Diurnal and Seasonal Patterns of Photosynthesis and Respiration by Stems of Populus tremuloides Michx

The photosynthetic and respiratory rates of 5- to 7-year-old aspen stems (Populus tremuloides Michx.) were monitored in the field for 1 year to determine the seasonal patterns. The stem was not capable of net photosynthesis, but the respiratory CO₂ loss from the stem was reduced by 0 to 100% depending on the time of year and the level of illumination as a result of bark photosynthesis. The monthly dark respiratory rate ranged from 0.24 mg CO₂/dm²· hr in January to a maximum 7.4 mg CO₂/dm²· hr in June. Individual measurements ranged from 0.02 mg CO₂/dm²· hr in February to 12.3 mg CO₂/dm²· hr in June. Gross photosynthesis followed a pattern similar to the dark respiratory rate. The mean monthly rate was highest in June (1.65 mg CO₂/dm²· hr) and lowest in December (0.02 mg CO₂/dm²· hr). Individual measurements ranged from 0.0 mg CO₂/dm²· hr in winter to 5.5 mg CO₂/dm²· hr in July.     

Photosynthesis and Respiration of Developing Populus tremuloides Internodes

The photosynthetic and respiratory performance of developing internodes of Populus tremuloides was evaluated by infrared gas analysis. Anatomical and morphological transitions were related to metabolic activity. Photosynthetic rates ranged from 6.0 to 10.0 milligrams CO₂ per decimeter squared per hour in the youngest internodes to 2.5 to 3.8 milligrams CO₂ per decimeter squared per hour in internodes with fully developed bark tissues. Respiration exceeded the rate of photosynthesis on the average by a factor of two. Stem photosynthesis increased with temperature up to 40°C and declined steeply between 40 and 50°C. Stem respiration increased nearly linearly to temperatures as high as 50°C.     

Genetic linkage mapping in aspen (Populus tremula L. and Populus tremuloides Michx.)

A large number of simple sequence repeat (SSR) marker-containing genetic maps are available for several Populus species. For aspen however, no SSR-containing map has been published so far. In this study, genetic linkage mapping was carried out with an interspecific mapping pedigree of 61 full-sib hybrids of European × quaking aspen (Populus tremula L. × Populus tremuloides Michx.), using the two-way pseudo-testcross strategy. Amplified fragment-length polymorphism (AFLP) and SSR markers were used for mapping, resulting in the first SSR-containing genetic linkage maps for aspen. The maps allow comparisons with a Populus consensus map and other published genetic maps of the genus Populus. The maps showed good collinearity to each other and to the Populus consensus map and provide a direct link to the Populus trichocarpa genomic sequence. Sex as a morphological trait was assessed in the mapping population and mapped on a non-terminal position of linkage group XIX on the male P. tremuloides map.     

Clonal and spatial genetic structures of aspen (Populus tremuloides Michx.)

To portray aspen clonal and spatial genetic structures, we mapped and genotyped trees in two 1-ha plots, each containing three aspen cohorts originating from fire or subsequent secondary disturbances. We used four microsatellite loci to identify aspen clones and increment core analysis to determine tree age. Clonal dimensions were measured by the maximum distance between two ramets and the number of ramets per genet. Standard normal deviate (SND) was used to assess the spatial distribution of aspen genets and cohorts, and multivariate spatial genetic autocorrelations to assess the spatial distribution of aspen genetic variation. Most aspen genets consisted of only one ramet (> 75%). Median clonal dimensions were 19 and 29 m (maxima: 104 and 72 m in the two plots). No segregation was observed between clones. Aspen cohorts were spatially segregated but trees were spatially aggregated within old and medium-aged cohorts. In contrast, trees were more randomly distributed within the youngest cohorts. This coincided with a spatial genetic autocorrelation at small scales (up to 30 m) in the older cohorts and a more random genetic distribution in the youngest ones. Our results suggest that aspen spatial genetic structuring reflects the spatial patterns produced by the regeneration of discrete cohorts at different stages of succession. Vegetative reproduction leads to aspen genetic spatial structuring at small scales (few metres) until midsuccession. However, as the stand gets older, the spatial distribution of aspen trees and genetic structure evolve from a structured pattern to a more random one under a gap disturbances regime.     

