Barb R. Thomas

Clone history shapes Populus drought responses

Just as animal monozygotic twins can experience different environmental conditions by being reared apart, individual genetically identical trees of the genus Populus can also be exposed to contrasting environmental conditions by being grown in different locations. As such, clonally propagated Populus trees provide an opportunity to interrogate the impact of individual environmental history on current response to environmental stimuli. To test the hypothesis that current responses to an environmental stimulus, drought, are contingent on environmental history, the transcriptome- level drought responses of three economically important hybrid genotypes—DN34 (Populus deltoides × Populus nigra), Walker [P. deltoides var. occidentalis × (Populus laurifolia × P. nigra)], and Okanese [Walker × (P. laurifolia × P. nigra)]—derived from two different locations were compared. Strikingly, differences in transcript abundance patterns in response to drought were based on differences in geographic origin of clones for two of the three genotypes. This observation was most pronounced for the genotypes with the longest time since establishment and last common propagation. Differences in genome-wide DNA methylation paralleled the transcriptome level trends, whereby the clones with the most divergent transcriptomes and clone history had the most marked differences in the extent of total DNA methylation, suggesting an epigenomic basis for the clone history-dependent transcriptome divergence. The data provide insights into the interplay between genotype and environment in the ecologically and economically important Populus genus, with implications for the industrial application of Populus trees and the evolution and persistence of these important tree species and their associated hybrids.

The mycorrhizal status of selected poplar clones introduced in Alberta

Abstract Mycorrhizal associations are essential in the establishment and growth of plant species. The types of mycorrhizas were investigated in the roots of poplar clones introduced in Alberta. We sampled roots of 28 poplar clones from a 5-year old trial established at the Alberta-Pacific Forest Industries Inc. mill site located 40 km North of Boyle, Alberta. The two most common types of mycorrhizas were quantified in plant roots. The poplar clones showed variable degrees of colonization by both ectomycorrhizal and arbuscular mycorrhizal fungi, suggesting differential host susceptibility. This highlights the potential use of selected strains of both ectomycorrhizal and arbuscular mycorrhizal fungi for inoculation trials in populiculture to improve the establishment of introduced poplars on to previously cleared agricultural or disturbed sites in the province of Alberta in particular and in the Canadian prairie provinces in general.

Genetic variation of hydraulic and wood anatomical traits in hybrid poplar and trembling aspen

Intensive forestry systems and breeding programs often include either native aspen or hybrid poplar clones, and performance and trait evaluations are mostly made within these two groups. Here, we assessed how traits with potential adaptive value varied within and across these two plant groups. Variation in nine hydraulic and wood anatomical traits as well as growth were measured in selected aspen and hybrid poplar genotypes grown at a boreal planting site in Alberta, Canada. Variability in these traits was statistically evaluated based on a blocked experimental design. We found that genotypes of trembling aspen were more resistant to cavitation, exhibited more negative water potentials, and were more water-use-efficient than hybrid poplars. Under the boreal field test conditions, which included major regional droughts, height growth was negatively correlated with branch vessel diameter (Dv ) in both aspen and hybrid poplars and differences in Dv were highly conserved in aspen trees from different provenances. Differences between the hybrid poplars and aspen provenances suggest that these two groups employ different water-use strategies. The data also suggest that vessel diameter may be a key trait in evaluating growth performance in a boreal environment.

Intraspecific variation in the Populus balsamifera drought transcriptome

Drought is a major limitation to the growth and productivity of trees in the ecologically and economically important genus Populus. The ability of Populus trees to contend with drought is a function of genome responsiveness to this environmental insult, involving reconfiguration of the transcriptome to appropriately remodel growth, development and metabolism. Here we test hypotheses aimed at examining the extent of intraspecific variation in the drought transcriptome using six different Populus balsamifera L. genotypes and Affymetrix GeneChip technology. Within a given genotype there was a positive correlation between the magnitude of water-deficit induced changes in transcript abundance across the transcriptome, and the capacity of that genotype to maintain growth following water deficit. Genotypes that had more similar drought-responsive transcriptomes also had fewer genotypic differences, as determined by microarray-derived single feature polymorphism (SFP) analysis, suggesting that responses may be conserved across individuals that share a greater degree of genotypic similarity. This work highlights the fact that a core species-level response can be defined; however, the underpinning genotype-derived complexities of the drought response in Populus must be taken into consideration when defining both species- and genus-level responses.

A comparison of pre-planting treatments on hardwood cuttings of four hybrid poplar clones

Rooting and early growth of four hybrid poplar clones (Populus spp.) planted in a greenhouse were examined after applying 40 pre-rooting treatment combinations to dormant cuttings. Treatments included 2 cutting lengths (5 and 10 cm), 5 soaking times (0, 2, 4, 8, and 12 days), and 4 dips (chitosan, rooting hormone powder, liquid rooting hormone added to the soaking water, and none). Significant differences in both rooting percentages and growth were shown between clones after 7 weeks of growth. Ten cm cuttings had 29% greater rooting success, 28% more above-ground growth, and 12% lower root/shoot ratios than 5 cm cuttings. Cuttings planted without soaking had the lowest rooting success, at less than 45% on average. Commercial rooting hormones decreased the number of rooted cuttings but increased root/shoot ratios. For optimal rooting, we recommend using 10 cm cuttings, soaked for 2 days in water (4 days for the Jackii10 clone) without any additional dipping/hormone substance.

