leaf) was correlated positively with the width of xylem conduits in the midrib, but negatively with the overall width of the midrib xylem, number of lateral ribs, leaf dry mass per area and bulk modulus of elasticity. The ontogenetic maximum in K(leaf) was also correlated positively with the proportion of intercellular spaces and leaf osmotic potential, but negatively with leaf thickness, volume of mesophyll cells and epidermis and number of cells per total mesophyll cell volume, the closest relationships being between leaf osmotic potential and number of cells per total mesophyll cell volume. It was concluded that differences in protoplast traits are more important than differences in xylem or parenchymal cell wall traits in determining the variability in K(leaf) among leaves growing under different environmental conditions.

Plant Leaves↗

Effects of two iron sources on iron and cadmium allocation in poplar (Populus alba) plants exposed to cadmium.

Effects of 10 microM cadmium (supplied as Cd nitrate) on the utilization and allocation of iron (Fe) were investigated in poplar (Populus alba L.) plants grown in nutrient solution with Fe(III)-EDTA or Fe(III)-citrate as the Fe source. The effects of Cd were also compared with those of Fe deprivation. The accumulation of Fe in roots was 10-fold higher in plants grown with Fe-citrate than with Fe-EDTA. Cadmium decreased leaf chlorophyll concentrations and photosynthetic rates, and these decreases were more marked in plants grown with Fe-citrate than with Fe-EDTA. In both Fe treatments, addition of Cd caused large increases in root and shoot apoplasmic and non-apoplasmic Cd contents and increases in root Fe content; however, Cd decreased shoot Fe content, especially in plants grown with Fe-citrate. New leaves of plants grown with Fe-citrate had small cellular (non-apoplasmic) Fe pools, whereas these pools were large in new leaves of plants grown with Fe-EDTA. Non-apoplasmic Cd pools in new leaves were smaller in plants grown with Fe-citrate than with Fe-EDTA, indicating that inactivation of non-apoplasmic Cd pools is facilitated more by Fe-EDTA than by Fe-citrate. In the presence of Cd, Fe-EDTA was also superior to Fe-citrate in maintaining an adequate Fe supply to poplar shoots. Differences in plant responses to Fe-EDTA and Fe-citrate may reflect differences in long-distance transport of Fe rather than in acquisition of Fe by roots.

Cadmium↗

Signals controlling root suckering and adventitious shoot formation in aspen (Populus tremuloides).

We determined the effects of removal of leaves, stem axillary buds, or the entire shoot on root suckering (adventitious shoot formation by roots) and basal stem sprouts in 3- and 4-year-old potted seedlings of aspen (Populus tremuloides Michx.). The greatest number of root suckers (67.9 +/- 8.5 per plant) emerged after excision of the entire shoot. Defoliated and debudded stems were the major source of inhibitory agents for root suckering, although axillary buds and developing new leaves also exerted a significant inhibitory effect. Removal of mature leaves had only a minor effect on root suckering. Removal of a continuous band of bark (girdling) at the base of the stem consistently stimulated growth of adventitious shoots from the stem below the girdle and occasionally promoted root suckering. Exogenous application of indole-3-acetic acid to excised stumps inhibited root suckering and basal stem sprouting. Naphthylphthalamic acid (NPA), an auxin polar transport inhibitor, had no effect on root suckering or stem sprouting when it was applied to the bark of the basal stem. However, NPA significantly increased root suckering when it was applied to the exposed surface of xylem after girdling. These results suggest that polar transport of auxin in the xylem parenchyma is an important inhibitor of root suckering. On decapitated stems, vacuum extraction of xylem sap from the root system lowered the frequency of root suckering compared with decapitation alone, indicating that substance(s) originating in the root system also play a significant role in controlling root suckering.

Indoleacetic Acids↗

Terminal bud failure of black cottonwood (Populus trichocarpa) exposed to salt-laden winter storms.