Pit Membrane Degradation and Air-Embolism Formation in Ageing Xylem Vessels of Populus tremuloides Michx

Air-embolism formation in xylem vessels of Populus tremuloidesMichx. was quantified by its reduction of hydraulic conductivityin branch segments. Embolism was induced by increasing xylemtension in drying stems, or by inserting one end of a hydratedstem in a pressure bomb and increasing air pressure in the bomb.Both treatments produced the same response suggesting that embolismby water stress was caused by air entering water-filled vessels,presumably through inter-vessel pits. In rapidly-growing P.tremuloides branches, the vessels of the outer growth ring werefunctional whereas vessels in older xylem were mostly embolized.This selective embolizing of older vessels was associated witha marked increase in permeability of their inter-vessel pitsto air, relative to pits of younger vessels. Air-injection pressuresless than 1·0 MPa completely embolized older vesselsthat had been re-filled in the laboratory, whereas pressuresover 4·0 MPa were required to embolize young vessels.Greater permeability of old vessels was due to degradation oftheir pit membranes as seen in the scanning electron microscope;large openings were present that were not seen in pit membranesof young vessels. These holes would allow air to penetrate vesselends at low pressure differences causing embolism. Degradationof pit membranes causing the selective dysfunction of oldersapwood may be a general phenomenon initiating heartwood formationin many species. Key words: Xylem embolism, hydraulic conductivity, heartwood formation, cavitation, Populus tremuloides, Michx     

An Examination of the Leaf Quaking Adaptation and Stomatal Distribution in Populus tremuloides Michx

The leaves of quaking aspen (Populus tremuloides Michx.) have a flattened petiole that allows them to quake (oscillate and roll) under low wind velocities. It was hypothesized that this adaptation might enable the plant to respond to windy conditions that would increase transpirational losses. No effects of wind with or without leaf quaking on stomatal resistance were observed under controlled conditions in the field. If wind and leaf quaking affect stomatal resistance, such effects must be small in comparison to those caused by other factors such as leaf water potential and ambient humidity.     

High efficiency shoot regeneration from callus of quaking aspen (Populus tremuloides Michx.)

Callus was induced from leaf segments of aspen (Populus tremuloides Michx.) on modified B5 (mB5) medium with 0.1 mg/1 benzyladenine (BA) and 0.5 mg/1 2,4-dichlorophenoxyacetic acid (2,4-D). The resulting callus was either subcultured to solidified Woody Plant Medium (WPM) with 0.5 mg/1 BA directly for shoot regeneration or sieved into liquid mB5 medium for suspension culture. After 3 weeks of suspension culture, when the callus clumps grew to 3–4 mm in diameter, they were transferred onto solidified WPM with 0.5 mg/1 BA for shoot regeneration. Almost 100% of the clumps formed shoots on WPM when subcultured directly from mB5 with an average number of 6 shoots per callus. When transferred from suspension culture in mB5 to WPM, an average of 6 shoots per callus were produced from 51% of calli. These shoots could be easily rooted on either mB5 or WPM with 0.2 mg/1 indole-3-butyric acid (IBA) and transferred to pots. Transplanted plants were kept under intermittent mist for 2–4 weeks before normal growth in the green house.Abbreviations BA 6-Benzyl-adenine - IBA indole-3-butyric acid - 2,4-D 2,4-dichlorophenoxyacetic acid - mB5 medium modified B5 medium - WPM Woody Plant medium     

The Role of Microsite Conditions in Restoring Trembling Aspen (Populus tremuloides Michx) from Seed

Encouraging natural regeneration of Populus tremuloides Michx (trembling aspen) from seed is a largely unexplored means for reintroducing the species into reclamation areas. We evaluated the effects of microsite (surface contour and substrate type) on aspen seedling establishment and growth on a reclaimed coal mine. The 4.6 ha study site was divided into six 48 m‐wide strips that had 15 or 40 cm capping material salvaged from a nearby forest floor added to the mine surface. We surveyed 126 m long transects located in the center of each strip for microsite conditions, and the presence and height of aspen seedlings. We found that aspen seedlings generally preferred mineral‐organic substrates and concave microsites. To facilitate the regeneration of aspen by seed, we suggest that land managers increase small‐scale roughness and microtopographic diversity on reclaimed sites .     

