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Genetic variation of the bud and leaf phenology of seventeen poplar clones in a short rotation coppice culture.

Leaf phenology of 17 poplar ( Populus spp.) clones, encompassing spring phenology, length of growth period and end-of-year phenology, was examined over several years of different rotations. The 17 poplar clones differed in their latitude of origin (45 degrees 30'N to 51 degrees N) and were studied on a short rotation experimental field plantation, situated in Boom (province of Antwerpen, Belgium; 51 degrees 05'N, 04 degrees 22'E). A similar, clear pattern of bud burst was observed during the different years of study for all clones. Clones Columbia River, Fritzi Pauley, Trichobel (Populus trichocarpa) and Balsam Spire (Populus trichocarpa x Populus balsamifera) from 45 degrees 30'N to 49 degrees N reached bud burst (expressed as day of the year or degree day sums) almost every year earlier than clones Wolterson (Populus nigra), Gaver, Gibecq and Primo (Populus deltoides x Populus nigra) (50 degrees N to 51 degrees N). This observation could not be generalised to end-of-season phenology, for which a yearly returning pattern for all clones was lacking. Late bud burst and early leaf fall of some clones (Beaupré, Boelare, IBW1, IBW2, IBW3) was brought about by increasing rust incidence during the years of observation. For these clones, the variability in leaf phenology was reflected in high coefficients of variation among years. The patterns of genetic variation in leaf phenology have implications for short rotation intensive culture forestry and management of natural populations. Moreover, the variation in phenology reported here is relevant with regard to the genetic mapping of poplar.

Genetic Variation↗

Responses of Riparian Cottonwoods to Alluvial Water Table Declines.

/ Human demands for surface and shallow alluvial groundwater have contributed to the loss, fragmentation, and simplification of riparian ecosystems. Populus species typically dominate riparian ecosystems throughout arid and semiarid regions of North American and efforts to minimize loss of riparian Populus requires an integrated understanding of the role of surface and groundwater dynamics in the establishment of new, and maintenance of existing, stands. In a controlled, whole-stand field experiment, we quantified responses of Populus morphology, growth, and mortality to water stress resulting from sustained water table decline following in-channel sand mining along an ephemeral sandbed stream in eastern Colorado, USA. We measured live crown volume, radial stem growth, annual branch increment, and mortality of 689 live Populus deltoides subsp. monilifera stems over four years in conjunction with localized water table declines. Measurements began one year prior to mining and included trees in both affected and unaffected areas. Populus demonstrated a threshold response to water table declines in medium alluvial sands; sustained declines >/=1 m produced leaf desiccation and branch dieback within three weeks and significant declines in live crown volume, stem growth, and 88% mortality over a three-year period. Declines in live crown volume proved to be a significant leading indicator of mortality in the following year. A logistic regression of tree survival probability against the prior year's live crown volume was significant (-2 log likelihood = 270, chi2 with 1 df = 232, P < 0.0001) and trees with absolute declines in live crown volume of >/=30 during one year had survival probabilities <0.5 in the following year. In contrast, more gradual water table declines of thick similar0.5 m had no measurable effect on mortality, stem growth, or live crown volume and produced significant declines only in annual branch growth increments. Developing quantitative information on the timing and extent of morphological responses and mortality of Populus to the rate, depth, and duration of water table declines can assist in the design of management prescriptions to minimize impacts of alluvial groundwater depletion on existing riparian Populus forests.

Journal Article↗

Increased leaf area expansion of hybrid poplar in elevated CO2. From controlled environments to open-top chambers and to FACE.