At coastal sites, trees are exposed to marine aerosols that may cause foliar necrosis and shoot dieback, which can result in deformed crowns and contorted stems. A six-year study of leaf primordia in terminal buds of black cottonwood trees (Populus trichocarpa Torr. & Gray) on Heimaey Island off the south coast of Iceland was undertaken to elucidate the physiological events associated with salt-deposition-related bud failure. Leaf and bud lengths, dry mass, water content and chloride concentrations were monitored and related to four phenological stages: (1) bud set; (2) dormancy induction; (3) dormancy release; and (4) bud break. The trees set buds in July and shed their leaves by late September. Leaf primordia generally stopped growing by September 10 +/- 22 days and attained midwinter water content in late September. Leaf growth commenced in the terminal buds by March 2 +/- 16 days, but mean dates of bud swelling and bud break were April 29 +/- 19 and May 10 +/- 12 days. In summer and until November, chloride concentrations in leaf primordia were low, but increasing. Chloride concentrations remained stable from December to February, even though the dormant trees were exposed to large amounts of marine aerosols. In February and March, three events occurred more or less simultaneously: (1) leaf extension growth commenced; (2) chloride concentration surged in the leaf primordia; and (3) the leaf primordia began to hydrate. Following dormancy release, growth and hydration of leaf primordia were negatively related to chloride concentration in the leaf primordia, with inhibition of leaf growth, tissue hydration and chloride acquisition occurring at a chloride concentration threshold estimated at 7.3 mg Cl- g(-1) tissue water. Necrosis of leaf primordia was observed above 14 mg Cl- g(-1) tissue water. Growth and hydration of leaves at bud break in mid-May was explained by a three-parameter logistic model of chloride concentration in leaf primordia at the end of March. By mid-May, 90% of all buds remained non-necrotic, but only 56% the terminal buds had broken. Salt alone explained the observed growth suppression of leaf primordia in the buds and the resultant failure of terminal buds to break by mid-May.

Geography↗

Methylobacterium populi sp. nov., a novel aerobic, pink-pigmented, facultatively methylotrophic, methane-utilizing bacterium isolated from poplar trees (Populus deltoides x nigra DN34).

A pink-pigmented, aerobic, facultatively methylotrophic bacterium, strain BJ001T, was isolated from internal poplar tissues (Populus deltoidesxnigra DN34) and identified as a member of the genus Methylobacterium. Phylogenetic analyses showed that strain BJ001T is related to Methylobacterium thiocyanatum, Methylobacterium extorquens, Methylobacterium zatmanii and Methylobacterium rhodesianum. However, strain BJ001T differed from these species in its carbon-source utilization pattern, particularly its use of methane as the sole source of carbon and energy, an ability that is shared with only one other member of the genus, Methylobacterium organophilum. In addition, strain BJ001T is the only member of the genus Methylobacterium to be described as an endophyte of poplar trees. On the basis of its physiological, genotypic and ecological properties, the isolate is proposed as a member of a novel species of the genus Methylobacterium, Methylobacterium populi sp. nov. (type strain, BJ001T=ATCC BAA-705T=NCIMB 13946T).

Aerobiosis↗

Populus euphratica displays apoplastic sodium accumulation, osmotic adjustment by decreases in calcium and soluble carbohydrates, and develops leaf succulence under salt stress.

Populus euphratica Olivier is known to exist in saline and arid environments. In this study we investigated the physiological mechanisms enabling this species to cope with stress caused by salinity. Acclimation to increasing Na+ concentrations required adjustments of the osmotic pressure of leaves, which were achieved by accumulation of Na+ and compensatory decreases in calcium and soluble carbohydrates. The counterbalance of Na+/Ca2+ was also observed in mature leaves from field-grown P. euphratica trees exposed to an environmental gradient of increasing salinity. X-ray microanalysis showed that a primary strategy to protect the cytosol against sodium toxicity was apoplastic but not vacuolar salt accumulation. The ability to cope with salinity also included maintenance of cytosolic potassium concentrations and development of leaf succulence due to an increase in cell number and cell volume leading to sodium dilution. Decreases in apoplastic and vacuolar Ca2+ combined with suppression of calcineurin B-like protein transcripts suggest that Na+ adaptation required suppression of calcium-related signaling pathways. Significant increases in galactinol synthase and alternative oxidase after salt shock and salt adaptation point to shifts in carbohydrate metabolism and suppression of reactive oxygen species in mitochondria under salt stress.