Phenology and climate relationships in aspen (Populus tremuloides Michx.) forest and woodland communities of southwestern Colorado

Trembling aspen (Populus tremuloides Michx.) occurs over wide geographical, latitudinal, elevational, and environmental gradients, making it a favorable candidate for a study of phenology and climate relationships. Aspen forests and woodlands provide numerous ecosystem services, such as high primary productivity and biodiversity, retention and storage of environmental variables (precipitation, temperature, snow–water equivalent) that affect the spring and fall phenology of the aspen woodland communities of southwestern Colorado. We assessed the land surface phenology of aspen woodlands using two phenology indices, start of season time (SOST) and end of season time (EOST), from the U.S. Geological Survey (USGS) database of conterminous U.S. phenological indicators over an 11-year time period (2001–2011). These indicators were developed with 250 m resolution remotely sensed data from the Moderate Resolution Imaging Spectroradiometer processed to highlight vegetation response. We compiled data on SOST, EOST, elevation, precipitation, air temperature, and snow water equivalent (SWE) for selected sites having more than 80% cover by aspen woodland communities. In the 11-year time frame of our study, EOST had significant positive correlation with minimum fall temperature and significant negative correlation with fall precipitation. SOST had a significant positive correlation with spring SWE and spring maximum temperature.     

Populations of aspen (Populus tremuloides Michx.) with different evolutionary histories differ in their climate occupancy

Quaking aspens (Populus tremuloides Michx.) are found in diverse habitats throughout North America. While the biogeography of aspens' distribution has been documented, the drivers of the phenotypic diversity of aspen are still being explored. In our study, we examined differences in climate between northern and southwestern populations of aspen, finding large‐scale differences between the populations. Our results suggest that northern and southwestern populations live in distinct climates and support the inclusion of genetic and phenotypic data with species distribution modeling for predicting aspens' distribution.     

Effects of genotype and nutrient availability on phytochemistry of trembling aspen (Populus tremuloides Michx.) during leaf senescence

We documented temporal patterns in phytochemical composition of Populus tremuloides Michx. during leaf senescence, and the influence of genotype and soil nutrient availability on such patterns. Levels of foliar nitrogen, carbohydrates, phenolic glycosides and condensed tannins were quantified for four aspen genotypes grown in a common garden, with low and high levels of soil nutrients. Levels of all compounds tended to decline over time, although the magnitude of change was influenced by plant genotype and nutrient availability. Genetic variation in concentrations of phytochemicals was much greater for phenolic glycosides and tannins than for nitrogen and carbohydrates, and these phenolic signatures generally persisted through leaf abscission. Our results suggest that genotypic and nutrient effects on patterns of chemical change during senescence will likely influence the performance of late-season herbivores on aspen. Moreover, nutrient and especially genotypic variation in phytochemistry of abscised leaves is likely to affect litter decomposition rates.     

A Novel Metabolic Pathway for Indole-3-Acetic Acid in Apical Shoots of Populus tremula (L.) x Populus tremuloides (Michx.)

Metabolism of indole-3-acetic acid (IAA) in apical shoots of Populus tremula (L.) x Populus tremuloides (Michx.) was investigated by feeding a mixture of [12C]IAA, [13C6]IAA, and [1[prime]-14C]IAA through the base of the excised stem. HPLC of methanolic plant extracts revealed eight major radiolabeled metabolites after a 24-h incubation period. Comparison between feeds with [5-3H]IAA and [1[prime]-14C]IAA showed that all detectable metabolites were nondecarboxylative products. The purified radiolabeled HPLC fractions were screened by frit-fast atom bombardment liquid chromatography-mass spectrometry for compounds with characteristic fragment pairs originating from the application with 12C and 13C isotopes. Samples of interest were further characterized by gas chromatography-mass spectrometry. Using this procedure, oxindole-3-acetic acid (OxIAA), indole-3-acetyl-N-aspartic acid (IAAsp), oxindole-3-acetyl-N-aspartic acid (OxIAAsp), and ring-hydroxylated oxindole-3-acetic acid were all identified as IAA metabolites. Furthermore, a novel metabolic pathway from IAA via IAAsp and OxIAAsp to OxIAA was established on the basis of refeeding experiments with the different IAA metabolites.     

