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PubMed2002

Genomic DNA methylation-demethylation during aging and reinvigoration of Pinus radiata.

In animals, DNA methylation is related to gene silencing during ontogenic development. Little is known about DNA methylation in plants, although occasional changes in the DNA methylation state of specific gene promoters have been reported in angiosperms during some developmental processes. We found large differences in the extent of DNA methylation between meristematic areas of juvenile and mature Pinus radiata D. Don. trees, whereas differences in the extent of DNA methylation between differentiated tissues of juvenile and mature trees were small. In meristematic areas, there was a gradual decrease in extent of DNA methylation as the degree of reinvigoration increased. The observed changes in extent of DNA methylation during aging and reinvigoration indicate that reinvigoration could be a consequence of epigenetic modifications opposite in direction to those that occur during aging.

DNA Methylation↗

PubMed2002

Characterization of a Pinus pinaster cDNA encoding an auxin up-regulated putative peroxidase in roots.

As part of a study to identify host plant genes regulated by fungal auxin during ectomycorrhiza formation, we differentially screened a cDNA library constructed from roots of auxin-treated Pinus pinaster (Ait.) Sol. seedlings. We identified three cDNAs up-regulated by auxin. Sequence analysis of one of these cDNAs, PpPrx75, revealed the presence of an open reading frame of 216 amino acids with the characteristic consensus sequences of plant peroxidases. The deduced amino acid sequence showed homology with Arabidopsis thaliana (L.) Heynh., Arachis hypogaea L. and Stylosanthes humilis HBK cationic peroxidases. Amino acid sequence identities in the conserved domains of plant peroxidases ranged from 60 to 100%. In PpPrx75, there are five cysteine residues and one histidine residue that are found at conserved positions among other peroxidases. A potential glycosylation site (NTS) is present in the deduced sequence. Phylogenetic analysis showed that PpPrx75 is closely related to two A. thaliana peroxidases. The PpPrx75 cDNA was induced by active auxins, ethylene, abscisic acid and quercetin, a flavonoid possibly involved in plant-microorganism interactions. Transcript accumulation was detected within 3 h following root induction by auxin, and the amount of mRNA increased over the following 24 h. The protein synthesis inhibitor cycloheximide did not inhibit indole-3-acetic acid-induced transcript accumulation, suggesting that PpPrx75 induction is a primary (direct) response to auxin. This cDNA can be used to study expression of an auxin-regulated peroxidase during ectomycorrhiza formation.

Amino Acid Sequence↗

PubMed2003

Sapwood development in Pinus radiata trees grown for three years at ambient and elevated carbon dioxide partial pressures.

Clonal trees of Pinus radiata D. Don were grown in open-top chambers at a field site in New Zealand for 3 years at ambient (37 Pa) or elevated (65 Pa) carbon dioxide (CO2) partial pressure. Nitrogen (N) was supplied to half of the trees in each CO2 treatment, at 15 g N m-2 in the first year and 60 g N m-2 in the subsequent 2 years (high-N treatment). Trees in the low-N treatment were not supplied with N but received the same amount of other nutrients as trees in the high-N treatment. In the first year, stem basal area increased more in trees growing at elevated CO2 partial pressure and high-N supply than in control trees, suggesting a positive interaction between these resources. However, the relative rate of growth became the same across trees in all treatments after 450 days, resulting in trees growing at elevated CO2 partial pressure and high-N supply having larger basal areas than trees in the other treatments. Sapwood N content per unit dry mass was consistently about 0.09% in all treatments, indicating that N status was not suppressed by elevated CO2 partial pressure. Thus, during the first year of growth, an elevated CO2 partial pressure enhanced carbon (C) and N storage in woody stems, but there was no further stimulus to C and N deposition after the first year. The chemical composition of sapwood was unaffected by elevated CO2 partial pressure, indicating that no additional C was sequestered through lignification. However, independent of the treatments, early wood was 13% richer in lignin than late wood. Elevated CO2 partial pressure decreased the proportion of sapwood occupied by the lumina of tracheids by up to 12%, indicating increased sapwood density in response to CO2 enrichment. This effect was probably a result of thicker tracheid walls rather than narrower lumina.