Adaptation, Physiological↗

The cellulose synthase gene superfamily and biochemical functions of xylem-specific cellulose synthase-like genes in Populus trichocarpa.

Wood from forest trees modified for more cellulose or hemicelluloses could be a major feedstock for fuel ethanol. Xylan and glucomannan are the two major hemicelluloses in wood of angiosperms. However, little is known about the genes and gene products involved in the synthesis of these wood polysaccharides. Using Populus trichocarpa as a model angiosperm tree, we report here a systematic analysis in various tissues of the absolute transcript copy numbers of cellulose synthase superfamily genes, the cellulose synthase (CesA) and the hemicellulose-related cellulose synthase-like (Csl) genes. Candidate Csl genes were characterized for biochemical functions in Drosophila Schneider 2 (S2) cells. Of the 48 identified members, 37 were found expressed in various tissues. Seven CesA genes are xylem specific, suggesting gene networks for the synthesis of wood cellulose. Four Csl genes are xylem specific, three of which belong to the CslA subfamily. The more xylem-specific CslA subfamily is represented by three types of members: PtCslA1, PtCslA3, and PtCslA5. They share high sequence homology, but their recombinant proteins produced by the S2 cells exhibited distinct substrate specificity. PtCslA5 had no catalytic activity with the substrates for xylan or glucomannan. PtCslA1 and PtCslA3 encoded mannan synthases, but PtCslA1 further encoded a glucomannan synthase for the synthesis of (1-->4)-beta-D-glucomannan. The expression of PtCslA1 is most highly xylem specific, suggesting a key role for it in the synthesis of wood glucomannan. The results may help guide further studies to learn about the regulation of cellulose and hemicellulose synthesis in wood.

Animals↗

Gradual soil water depletion results in reversible changes of gene expression, protein profiles, ecophysiology, and growth performance in Populus euphratica, a poplar growing in arid regions.

The responses of Populus euphratica Oliv. plants to soil water deficit were assessed by analyzing gene expression, protein profiles, and several plant performance criteria to understand the acclimation of plants to soil water deficit. Young, vegetatively propagated plants originating from an arid, saline field site were submitted to a gradually increasing water deficit for 4 weeks in a greenhouse and were allowed to recover for 10 d after full reirrigation. Time-dependent changes and intensity of the perturbations induced in shoot and root growth, xylem anatomy, gas exchange, and water status were recorded. The expression profiles of approximately 6,340 genes and of proteins and metabolites (pigments, soluble carbohydrates, and oxidative compounds) were also recorded in mature leaves and in roots (gene expression only) at four stress levels and after recovery. Drought successively induced shoot growth cessation, stomatal closure, moderate increases in oxidative stress-related compounds, loss of CO2 assimilation, and root growth reduction. These effects were almost fully reversible, indicating that acclimation was dominant over injury. The physiological responses were paralleled by fully reversible transcriptional changes, including only 1.5% of the genes on the array. Protein profiles displayed greater changes than transcript levels. Among the identified proteins for which expressed sequence tags were present on the array, no correlation was found between transcript and protein abundance. Acclimation to water deficit involves the regulation of different networks of genes in roots and shoots. Such diverse requirements for protecting and maintaining the function of different plant organs may render plant engineering or breeding toward improved drought tolerance more complex than previously anticipated.

Climate↗

Structural and kinetic basis for substrate selectivity in Populus tremuloides sinapyl alcohol dehydrogenase.