The localization of pentosans within the cell wall of aspen (Populus tremuloides Michx.) by high resolution autoradiography

Summary Radiochemical studies of Populus tremuloides xylem tissue administered l-[1-³H]arabinose, d-[1-³H]glucose, and d-[6-³H]glucose demonstrate that l-[1-³H]arabinose is an excellent precursor for pentosan in this tissue. Transverse sections of first-year xylem (from cambial zone to pith) were examined by light and electron microscope autoradiography. Relatively large amounts of labeled pentosan are found in parenchyma cell walls, including the protective layer of ray parenchyma. Computeraided analyses of grain distributions in electron micrographs of cell walls of individual fibers localized the labeled wall components after different periods of incubation by comparison to model behavior. These analyses indicate that pentosan is added to the secondary cell wall of developing fibers by an appositional mechanism.     

Microsatellite DNA markers in Populus tremuloides.

Markers for eight new microsatellite DNA or simple sequence repeat (SSR) loci were developed and characterized in trembling aspen (Populus tremuloides) from a partial genomic library. Informativeness of these microsatellite DNA markers was examined by determining polymorphisms in 38 P. tremuloides individuals. Inheritance of selected markers was tested in progenies of controlled crosses. Six characterized SSR loci were of dinucleotide repeats (two perfect and four imperfect), and one each of trinucleotide and tetranucleotide repeats. The monomorphic SSR locus (PTR15) was of a compound imperfect dinucleotide repeat. The primers of one highly polymorphic SSR locus (PTR7) amplified two loci, and alleles could not be assigned to a specific locus. At the other six polymorphic loci, 25 alleles were detected in 38 P. tremuloides individuals; the number of alleles ranged from 2 to 7, with an average of 4.2 alleles per locus, and the observed heterozygosity ranged from 0.05 to 0.61, with an average of 0.36 per locus. The two perfect dinucleotide and one trinucleotide microsatellite DNA loci were the most informative. Microsatellite DNA variants of four SSR loci characterized previously followed a single-locus Mendelian inheritance pattern, whereas those of PTR7 from the present study showed a two-locus Mendelian inheritance pattern in controlled crosses. The microsatellite DNA markers developed and reported here could be used for assisting various genetic, breeding, biotechnology, genome mapping, conservation, and sustainable forest management programs in poplars.     

杨树人工林凋落物养分归还功能研究

对杨树中龄林、成熟林凋落物量、组成特征、季节动态及凋落物中N、P、K、Fe、Ca、Mg含量进行了定位观测和实验测定。结果表明:杨树成熟林的年凋落物量为4.45t·hm^-2·a^-1,是中龄林的1.42倍,两者存在显著差异(p<0.05);叶是凋落物的主要形式,分别占中龄林、成熟林总凋落物量的70.1%和81.5%;凋落物量具有明显的季节动态,表现为"双峰"型;成熟林养分归还量是68.4kg·hm^-2·a^-1,是中龄林48.96kg·hm^-2·a^-1的1.4倍;两林分各养分年归还量的大小顺序为N>Ca>K>Mg>Fe>P;杨树成熟林的N、P、Fe、Mg4种元素的利用率大于中龄林的,而K和Ca元素的利用率却相反。     

Sulfide-Resistant Respiration in Leaves of Elodea canadensis Michx: Comparison with Cyanide-Resistant Respiration

The rate of dark O₂ uptake of Elodea canadensis leaves was titrated with either cyanide or sulfide in the presence and in the absence of 5 millimolar salicylhydroxamic acid (SHAM), an inhibitor of the alternative oxidase. The inhibition of O₂ uptake by SHAM alone was very small (3-6%), suggesting that actual respiration mainly occurred through the cytochrome pathway. O₂ uptake was slightly stimulated by cyanide at concentrations of 50 micromolar or higher, but in the presence of SHAM respiration was strongly suppressed. The effects of sulfide on O₂ uptake were similar to those of cyanide, except that the percent stimulation of O₂ uptake by sulfide alone was somewhat higher than that of cyanide. However, the estimates of the capacity of the alternative pathway were similar with both inhibitors. Another difference is that maximal inhibition of respiration in the presence of SHAM was observed with lower concentrations of sulfide (50 micromolar) than cyanide (250 micromolar). The results suggest that sulfide can be used as a suitable inhibitor of cytochrome c oxidase in studies with intact plant tissues, and that sulfide does not apparently inhibit the alternative oxidase.