Carbon Dioxide↗

PubMed2003

Horizontal and vertical variations in photosynthetic capacity in a Pinus densiflora crown in relation to leaf nitrogen allocation and acclimation to irradiance.

We measured horizontal and vertical gradients of light (rPPFD) along four first-order branches of a Pinus densiflora Sieb. & Zucc. crown, and compared variations in specific leaf area (SLA), needle nitrogen concentration (N), chlorophyll concentration (Chl) and photosynthetic capacity (i.e., maximum rate of carboxylation (V(cmax))) along the two axes. The horizontal gradient of rPPFD along first-order branches was similar in magnitude to the vertical gradient of rPPFD from the upper to the lower crown. None of the measured parameters (i.e., SLA, N, Chl and Vcmax) were strictly proportional to rPPFD, although they were more or less correlated with light when data obtained for all of the crown were pooled (r(2) = 0.31-0.80). The slope of rPPFD against N on an area basis (Narea) for a branch in the middle of the crown orientated northward was significantly greater than the slope for a similar branch orientated southward. Horizontal variations were unrelated to age effects because measurements were all on 1-year-old needles. We conclude that factors other than light (i.e., orientation) may influence N allocation within branches. There was considerably less variation in the relationship of Vcmax to Narea (r2 = 0.58) than in the relationship of Vcmax to rPPFD (r2 = 0.41). Fractional N distribution among components of the photosynthetic machinery was constant within the crown. Together with the relationships between rPPFD and N on a mass basis (r2 = 0.80) and SLA and Vcmax (r2 = 0.60), these findings suggest that most light acclimation in P. densiflora occurs through changes in needle morphology (e.g., SLA) during development.

Acclimatization↗

PubMed2003

Wood properties of Scots pines (Pinus sylvestris) grown at elevated temperature and carbon dioxide concentration.

Impacts of elevated temperature and carbon dioxide concentration ([CO2]) on wood properties of 15-year-old Scots pines (Pinus sylvestris L.) grown under conditions of low nitrogen supply were investigated in open-top chambers. The treatments consisted of (i) ambient temperature and ambient [CO2] (AT+AC), (ii) ambient temperature and elevated [CO2] (AT+EC), (iii) elevated temperature and ambient [CO2] (ET+AC) and (iv) elevated temperature and elevated [CO2] (ET+EC). Wood properties analyzed for the years 1992-1994 included ring width, early- and latewood width and their proportions, intra-ring wood density (minimum, maximum and mean, as well as early- and latewood densities), mean fiber length and chemical composition of the wood (cellulose, hemicellulose, lignin and acetone extractive concentration). Absolute radial growth over the 3-year period was 54% greater in AT+EC trees and 30 and 25% greater in ET+AC and ET+EC trees, respectively, than in AT+AC trees. Neither elevated temperature nor elevated [CO2] had a statistically significant effect on ring width, early- and latewood widths or their proportions. Both latewood density and maximum intra-ring density were increased by elevated [CO2], whereas fiber length was increased by elevated temperature. Hemicellulose concentration decreased and lignin concentration increased significantly in response to elevated temperature. There were no statistically significant interaction effects of elevated temperature and elevated [CO2] on the wood properties, except on earlywood density.

Carbon Dioxide↗

PubMed2003

Aux/IAA gene family is conserved in the gymnosperm, loblolly pine (Pinus taeda).