We examined the response of hybrid poplar to elevated CO2 in contrasting growth environments: controlled environment chamber (CE). open-top chamber (OTC) and poplar free air CO2 enrichment (POPFACE) in order to compare short versus long-term effects and to determine whether generalisations in response are possible for this fast growing tree. Leaf growth, which for poplar is an important determinant of stemwood productivity was followed in all environments, as were the determinants of leaf growth-cell expansion and cell production. Elevated CO2 (550-700 micromol mol(-1), depending on environment) resulted in an increase in final leaf size for Populus trichocarpa x Populus deltoides (Populus x interamericana) and P. deltoides x Populus nigra (Populus x euramericana), irrespective of whether plants were exposed during a short-term CE glasshouse study (90 days), a long-term OTC experiment (3 years) or during the first year of a POPFACE experiment. An exception was observed in the closed canopy POPFACE experiment, where final leaf size remained unaltered by CO2. Increased leaf extension rate was observed in elevated CO2 in all experiments, at some point during leaf development, as determined by leaf length. Again the exception were the POPFACE experiment, where effects were not statistically significant. Leaf production and specific leaf area (SLA) were increased and decreased, respectively, on five out of six occasions, although both were only statistically significant on two occasions and interestingly for SLA never in the FACE experiment. Although both cell expansion and cell production were sensitive to CO2 concentration, effects appeared highly dependent on growth environment and genotype. However, increased leaf cell expansion in elevated CO2 was often associated with changes in the biophysical properties of the cell wall, usually increased cell wall plasticity. This research has shown that enhanced leaf area development was a consistent response to elevated CO2 but that the magnitude of this response is likely to decline, in long-term exposure to elevated CO2. Effects on SLA and leaf production suggest that CE and OTC experiments may not always provide good predictors of the 'qualitative' effects of elevated CO2 in long-term ecosystem experiments.

Air Pollutants↗

Detritus processing and microbial dynamics of an aquatic macrophyte and terrestrial leaf in a thermally constant, spring-fed stream.

Past studies of organic matter processing in stream ecosystems have focused on the fate of allochthonous terrestrial leaf detritus. In streams with a reduced canopy, submerged macrophytes may provide a significant source of organic matter to the microbial community and higher trophic levels. We compared mass loss patterns and microbial dynamics between a submerged macrophyte, Sagittaria platyphylla, and a deciduous leaf, Populus deltoides. Mass loss rates were higher for the submerged macrophyte, though exponential decay values indicated that both are "fast" decomposers. Bacterial abundance was not significantly different between plant types, but bacterial productivity was significantly higher in Sagittaria. Although fungal biomass was higher overall for Populus, it was not significantly different from that of Sagittaria until day 30. Relative to fungi, bacteria made up 4% and 7% of the peak microbial biomass on Populus and Sagittaria, respectively. Aquatic hyphomycete sporulation was detected only on Populus. These results suggest that in systems where submerged macrophytes are abundant, they can provide a carbon source quantitatively comparable to that of riparian leaf detritus, but that qualitative differences in leaf structural composition cause a shift toward a more significant role for bacterial decomposers.

Bacteria↗

A method for describing the canopy architecture of coppice poplar with allometric relationships.

A multi-scale biometric methodology for describing the architecture of fast-growing short-rotation woody crops is used to describe 2-year-old poplar clones during the second rotation. To allow for expressions of genetic variability observed within this species (i.e., growth potential, leaf morphology, coppice and canopy structure), the method has been applied to two clones: Ghoy (Gho) (Populus deltoides Bartr. ex Marsh. x Populus nigra L.) and Trichobel (Tri) (Populus trichocarpa Torr. & A. Gray x Populus trichocarpa). The method operates at the stool level and describes the plant as a collection of components (shoots and branches) described as a collection of metameric elements, themselves defined as a collection of elementary units (internode, petiole, leaf blade). Branching and connection between the plant units (i.e., plant topology) and their spatial location, orientation, size and shape (i.e., plant geometry) describe the plant architecture. The methodology has been used to describe the plant architecture of 15 selected stools per clone over a 5-month period. On individual stools, shoots have been selected from three classes (small, medium and large) spanning the diameter distribution range. Using a multi-scale approach, empirical allometric relationships were used to parameterize elementary units of the plant, topological relationships and geometry (e.g., distribution of shoot diameters on stool, shoot attributes from shoot diameter). The empirical functions form the basis of the 3-D Coppice Poplar Canopy Architecture model (3-D CPCA), which recreates the architecture and canopy structure of fast-growing coppice crops at the plot scale. Model outputs are assessed through visual and quantitative comparisons between actual photographs of the coppice canopy and simulated images. Overall, results indicate a good predictive ability of the 3-D CPCA model.