We describe the three-dimensional structure of sinapyl alcohol dehydrogenase (SAD) from Populus tremuloides (aspen), a member of the NADP(H)-dependent dehydrogenase family that catalyzes the last reductive step in the formation of monolignols. The active site topology revealed by the crystal structure substantiates kinetic results indicating that SAD maintains highest specificity for the substrate sinapaldehyde. We also report substantial substrate inhibition kinetics for the SAD-catalyzed reduction of hydroxycinnamaldehydes. Although SAD and classical cinnamyl alcohol dehydrogenases (CADs) catalyze the same reaction and share some sequence identity, the active site topology of SAD is strikingly different from that predicted for classical CADs. Kinetic analyses of wild-type SAD and several active site mutants demonstrate the complexity of defining determinants of substrate specificity in these enzymes. These results, along with a phylogenetic analysis, support the inclusion of SAD in a plant alcohol dehydrogenase subfamily that includes cinnamaldehyde and benzaldehyde dehydrogenases. We used the SAD three-dimensional structure to model several of these SAD-like enzymes, and although their active site topologies largely mirror that of SAD, we describe a correlation between substrate specificity and amino acid substitution patterns in their active sites. The SAD structure thus provides a framework for understanding substrate specificity in this family of enzymes and for engineering new enzyme specificities.

Acrolein↗

The genetics and genomics of the drought response in Populus.

The genetic nature of tree adaptation to drought stress was examined by utilizing variation in the drought response of a full-sib second generation (F(2)) mapping population from a cross between Populus trichocarpa (93-968) and P. deltoides Bart (ILL-129) and known to be highly divergent for a vast range of phenotypic traits. We combined phenotyping, quantitative trait loci (QTL) analysis and microarray experiments to demonstrate that 'genetical genomics' can be used to provide information on adaptation at the species level. The grandparents and F(2) population were subjected to soil drying, and contrasting responses to drought across genotypes, including leaf coloration, expansion and abscission, were observed, and QTL for these traits mapped. A subset of extreme genotypes exhibiting extreme sensitivity and insensitivity to drought on the basis of leaf abscission were defined, and microarray experiments conducted on these genotypes and the grandparent species. The extreme genotype groups induced a different set of genes: 215 and 125 genes differed in their expression response between groups in control and drought, respectively, suggesting species adaptation at the gene expression level. Co-location of differentially expressed genes with drought-specific and drought-responsive QTLs was examined, and these may represent candidate genes contributing to the variation in drought response.

Disasters↗

Productivity, leaf traits and carbon isotope discrimination in 29 Populus deltoides x P. nigra clones.

Here we tested whether some leaf traits could be used as predictors for productivity in a range of Populus deltoides x P. nigra clones. These traits were assessed in 3-yr-old rooted cuttings from 29 clones growing in an open field trial, in a five randomized complete block design, under optimal irrigation. Variables were assigned to four groups describing productivity (above-ground biomass, total leaf area), leaf growth (total number of leaves increment rate), leaf structure (area of the largest leaf, specific leaf area, carbon and nitrogen contents), and carbon isotope discrimination in the leaves (Delta). High-yielding clones displayed larger total leaf area and individual leaf area, while no correlation could be detected between productivity and either leaf structure or Delta. By contrast, Delta was negatively correlated with number of leaves increment rate and leaf N content. Our study shows that there is a potential to improve water-use efficiency in poplar without necessarily reducing the overall productivity.

Carbon Isotopes↗

The transcriptome of Populus in elevated CO2.