We isolated five members of the Aux/IAA gene family in loblolly pine (Pinus taeda L.). Degenerate primers complementary to conserved regions of angiosperm Aux/IAA genes were used to amplify fragments that were, in turn, used as probes to screen a cDNA library constructed from auxin-treated hypocotyls. The five unique clones, named PTIAA1-5, contain the four highly conserved domains that are characteristic of the Aux/IAA proteins. All clones contain the bipartite nuclear localization signal (NLS) between Domains I and II that is predicted in most angiosperm Aux/IAA genes, but only one, PTIAA2, contains the conserved NLS in Domain IV. The five invariant residues in Domain II that have been found to constitute part of a protein destabilization element in Arabidopsis thaliana (L.) Heynh. are conserved in all the PTIAAs. A postulated phosphorylation site located between Domains I and II and proximal to the conserved bipartite NLS was conserved in 20 out of 36 genes in this analysis, including the pine genes. Transcripts of all five PTIAAs accumulated specifically in the hypocotyls in response to exogenous auxin treatment and were induced by all auxins tested. Transcript abundance above basal levels in response to 1-naphthaleneacetic acid treatment was first detected after 10 min (PTIAA3) to 3 h (PTIAA2) in the different genes and remained above basal levels throughout 7 days. Induction of PTIAA2 was inhibited by the protein synthesis inhibitor cycloheximide, indicating that PTIAA2 is a secondary response gene. Phylogenetic analysis showed that all five pine genes clustered within a single class (Class I) of the dendrogram. Clone PTIAA2 has a sequence that is relatively distinct from the other four and is the most closely related to the angiosperm genes of Class I. Class I contains both primary and secondary auxin response genes, suggesting that it is the original lineage and that other gene classes have evolved subsequent to the angiosperm/gymnosperm divergence.

Amino Acid Sequence↗

PubMed2003

Reinvigoration of Pinus radiata is associated with partial recovery of juvenile-like polyamine concentrations.

Polyamine concentrations in consecutive radiata pine (Pinus radiata D. Don) grafts of 30-year-old trees on 1-year-old seedling rootstocks were analyzed by high performance liquid chromatography coupled with fluorescence detection to determine whether reinvigoration is associated with the recovery of polyamine concentrations typical of those in juvenile tissues. Reinvigoration of radiata pine was correlated with the attainment of some, but not all, polyamines characteristic of juvenile trees. In response to reinvigoration, free putrescine, the ratio of free polyamines to low molecular weight polyamine conjugates, and the relative content of putrescine versus spermidine plus spermine increased to values approaching those characteristic of juvenile tissue. In contrast, there was no noticeable change in the concentrations of low molecular weight polyamine conjugates during reinvigoration even though these conjugates increased greatly during tree maturation. We conclude that certain polyamines defined as reinvigoration markers can be used in forestry upgrading programs to assess the morphogenic ability of reinvigorated trees.

Biogenic Polyamines↗

PubMed2003

Variability of stem and branch maintenance respiration in a Pinus pinaster tree.

The relationship between maintenance respiration (Rm) of woody organs and their structural characteristics was explored in adult Pinus pinaster Ait. trees. We measured Rm on 75 stem and branch segments of different ages (from 3 to 24 years) and diameters (from 1 to 35 cm). The temperature response of Rm was derived from field measurements based on a classical exponential function with Q10 = 2.13. Relationships between Rm and the dimensions of the woody organs were analyzed under controlled conditions in the laboratory. The surface area of a woody organ was a better predictor of Rm than volume, but surface area failed to account for the observed within-tree variability of Rm among stems, branches and twigs. Two simple models were proposed to predict the variability of Rm at 15 degrees C in an adult tree. Model 1, a linear function model based on the dry mass and nitrogen concentration of sapwood and phloem tissues, explained most of the variability of Rm in branches and stems (R2 = 0.97). We concluded that the respective contributions of the phloem and sapwood depend on the location and diameter of the woody organ. Model 2, a power-law function model based on the length, diameter and age of the sample, explained the same variance of Rm as Model 1 and is appropriate for scaling Rm to the stand level. Models 1 and 2 appear to explain a larger variability of Rm than models based on stem area or sapwood mass.

Cell Respiration↗

PubMed2003

Osmotic adjustment in three-year-old seedlings of five provenances of maritime pine (Pinus pinaster) in response to drought.

Three-year-old seedlings of five provenances of Pinus pinaster Ait. that differed in climatic conditions at their geographical origin were subjected to decreasing soil water availability. The degree of needle osmotic adjustment (OA) was estimated based on logarithmic plots of needle relative water content (RWC) against needle osmotic potential (Psi(pi)); i.e., lnRWC versus -ln(-Psi(pi)). There were significant differences among provenances in active OA (0.13 to 0.30 MPa for a decrease in RWC to 80%), and a clear negative relationship was found between OA and precipitation (650 to 1280 mm of mean annual rainfall) at the geographical origins of the provenances. A high osmoregulatory capacity contributes to the maintainance of positive turgor at low water potentials. We conclude that OA is one of the mechanisms underlying adaptation to drought in P. pinaster. Solute accumulation was about 2.3 times higher in the provenance from the driest site than in the provenance from the wettest site. The contribution of osmotic adjustment to differences in drought tolerance mechanisms among provenances is discussed.