Imaging, Three-Dimensional↗

A transcriptional timetable of autumn senescence.

BACKGROUND: We have developed genomic tools to allow the genus Populus (aspens and cottonwoods) to be exploited as a full-featured model for investigating fundamental aspects of tree biology. We have undertaken large-scale expressed sequence tag (EST) sequencing programs and created Populus microarrays with significant gene coverage. One of the important aspects of plant biology that cannot be studied in annual plants is the gene activity involved in the induction of autumn leaf senescence. RESULTS: On the basis of 36,354 Populus ESTs, obtained from seven cDNA libraries, we have created a DNA microarray consisting of 13,490 clones, spotted in duplicate. Of these clones, 12,376 (92%) were confirmed by resequencing and all sequences were annotated and functionally classified. Here we have used the microarray to study transcript abundance in leaves of a free-growing aspen tree (Populus tremula) in northern Sweden during natural autumn senescence. Of the 13,490 spotted clones, 3,792 represented genes with significant expression in all leaf samples from the seven studied dates. CONCLUSIONS: We observed a major shift in gene expression, coinciding with massive chlorophyll degradation, that reflected a shift from photosynthetic competence to energy generation by mitochondrial respiration, oxidation of fatty acids and nutrient mobilization. Autumn senescence had much in common with senescence in annual plants; for example many proteases were induced. We also found evidence for increased transcriptional activity before the appearance of visible signs of senescence, presumably preparing the leaf for degradation of its components.

Aging↗

Comparative water relations of three successional hardwood species in central Wisconsin.

Water relations of co-occurring understory saplings of Quercus ellipsoidalis E.J. Hill, an early successional, xeric species, Populus tremuloides Michx., an early successional, mesic species, and Acer rubrum L., a late successional species that occurs on both wet and dry sites, were evaluated on four dates during the 1986 growing season. The understory was characterized by high soil water content, low irradiance and low vapor pressure deficit throughout the growing season. Stomatal conductance and calculated transpiration flux were lowest for A. rubrum and highest for P. tremuloides and Q. ellipsoidalis. Except early in the growing season, leaf water potentials were lower in P. tremuloides than in the other species. Populus tremuloides had the highest bulk modulus of elasticity, Q. ellipsoidalis the lowest. Over the growing season, Populus tremuloides and Q. ellipsoidalis, but not A. rubrum, exhibited a decrease in osmotic potential at both full and zero turgor. Of the three species, Populus tremuloides exhibited the sharpest decrease in leaf water potential and turgor pressure with decreasing relative water content.

Journal Article↗

Xylem embolism in response to freeze-thaw cycles and water stress in ring-porous, diffuse-porous, and conifer species.

Vulnerability to xylem embolism by freeze-thaw cycles and water stress was quantified in ring-porous (Quercus gambelii Nutt.), diffuse-porous (Populus tremuloides Michx., Betula occidentalis Hook.), and conifer species (Abies lasiocarpa Nutt., Juniperus scopulorum Sarg.). Embolism was measured by its reduction of xylem hydraulic conductivity; it was induced by xylem tension (water-stress response) and by a tension plus a freeze-thaw cycle (freeze response). Conifers showed little (Juniperus) or no (Abies) freeze response even to repeated cycles. In contrast, Quercus embolized more than 90% by freezing at tensions below 0.2 MPa, whereas similar embolism without freezing required tensions above 4.5 MPa. Diffuse-porous trees (Betula, Populus) showed an intermediate freeze response. The magnitude of the freeze response was correlated with conduit volume but occurred at higher tensions than predicted from theory. Large early-wood vessels (2.8 x 10(-9) m(3)) in oak were most vulnerable to embolism by freezing, small vessels in Populus and Betula were intermediate (approximately 7 x 10(-11) m(3)), and tracheids in conifers (about 3 x 10(-13) m(3)) were most resistant. The same trend was found within a stem: embolism by freeze-thawing occurred preferentially in wider conduits. The water-stress response was not correlated with conduit volume; previous work indicates it is a function of interconduit pit membrane structure. Native embolism levels during winter corroborated laboratory results on freezing: Quercus embolized 95% with the first fall freeze, Populus and Betula showed gradual increases to more than 90% embolism by winter's end, and Abies remained below 30%.