Drought induces alterations in the stomatal development program in Populus

Much is known about the physiological control of stomatal aperture as a means by which plants adjust to water availability. By contrast, the role played by the modulation of stomatal development to limit water loss has received much less attention. The control of stomatal development in response to water deprivation in the genus Populus is explored here. Drought induced declines in stomatal conductance as well as an alteration in stomatal development in two genotypes of Populus balsamifera. Leaves that developed under water-deficit conditions had lower stomatal indices than leaves that developed under well-watered conditions. Transcript abundance of genes that could hypothetically underpin drought-responsive changes in stomatal development was examined, in two genotypes, across six time points, under two conditions, well-watered and with water deficit. Populus homologues of STOMAGEN, ERECTA (ER), STOMATA DENSITY AND DISTRIBUTION 1 (SDD1), and FAMA had variable transcript abundance patterns congruent with their role in the modulation of stomatal development in response to drought. Conversely, there was no significant variation in transcript abundance between genotypes or treatments for the Populus homologues of YODA (YDA) and TOO MANY MOUTHS (TMM). The findings highlight the role that could be played by stomatal development during leaf expansion as a longer term means by which to limit water loss from leaves. Moreover, the results point to the key roles played by the regulation of the homologues of STOMAGEN, ER, SDD1, and FAMA in the control of this response in poplar.

Frost hardiness vs. growth performance in trembling aspen: an experimental test of assisted migration

Summary 1. According to the range limit hypothesis, the distribution of many temperate species is restricted by a trade-off between their capacity to survive winter extremes in the north (or high elevation) and their ability to compete with better-adapted species in the south (or low elevation range limits). This trade-off has important implications in forestry, particularly in the context of managed seed movement under climate change. 2. In this study, we aim to quantify trade-offs among growth, frost hardiness and timing of leaf senescence and bud break in populations of trembling aspen, Populus tremuloides Michx., which were observed in a large reciprocal transplant experiment across five planting sites in western Canada, including additional provenances from Minnesota. 3. After 10 years, we found pronounced increases in productivity as a result of long-distance transfers in a north-west direction. For example, provenances moved 1600 km north-west from Minnesota to central Alberta (a shift of 7° latitude to the north) produced almost twice the biomass of local sources. Similarly, provenances moved 800 km from central Alberta to north-east British Columbia (4° latitude north) also produced twice the biomass of local sources. 4. We further found that increased growth was not associated with lower survival rates. Bud break in provenances transferred north-west generally occurred slightly later than in local sources, suggesting decreased risk of spring frost injury. Leaf abscission was later in provenances transferred in a north-west direction, but they appeared to be very frost hardy, well ahead of very rare early fall frost events. 5. Synthesis and applications. This study demonstrated that assisted migration prescriptions have considerable potential to enhance forest productivity. In the case of aspen, even longdistance seed transfers in a north-west direction were successful. We conclude that benefits in productivity outweigh potential risks associated with northward transfer of aspen planting stock under both current and projected future climate conditions.

Stomatal factors and vulnerability of stem xylem to cavitation in poplars

The relationships between the vulnerability of stem xylem to cavitation, stomatal conductance, stomatal density, and leaf and stem water potential were examined in six hybrid poplar (P38P38, Walker, Okanese, Northwest, Assiniboine and Berlin) and balsam poplar (Populus balsamifera) clones. Stem xylem cavitation resistance was examined with the Cavitron technique in well-watered plants grown in the greenhouse. To investigate stomatal responses to drought, plants were subjected to drought stress by withholding watering for 5 (mild drought) and 7 (severe drought) days and to stress recovery by rewatering severely stressed plants for 30 min and 2 days. The clones varied in stomatal sensitivity to drought and vulnerability to stem xylem cavitation. P38P38 reduced stomatal conductance in response to mild stress while the balsam poplar clone maintained high leaf stomatal conductance under more severe drought stress conditions. Differences between the severely stressed clones were also observed in leaf water potentials with no or relatively small decreases in Assiniboine, P38P38, Okanese and Walker. Vulnerability to drought-induced stem xylem embolism revealed that balsam poplar and Northwest clones reached loss of conductivity at lower stem water potentials compared with the remaining clones. There was a strong link between stem xylem resistance to cavitation and stomatal responsiveness to drought stress in balsam poplar and P38P38. However, the differences in stomatal responsiveness to mild drought suggest that other drought-resistant strategies may also play a key role in some clones of poplars exposed to drought stress.

Sixteen years of winter stress: an assessment of cold hardiness, growth performance and survival of hybrid poplar clones at a boreal planting site

In recent years, thousands of hectares of hybrid poplar plantations have been established in Canada for the purpose of carbon sequestration and wood production. However, boreal planting environments pose special challenges that may compromise the long-term survival and productivity of such plantations. In this study, we evaluated the effect of winter stress, that is, frequent freeze-thaw and extreme cold events, on growth and survival of 47 hybrid poplar clones in a long-term field experiment. We further assessed physiological and structural traits that are potentially important for cold tolerance for a selected set of seven clones. We found that trees with narrow xylem vessels showed reduced freezing-induced embolism and showed superior productivity after 16 growing seasons. With respect to cold hardiness of living tissues, we only observed small differences among clones in early autumn, which were nonetheless significantly correlated to growth. Maximum winter cold hardiness and the timing of leaf senescence and budbreak were not related to growth or survival. In conclusion, our data suggest that reduction of freezing-induced embolism due to small vessel diameters is an essential adaptive trait to ensure long-term productivity of hybrid poplar plantations in boreal planting environments.

The interaction between nitrogen source, soil pH, and drought in the growth and physiology of three poplar clonesThis article is one of a selection of papers published in the Special Issue on Poplar Research in Canada.

The objective of this study was to understand how nitrogen (N) source, soil pH, and drought interact in the growth and water-use efficiency of poplars. Rooted cuttings from two hybrid poplar clones...