Adaptation, Physiological↗

PubMed2003

Timing of GA4/7 application and the flowering of Pinus sylvestris grafts in the greenhouse.

Effects of varying the time of stem injection of gibberellin A4/7 (GA4/7) on male and female flowering in potted grafts of Pinus sylvestris L. in a greenhouse were monitored in two consecutive years. Flowering results for each application time were correlated with phenological characteristics and abiotic factors in an attempt to find reproducible indicators of the most effective time of GA4/7 treatment. The GA4/7 treatment had a significant effect on female flowering, but no effect on male flowering. In both years, the largest effect on female flowering was obtained with the latest application. Percent shoot and needle elongation (relative to final length) were poor indicators of the optimal time of GA4/7 application. Three heat sum accumulation models gave equally accurate predictions of the optimal time for GA4/7 application; therefore, the simplest model, i.e., the sum of degree-days, is recommended. The timing window for stimulation of female flowering with GA4/7 had a lower limit of about 500 degree-days (threshold +5 degrees C), and the optimal time of GA4/7 application was after about 700 degree-days.

Flowers↗

PubMed2003

Variation in needle longevity of Pinus tabulaeformis forests at different geographic scales.

Needle longevity of conifer species is known to increase with latitude, but little is known about intraspecific variation and associated factors within a location. Chinese pine (Pinus tabulaeformis Carr.) forests were investigated to identify patterns of needle longevity at seven sites with distinctive climatic conditions along a latitudinal gradient (33-38 degrees N) in Shaanxi province, northwest China. A demographic approach was used to quantify needle longevity as an index of entire foliage population adjusted to needle-age-specific mortality rates. There were significant differences in needle longevity of Chinese pine stands across sites and across sample plots within a site. Individual tree needle longevity ranged from 0.62 to 3.75 years for 276 samples across sites. Needle longevity increased with latitude (R2 = 0.40, P < 0.0001), but decreased with mean January temperature (R2 = 0.63, P < 0.0001). Foliage retention of Chinese pine stands at the regional level was generally associated with climatic variability, indicating that variation in needle longevity was primarily an environmental acclimation to low temperature in winter. Stand characteristics were closely associated with needle longevity at three sites located within the same climatic zone. Needle longevity was positively correlated with tree age (R2 = 0.48, P < 0.0001) and stand density (R2 = 0.26, P = 0.0015) at Huanglong and Huangling, respectively, whereas it was negatively associated with total tree height at Zhidan (R2 = 0.50, P < 0.0001). It is concluded that, at the stand and individual tree level, intraspecific variation in needle longevity is most likely a result of adaptation to patchy microsite environments.

China↗

PubMed2003

Response of photosynthesis in second-generation Pinus radiata trees to long-term exposure to elevated carbon dioxide partial pressure.

Second-generation Pinus radiata D. Don trees, propagated from cuttings of 4-year-old trees previously grown at ambient (36 Pa) and elevated (65 Pa) CO2 partial pressure (Ca) were grown under the same conditions in open-top chambers for a further year. As cuttings of the original trees, these second-generation trees were physiologically the same age as the first-generation trees with the only difference between the two being size. This allowed us to test the effects of tree size independently of age or duration of exposure. Total non-structural carbohydrate concentration, area-based nitrogen concentration, leaf mass per unit area and chlorophyll concentration measured in three foliage age cohorts were unaffected by either age or Ca. There were no signs of photosynthetic down-regulation in trees grown at elevated Ca. When measured at the growth Ca, photosynthetic rate in young needles during summer, autumn and spring was 34, 43 and 38% higher, respectively, in trees grown at elevated Ca than in trees grown at ambient Ca. In older needles, the corresponding photosythetic rate increases were 26, 47 and 49%. Water-use efficiency, determined by stable carbon isotope analysis, was 49% higher in foliage in the elevated Ca treatment than in foliage in the ambient Ca treatment. This increase was entirely due to photosynthetic enhancement, because stomatal conductance did not differ between treatments. We conclude that down-regulation of photosynthesis at elevated Ca is related to tree size rather than tree age or duration of exposure, and that enhanced photosynthetic rates can be maintained while sink strength is high enough to use the excess photosynthates.elevated CO2, needle age, photosynthetic down-regulation, photosynthetic enhancement, sink strength, water-use efficiency.