Journal Article↗

Processesof Tamarix invasion and floodplain development along the lower Green River, Utah.

Significant ecological, hydrologic, and geomorphic changes have occurred during the 20th century along many large floodplain rivers in the American Southwest. Native Populus forests have declined, while the exotic Eurasian shrub, Tamarix, has proliferated and now dominates most floodplain ecosystems. Photographs from late 19th and early 20th centuries illustrate wide river channels with largely bare in-channel landforms and shrubby higher channel margin floodplains. However, by the mid-20th century, floodplains supporting dense Tamarix stands had expanded, and river channels had narrowed. Along the lower Green River in eastern Utah, the causal mechanism of channel and floodplain changes remains ambiguous due to the confounding effects of climatically driven reductions in flood magnitude, river regulation by Flaming Gorge Dam, and Tamarix invasion. This study addressed whether Tamarix establishment and spread followed climate- or dam-induced reductions in annual peak flows or whether Tamarix was potentially a driver of floodplain changes. We aged 235 Tamarix and 57 Populus individuals, determined the hydrologic and geomorphic processes that controlled recruitment, identified the spatial relationships of germination sites within floodplain stratigraphic transects, and mapped woody riparian vegetation cohorts along three segments of the lower Green River. The oldest Tamarix established along several sampling reaches in 1938, and 1.50-2.25 m of alluvium has accreted above their germination surfaces. Nearly 90% of the Tamarix and Populus samples established during flood years that exceeded the 2.5-year recurrence interval. Recruitment was most common when large floods were followed by years with smaller peak flows. The majority of Tamarix establishment and Green River channel narrowing occurred long before river regulation by Flaming Gorge Dam. Tamarix initially colonized bare instream sand deposits (e.g., islands and bars), and most channel and floodplain changes followed the establishment of Tamarix. Our results suggest that Tamarix recruitment was triggered by large annual floods that were followed by years with lower peak flows, not by periods of low flow alone. Tamarix appears to have actively invaded floodplains, while Populus colonization has been limited. Thus, Tamarix invasion may have greatly influenced floodplain development and riparian vegetation composition along the lower Green River since the early 20th century.

Disasters↗

[Physioecological characteristics of four dominant plant species in Kerqin sandy land].

The photosynthetic and water physioecological characteristics of four dominant species, Caragana microphylla, Artemisia halodendron, Artemisia frigida and Populus spp(a hybrid), in Kerqin sandy land were compared. The photosynthesis rates of C. microphylla, A. halodendron, and Populus spp. reached highest at 6:00, rapidly fall at 8:00, remained at low level from 8:00 to 16:00, then rose slowly after 16:00. The photosynthesis rate of A. frigida was comparatively low at 6:00, so the range of declining speed was small. Diurnal photosynthesis rate of Artemisia frigida decreased at 6:00-8:00, C. microphylla at 8:00-10:00, Populus spp and. A. halodendron at 6:00-10:00, and was controlled by stomatal limitation; in rest time, it was controlled by non-stomatal limitation. The four species also adapted different ways to resist drought stress. C. microphylla and A. frigida resisted drought stress through their low transpiration, low water potential and high ratio of bound water to free water(BW/FW); Populus spp. resisted drought stress through its rapid fall of stomatal conductivity to reduce its transpiration in spite of its high water potential and high BW/FW, while A. halodendron resisted drought stress weakly because of its high transpiration, high water consumption and low water use efficiency.

Artemisia↗

Botryozyma mucatilis sp. nov., an anamorphic ascomycetous yeast associated with nematodes in poplar slime flux.

A new species of Botryozyma, Botryozyma mucatilis, was isolated from the surface of free-living nematodes, Panagrellus dubius, inhabiting slime flux from hybrid poplars, Populus deltoidesxtrichocarpa, in Oregon, USA. This species was discovered in relatively close proximity to the teleomorphic species Ascobotryozyma americana and Ascobotryozyma cognata, both collected from P. dubius nematodes inhabiting beetle galleries in Populus spp. and Populus and Salix spp., respectively. B. mucatilis is recognized as a distinct species based on molecular and morphological data. Sequence divergence in both the D1/D2 domain of the nuclear large-subunit rDNA and internal transcribed spacer region rDNA, low DNA reassociation values, notably different amplified fragment-length polymorphic fingerprints, and significantly longer cells all support the designation of a novel species.