The consequences of increasing atmospheric carbon dioxide for long-term adaptation of forest ecosystems remain uncertain, with virtually no studies undertaken at the genetic level. A global analysis using cDNA microarrays was conducted following 6 yr exposure of Populus x euramericana (clone I-214) to elevated [CO(2)] in a FACE (free-air CO(2) enrichment) experiment. Gene expression was sensitive to elevated [CO(2)] but the response depended on the developmental age of the leaves, and < 50 transcripts differed significantly between different CO(2) environments. For young leaves most differentially expressed genes were upregulated in elevated [CO(2)], while in semimature leaves most were downregulated in elevated [CO(2)]. For transcripts related only to the small subunit of Rubisco, upregulation in LPI 3 and downregulation in LPI 6 leaves in elevated CO(2) was confirmed by anova. Similar patterns of gene expression for young leaves were also confirmed independently across year 3 and year 6 microarray data, and using real-time RT-PCR. This study provides the first clues to the long-term genetic expression changes that may occur during long-term plant response to elevated CO(2).

Adaptation, Physiological↗

Competition- and resource-mediated tradeoffs between growth and defensive chemistry in trembling aspen (Populus tremuloides).

Costs of defense are thought to maintain genetic variations in the expression of defense within plant populations. As with many plant species, aspen exhibits considerable variation in allocation to secondary metabolites. This study examined the independent and interactive effects of genotype, soil fertility and belowground competition on defensive chemistry and growth in trembling aspen (Populus tremuloides). Four aspen genotypes were grown with high and low soil fertility, and with and without root competition. Physiological, morphological and allocational determinants of growth were measured to identify growth-defense tradeoffs. Nutrient limitation and competition decreased growth, leaf mass ratio, leaf nitrogen concentration and photosynthesis, and increased root : shoot ratio and leaf condensed tannin concentrations. The competition treatment also resulted in increased leaf phenolic glycoside (PG) concentrations. Aspen growth was negatively correlated with PG concentrations under low fertility with competition. The relationship between growth and its major determinants was also negatively related to foliar condensed tannins expressed as a proportion of tree mass, indicating an additional indirect cost of allocation to secondary metabolites.

Biomass↗

Developmental and seasonal expression of PtaHB1, a Populus gene encoding a class III HD-Zip protein, is closely associated with secondary growth and inversely correlated with the level of microRNA (miR166).

In contrast to our knowledge of the shoot apical meristem, our understanding of cambium meristem differentiation and maintenance is limited. Class III homeodomain leucine-zipper (HD-Zip) proteins have been shown to play a regulatory role in vascular differentiation. The hybrid aspen (Populus tremulaxPopulus alba) class III HD-Zip transcription factor (PtaHB1) and microRNA 166 (Pta-miR166) family were cloned from hybrid aspen using a combination of in silico and polymerase chain reaction methods. Expression analyses of PtaHB1 and Pta-miR166 were performed by Northern blot analysis. The expression of PtaHB1 was closely associated with wood formation and regulated both developmentally and seasonally, with the highest expression during the active growing season. Also, its expression was inversely correlated with the level of Pta-miR166. Pta-miR166-directed cleavage of PtaHB1 in vivo was confirmed using modified 5'-rapid amplification of cDNA ends (RACE). The expression of Pta-miR166 was much higher in the winter than in the growing seasons, suggesting seasonal and developmental regulation of microRNA in this perennial plant species.

Amino Acid Sequence↗

Nitrogen uptake and metabolism in Populus x canescens as affected by salinity.

External salinization can affect different steps of nitrogen (N) metabolism (ion uptake, N assimilation, and amino acid and protein synthesis) depending on the inorganic N source. Here, we assessed the net uptake of N supplied as nitrate or ammonium and N assimilation (combining metabolite analyses with molecular biological approaches) in grey poplar (Populus x canescens) plants grown under saline (75 mM NaCl) and control conditions. The specific (micromol N g(-1) dry weight fine roots h(-1)) and total plant (micromol N per plant h(-1)) N net uptake rates, total plant N content, total plant biomass and total leaf protein concentration were reduced under saline conditions when plants were supplied with ammonium. In both nutritional groups, salt treatment caused pronounced accumulation of soluble N compounds in the leaves. The mRNAs of genes coding for enzymes catalyzing rate-limiting steps of both proline synthesis and degradation (delta-1-pyrroline-5-carboxylate synthase and proline dehydrogenase) as well as for NADH-dependent glutamate synthase were accumulated under saline conditions. Whereas under control conditions the plant N status seemed to be superior when ammonium was supplied, the N balance of ammonium-fed plants was more severely affected by salt stress than that of plants supplied with nitrate. Possible metabolic implications of stress-related accumulation of particular amino acids are discussed.