Carbon Dioxide↗

PubMed2004

Effectiveness of mycorrhizal inoculation in the nursery on growth and water relations of Pinus radiata in different water regimes.

We compared the performance of mycorrhiza-inoculated and non-inoculated radiata pine (Pinus radiata D. Don) seedlings in two field studies. In the first study, above ground growth of 2-year-old container-grown trees was compared after planting in two areas that differed in water availability. Mycorrhizal inoculation significantly improved above ground growth in the 2 years after planting in both areas, especially at the drier site. In the second study, the influence of Rhizopogon roseolus (Corda ex Sturm) Th. Fries and Scleroderma citrinum Pers. on xylem water potential and hydraulic conductance at the soil-root interface of trees established at the same sites was assessed. Nursery inoculations with R. roseolus and S. citrinum improved tree growth during the first 2 years after field outplanting, particularly at the drier site.

Chlorophyll↗

PubMed2004

Changes in polyamine concentration associated with aging in Pinus radiata and Prunus persica.

Endogenous polyamine concentrations in apical buds of peach (Prunus persica (L.) Batsch cv. 'Hakuho') and monterey pine (Pinus radiata D. Don) seedlings, saplings and mature trees were analyzed by high performance liquid chromatography coupled with fluorescence detection to establish physiological correspondences and disparities between the species. Although there were many differences in polyamine metabolism between peach and monterey pine, several similarities were found. Specifically, in both species, the ratio between concentrations of free polyamines and low molecular-weight polyamine conjugates was greater than and less than unity in juvenile and adult plants, respectively. Free putrescine concentration decreased with age in both species.

Biogenic Polyamines↗

PubMed2004

Effect of parent genotype on somatic embryogenesis in Scots pine (Pinus sylvestris).

Controlled crosses of seven Scots pine (Pinus sylvestris L.) trees produced 49 families that included both reciprocals and selfings. Embryogenic cultures were initiated from immature megagametophytes and after 6 months in maintenance culture, mature somatic embryos were produced from the surviving 166 lines. The effect of parent genotypes on the cultures was evaluated at initiation of the tissue culture period, after 6 months in maintenance culture and at embryo maturation. The effect of the maternal parent was most pronounced at culture initiation. After 6 months in tissue culture, the maternal effect had decreased and the effects of both parents were significant. By the somatic embryo maturation stage, the maternal effect was still considerable but the paternal effect was no longer detectable. There was little correlation between the ranking of mothers and fathers, indicating that the maternal effect was caused by factors other than the paternal effect. No mother x father interaction was found, indicating that mothers successful at initiation and after 6 months in tissue culture, pollinated by any of the successful fathers, produced somatic lines and mature somatic embryos.

Genotype↗

PubMed2004

Characterization, expression and evolution of two novel subfamilies of Pinus monticola cDNAs encoding pathogenesis-related (PR)-10 proteins.