Animals↗

Molecular cloning of ten distinct hypervariable regions from the cellulose synthase gene superfamily in aspen trees.

Recent molecular genetic data suggest that cellulose synthase (CesA) genes coding for the enzymes that catalyze cellulose biosynthesis (CESAs) in Arabidopsis and other herbaceous plants belong to a large gene family. Much less is known about CesA genes from forest trees. To isolate new CesA genes from tree species, discriminative but easily obtainable homologous DNA probes are required. Hypervariable regions (HVRII) of CesA genes represent highly divergent DNA sequences that can be used to examine structural, expressional and functional relationships among CesA genes. We used a reverse transcriptase-polymerase chain reaction (RT-PCR)-based technique to identify HVRII regions from eight types of CesA genes and two types of CesA-like D (CslD) genes in quaking aspen (Populus tremuloides Michx.). Comparison of these aspen CESA/CSLD HVRII regions with the predicted proteins from eight full-length CesA/CslD cDNAs available in our laboratory and with searches for aspen CesA/CslD homologs in the recently released Populus trichocarpa Torr. & A. Gray. genome confirmed the utility of this approach in identifying several CesA/CslD gene members from the Populus genome. Phylogenetic analysis of 56 HVRII domains from a variety of plant species suggested that at least six distinct classes of CESAs exist in plants, supporting a previous proposal for renaming HVRII regions as class-specific regions (CSR). This method of CSR cloning could be applied to other crop plants and tree species, especially softwoods, for which the whole genome sequence is unlikely to become available in the near future because of the large size of these genomes.

Cloning, Molecular↗

Translocation of nitrogen in the xylem of field-grown cherry and poplar trees during remobilization.

Studies of small trees growing in pots have established that individual amino acids or amides are translocated in the xylem sap of a range of tree species following bud burst, as a consequence of nitrogen (N) remobilization from storage. This paper reports the first study of N translocation in the xylem of large, deciduous, field-grown trees during N remobilization in the spring. We applied 15N fertilizer to the soil around 10-year-old Prunus avium L. and Populus trichocharpa Torr. & Gray ex Hook var. Hastata (Dode) A. Henry x Populus balsamifera L. var. Michauxii (Dode) Farwell trees before bud burst to label N taken up by the roots. Recovery of unlabeled N in xylem sap and leaves was used to demonstrate that P. avium remobilizes N in both glutamine (Gln) and asparagine (Asn). Sap concentrations of both amides rose sharply after bud burst, peaking 14 days after bud burst for Gln, and remaining high some 45 days for Asn. There was no 15N enrichment of either amide until 21 days after bud burst. In the Populus trees, nearly all the N was translocated in the sap as Gln, the concentration of which peaked and then declined before the amide was enriched with 15N, 40 days after bud burst. Xylem sap of clonal P. avium trees was sampled at different positions in the crown to assess if the amino acid and amide composition of the sap varied within the crown. Sap was sampled during remobilization (when the concentration of Gln was maximal), at the end of remobilization and at the end of the experiment (68 days after bud burst). Although the date of sampling had a highly significant effect on sap composition, the effect of position of sampling was marginal. The results are discussed in relation to N translocation in adult trees and the possibility of measuring N remobilization by calculating the flux of N translocation in the xylem.

Amides↗

Phenotypic variation and quantitative trait locus identification for osmotic potential in an interspecific hybrid inbred F2 poplar pedigree grown in contrasting environments.