Biomass↗

Biogenic volatile organic compounds as potential carbon sources for microbial communities in soil from the rhizosphere of Populus tremula.

Catabolism of a (14)C-labelled volatile monoterpene compound (geraniol) to (14)CO(2) was investigated in soils taken from the rhizosphere at distances up to 200 cm from the trunks of three small Populus tremula trees growing at different sites in Slovenia. Emissions of limonene of up to 18 microg m(-2) h(-1) were detected from the soil surface at each site. Evolution of (14)C-labelled CO(2) was measured as a product of catabolism of (14)C-labelled geraniol introduced into the soil samples. Indigenous soil microorganisms degraded the geraniol rapidly. There was a significant difference in relative lag times and rates of catabolism along the gradient from the tree trunks, with relatively longer lag times and lower rates occurring in soil samples from the farthest point from the tree.

Acyclic Monoterpenes↗

Dynamic Rhizodeposition in the Woody Perennial Populus trichocarpa.

Plants undergo physiological and metabolic changes that release specific molecules into the surrounding soil, a process collectively known as rhizodeposition. These compounds play crucial roles in plant-microbe-soil interactions, such as supporting plant development and resilience in changing environments. Under nutrient-limited conditions, these plant-derived compounds modify the rhizosphere environment, mobilizing otherwise inaccessible nutrients and recruiting stress-adaptive microbial communities that support stress resilience. Currently, the chemical diversity of rhizodeposition has yet to be fully realized but is expected to be a complex mixture that includes soluble organic compounds excreted from root cells, along with products of root cell turnover, sloughed-off root cap and border cells, and mucilage. Here, we developed a methodological and conceptual framework for an in-depth measurement of rhizodeposition through critical advancements in untargeted metabolomics. This approach provided foundational insights into the dynamic changes in rhizodeposition for the woody perennial Populus trichocarpa and rhizodeposit profiles varying by genotype, time, location, and environment. More broadly, this study provides a framework that will help formulate the next steps to effectively study rhizodeposition.

Populus↗

Haplotype-resolved telomere-to-telomere genome assembly of Populus lasiocarpa unveils retrotransposon-driven centromere evolution.

Centromeres, essential for chromosome segregation, exhibit remarkable evolutionary dynamism in sequence composition and structural organization. Here, we report the first haplotype-resolved, telomere-to-telomere genome assembly of Populus lasiocarpa (PLAS) and precisely map all 38 functional centromeres through CENH3 ChIP-Seq. Unlike classical satellite-rich centromeres in model plants, PLAS centromeres lack abundant satellite arrays but are dominated by retrotransposons, particularly RLG and RIL elements, which form intricate nested TE arrays within the functional centromeric regions, disrupting their structural integrity and driving their evolution. Comparative analysis with P. trichocarpa reveals a conserved retrotransposon-dominated architecture, despite minimal sequence conservation. We propose a cyclic model of centromere evolution in which autonomous retrotransposons destabilize functional centromeres through epigenetic erosion, triggering neocentromere formation at pericentromeric sites enriched in transposable elements (TEs) and tandem repeats (TRs). These neocentromeres either succumb to recurrent retrotransposon invasions or stabilize through KARMA-mediated TR expansion, ultimately giving rise to satellite-rich centromeres. Our work redefines centromeres as dynamic, epigenetically plastic domains shaped by retrotransposon-TR antagonism, challenging the satellite-centric paradigm and offering novel insights into plant genome evolution.

Retroelements↗