Proteins of the pathogenesis-related (PR)-10 family are induced in many plants by phytopathogens and environmental stresses. A multi-gene family of PR10 proteins has previously been found in the genome of western white pine (Pinus monticola Dougl. ex D. Don). We isolated two novel subfamilies of PR10 cDNAs (PmPR10-2 and PmPR10-3) from P. monticola that are distinct from other PR10 genes (PmPR10-1.1-1.14) reported from the same species. The PmPR10 proteins are grouped in three subfamilies based on similarity in amino acid sequences. The sequence identities of PmPR10 proteins are much higher among members within a subfamily than among members of different subfamilies (86-99% versus 59-68%). Induction of both PmPR10-2 and PmPR10-3 mRNAs was detected by reverse transcriptase-polymerase chain reaction (RT-PCR) in needles in response to wounding treatment. PmPR10-3 was also expressed in needles during cold acclimation in winter. Transcript levels of both PmPR10-2 and PmPR10-3 were less than the detectable levels of constitutive expression in roots, stems and vegetative shoots, whereas PmPR10-1.10 mRNA of subfamily I was expressed at various levels. Phylogenetic analysis showed that PmPR10 and PR10 proteins from other conifers are grouped within one clade that is distinct from that of angiosperm PR10 proteins. In the conifer monophyletic group, PR10 sequences diversify into three distinct clusters. Among these three clusters, some PR10 proteins from single conifer species showed greater divergence distances than sequences from other conifer species, suggesting that, within the conifers, the multi-gene family underwent great diversification during evolution. Based on ratios of nonsynonymous to synonymous nucleotide substitutions (Ka/Ks), we speculate that positive selection resulted in the divergence of PmPR10 subfamilies I and III. Possible mechanisms and significance of PR10 gene evolution are discussed.

Amino Acid Sequence↗

PubMed2004

Regulation of organic and inorganic nitrogen uptake in Scots pine (Pinus sylvestris) seedlings.

Plants possess regulatory mechanisms that enhance nitrogen (N) uptake under conditions of spatial and temporal variation in N availability. Study of regulatory mechanisms has focused almost exclusively on the uptake of inorganic N sources (i.e., ammonium (NH4+), nitrate (NO3-). Several lines of evidence, however, suggest that amino acids may constitute a potential source of N for a number of plant species, including conifers. In the present study, we investigated the uptake of amino acids and inorganic N in Scots pine (Pinus sylvestris L.) seedlings grown at different N concentrations. We compared the uptake rate of the individual N sources using U-[13C2], [15N]-glycine, U-[13C6], [15N4]-arginine, 15NH4, or 15NO3, and tested the short-term effect of N supply on the uptake rate of glycine, arginine and in field-grown Scots pine seedlings. Our data indicate that Scots pine seedlings can absorb substantial amounts of N in the form of intact arginine and glycine molecules. The data also suggest that Scots pine seedlings down-regulate their uptake of NH4+-N and arginine-N, but not of glycine-N in response to increased endogenous N concentrations.

Nitrogen↗

PubMed2004

Assessing variation in the radial profile of sap flux density in Pinus species and its effect on daily water use.

We monitored sap flux density (v) diurnally in nine mature southeastern pine (Pinus spp.) trees with a thermal dissipation probe that spanned the sapwood radius. We found the expected pattern of high v near the cambium and decreasing v with depth toward the center of the tree; however, the pattern was not constant within a day or between trees. Radial profiles of trees were steeper earlier in the day and became less steep later in the day. As a result, time-dependent changes in the shape of the radial profile of v were sometimes correlated with daily changes in evaporative demand. As the radial profile became less steep, the inner xylem contributed relatively more to total tree sap flow than it did earlier in the day. We present a 3-parameter Gaussian function that can be used to describe the radial distribution of v in trees. Parameters in the function represent depth in the xylem from the cambium, maximum v, depth in the xylem where maximum v occurs, and the rate of radial change in v with radial depth (beta). Values of beta varied significantly between trees and with time, and were sometimes correlated with air vapor pressure deficit (D). We hypothesize that this occurred during periods of high transpiration when the water potential gradient became great enough to move water in the inner sapwood despite its probable high hydraulic resistance. We examined discrepancies among estimates of daily water use based on single-point, two-point and multi-point (i.e., every 20 mm in the sapwood) measurements. When radial distribution of v was not considered, a single-point measurement resulted in errors as large as 154% in the estimate of daily water use relative to the estimate obtained from a multi-point measurement. Measuring v at two close sample points (10 and 30 mm) did not improve the estimate; however, estimates derived from v measured at two distant sample points (10 and 70 mm) significantly improved the estimate of daily water use, although errors were as great as 32% in individual trees. The variability in v with depth in the xylem, over time, and between trees indicates that measurements of the radial distribution of v are necessary to accurately estimate water flow in trees with large sapwood areas.

Circadian Rhythm↗