Elucidation of the mechanisms of dehydration tolerance in poplar (Populus sp.) trees will permit development of biochemical and molecular indicators to identify dehydration-tolerant genotypes during genetic selection. The objectives of this study were to characterize the degree of phenotypic variation in osmotic potential (a determinant of dehydration tolerance), determine the relationship between osmotic potential at full turgor and relative growth rate, and identify quantitative trait loci (QTL) for osmotic potential in an advanced-generation, interspecific poplar pedigree established in contrasting environments. A three-generation, sib-mated black cottonwood (Populus trichocarpa Torr. & Gray) and eastern cottonwood (P. deltoides Bartr.) segregating F(2) family (Family 331) was analyzed at a dry site east of the Cascade Mountain Range (Boardman, OR) and at a wet site west of the mountains (Clatskanie, OR). At the Boardman site, 2-year-old trees (59 clones) were either irrigated everyday (wet) or every other day (dry), whereas 3- and 4-year-old trees (58 clones) at the Clatskanie site were unirrigated. At the Boardman site, the typically narrow range of osmotic potentials exhibited by grandparents and parents was greatly expanded in the F(2) population, spanning from -1.38 to -2.35 MPa under wet conditions, with a similar range under dry conditions (-1.40 to -2.15 MPa). Clones that had osmotic potentials < or = -1.90 MPa generally displayed full maintenance of stem relative growth rates under dry conditions in contrast to clones with osmotic potentials that were < or = -1.60 MPa, in which stem relative growth rates were reduced by an average of 38% in the dry treatment relative to the wet treatment. Although osmotic adjustments of 0.13 to 0.36 MPa were observed in nine out of 59 clones, adjustment typically occurred from relatively high baseline osmotic potentials. The range in osmotic potential at the wetter Clatskanie site at age three was higher (-1.27 to -1.84 MPa) and was further expanded the following year (-1.14 to -1.94 MPa), which had a wetter spring than the previous year, followed by a typically dry July. Seven QTL for osmotic potential were identified that each explained > 7.5% of the variation in osmotic potential. Given that four clones (7%) had osmotic potentials of -2.00 MPa or less and that QTL for osmotic potential have been identified, we suggest that there are opportunities to extend the limit of dehydration tolerance in Populus.

Adaptation, Physiological↗

EST data suggest that poplar is an ancient polyploid.

We analysed the publicly available expressed sequence tag (EST) collections for the genus Populus to examine whether evidence can be found for large-scale gene-duplication events in the evolutionary past of this genus. The ESTs were clustered into unigenes for each poplar species examined. Gene families were constructed for all proteins deduced from these unigenes, and K(S) dating was performed on all paralogs within a gene family. The fraction of paralogs was then plotted against the K(S) values, which resulted in a distribution reflecting the age of duplicated genes in poplar. Sufficient EST data were available for seven different poplar species spanning four of the six sections of the genus Populus. For all these species, there was evidence that a large-scale gene-duplication event had occurred. From our analysis it is clear that all poplar species have shared the same large-scale gene-duplication event, suggesting that this event must have occurred in the ancestor of poplar, or at least very early in the evolution of the Populus genus.

Biological Evolution↗

Differences in the kinetics and scale of signalling molecule production modulate the ozone sensitivity of hybrid poplar clones: the roles of H2O2, ethylene and salicylic acid.

Hydrogen peroxide (H(2)O(2)), ethylene, 1-aminocyclopropane-1-carboxylic acid (ACC) and salicylic acid (SA) concentrations and ACC synthase (ACS) gene expression were measured to establish whether the high sensitivity of the Populus deltoides x maximowiczii clone Eridano to ozone (O(3)) exposure, compared with the O(3)-resistant Populus deltoides x euramericana clone I-214, is attributable to differences in the modulation of signal transduction pathways. In a time-course experiment, Populus deltoides (poplar) clones were exposed to acute fumigation with 150 nl l(-1) O(3) for 5 h. The two poplar clones showed differences in ethylene evolution, I-214 displaying earlier and less pronounced ethylene emission than Eridano. In both clones, ethylene evolution was accompanied by increased ACS transcript levels and enhanced emission of free ACC. I-214 exhibited a greater basal concentration of free SA and a lower concentration of the conjugated pool. However, a slight accumulation of free SA at the end of the 5-h exposure was found only in Eridano, together with an earlier minimal increase in the concentration of conjugated SA. The results show that both clones react to O(3) by producing H(2)O(2), ethylene and SA, but the difference in sensitivity to the pollutant is probably attributable to differences in the kinetics and magnitude of this response.

Base Sequence↗