ALBERT

Description: Non-structural carbohydrate (NSC) pools fluctuate based on the interplay between photosynthesis, demand from various carbon (C) sinks and tree hydraulic status. Thus, it has been hypothesized that tree species with isohydric stomatal control (i.e., trees that close stomata rapidly in response to drought) rely heavily on NSC pools to sustain metabolism, which can lead to negative physiological consequences such as C depletion. Here, we seek to use a species’ degree of isohydry or anisohydry as a conceptual framework for understanding the interrelations between photosynthetic C supply, hydraulic damage and fluctuations in NSC pools. We conducted a 6-week experimental drought, followed by a 6-week recovery period, in a greenhouse on seven tree species that span the spectrum from isohydric to anisohydric. Throughout the experiment, we measured photosynthesis, hydraulic damage and NSC pools. Non-structural carbohydrate pools were remarkably stable across species and tissues—even highly isohydric species that drastically reduced C assimilation were able to maintain stored C. Despite these static NSC pools, we still inferred an important role for stored C during drought, as most species converted starches into sugars during water stress (and back again post-drought). Finally, we did not observe any linkages between C supply, hydraulic damage and NSC pools, indicating that NSC was maintained independent of variation in photosynthesis and hydraulic function. Our results advance the idea that C depletion is a rare phenomenon due to either active maintenance of NSC pools or sink limitation, and thus question the hypothesis that reductions in C assimilation necessarily lead to C depletion.

Description: To study the effects of slightly elevated temperature and ozone (O3) on leaf structural characteristics of silver birch (Betula pendula Roth), saplings of four clonal genotypes of this species were exposed to elevated temperature (ambient air temperature +0.8–1.0 °C) and elevated O3 (1.3–1.4× ambient O3), alone and in combination, in an open-air exposure field over two growing seasons (2007 and 2008). So far, the impacts of moderate elevation of temperature or the combination of elevated temperature and O3 on leaf structure of silver birch have not been intensively studied, thus showing the urgent need for this type of studies. Elevated temperature significantly increased leaf size, reduced non-glandular trichome density, decreased epidermis thickness and increased plastoglobuli size in birch leaves during one or both growing seasons. During the second growing season, O3 elevation reduced leaf size, increased palisade layer thickness and decreased the number of plastoglobuli in spongy cells. Certain leaf structural changes observed under a single treatment of elevated temperature or O3, such as increase in the amount of chloroplasts or vacuole, were no longer detected at the combined treatment. Leaf structural responses to O3 and rising temperature may also depend on timing of the exposure during the plant and leaf development as indicated by the distinct changes in leaf structure along the experiment. Genotype-dependent cellular responses to the treatments were detected particularly in the palisade cells. Overall, this study showed that even a slight but realistic elevation in ambient temperature can notably modify leaf structure of silver birch saplings. Leaf structure, in turn, influences leaf function, thus potentially affecting acclimation capacity under changing climate.

Description: Scots pine (Pinus sylvestris L.) is one of the most important conifers in Northern Europe. In boreal forests, over one-third of net primary production is allocated to roots. Pioneer roots expand the horizontal and vertical root systems and transport nutrients and water from belowground to aboveground. Fibrous roots, often colonized by mycorrhiza, emerge from the pioneer roots and absorb water and nutrients from the soil. In this study, we installed three flatbed scanners to detect the daily growth of both pioneer and fibrous roots of Scots pine during the growing season of 2018, a year with an unexpected summer drought in Southern Finland. The growth rate of both types of roots had a positive relationship with temperature. However, the relations between root elongation rate and soil moisture differed significantly between scanners and between root types indicating spatial heterogeneity in soil moisture. The pioneer roots were more tolerant to severe environmental conditions than the fibrous roots. The pioneer roots initiated elongation earlier and ceased it later than the fibrous roots. Elongation ended when the temperature dropped below the threshold temperature of 4 °C for pioneer roots and 6 °C for fibrous roots. During the summer drought, the fibrous roots halted root surface area growth at the beginning of the drought, but there was no drought effect on the pioneer roots over the same period. To compare the timing of root production and the aboveground organs’ production, we used the CASSIA model, which estimates the aboveground tree carbon dynamics. In this study, root growth started and ceased later than growth of aboveground organs. Pioneer roots accounted for 87% of total root productivity. We suggest that future carbon allocation models should separate the roots by root types (pioneer and fibrous), as their growth patterns are different and they have different reactions to changes in the soil environment.

Description: Climate change will increase the risk of flooding in several areas of the world where Populus deltoides Marshall (eastern cottonwood) is planted, so it would be desirable for this species to select for flooding tolerance. The aims of this work were to explore the variability in growth, leaf traits and flooding tolerance in an F1 full-sib intraspecific progeny of P. deltoides, to analyze the correlations of leaf and growth traits with flooding tolerance and to assess their suitability for use in breeding programs. Two-month-old parental clones and their progeny of 30 full-sib F1 genotypes were grown in pots and subjected to two treatments: (i) plants watered to field capacity (control) and (ii) plants flooded up to 10 cm above soil level for 35 days. Growth (height, diameter and biomass partition) and leaf traits (leaf size and number, specific leaf area, leaf senescence, abscission, stomatal conductance, carbon isotope discrimination, stomatal index) were measured. Flooding tolerance for each genotype was estimated as the ratio of the biomass of stressed plants to the biomass of control plants. Results showed segregation in terms of flooding tolerance in the F1 progeny. A significant genotype effect was found for leaf size and number, carbon isotopic discrimination and stomatal conductance, but it did not correlate with flooding tolerance. Height, diameter and root-to-shoot ratio had a positive phenotypic correlation with flooding tolerance, and there was a positive genetic correlation of height and diameter with biomass on both treatments. The narrow sense heritability values for the traits analyzed ranged from 0 to 0.56. We conclude that growth traits are more adequate than leaf traits for selection to increase flooding tolerance. A vigorous initial growth would increase flooding tolerance in young poplar plants.

Description: Vascular plants have two types of water-conducting cells, xylem vessel cells (in angiosperms) and tracheid cells (in ferns and gymnosperms). These cells are commonly characterized by secondary cell wall (SCW) formation and programmed cell death (PCD), which increase the efficiency of water conduction. The differentiation of xylem vessel cells is regulated by a set of NAC (NAM, ATAF1/2 and CUC2) transcription factors, called the VASCULAR-RELATED NAC-DOMAIN (VND) family, in Arabidopsis thaliana Linne. The VNDs regulate the transcriptional induction of genes required for SCW formation and PCD. However, information on the transcriptional regulation of tracheid cell differentiation is still limited. Here, we performed functional analysis of loblolly pine (Pinus taeda Linne) VND homologs (PtaVNS, for VND, NST/SND, SMB-related protein). We identified five PtaVNS genes in the loblolly pine genome, and four of these PtaVNS genes were highly expressed in tissues with tracheid cells, such as shoot apices and developing xylem. Transient overexpression of PtaVNS genes induced xylem vessel cell-like patterning of SCW deposition in tobacco (Nicotiana benthamiana Domin) leaves, and up-regulated the promoter activities of loblolly pine genes homologous to SCW-related MYB transcription factor genes and cellulose synthase genes, as well as to cysteine protease genes for PCD. Collectively, our data indicated that PtaVNS proteins possess transcriptional activity to induce the molecular programs required for tracheid formation, i.e., SCW formation and PCD. Moreover, these findings suggest that the VNS–MYB-based transcriptional network regulating water-conducting cell differentiation in angiosperm and moss plants is conserved in gymnosperms.

Description: Most bamboo species including Moso bamboo (Phyllostachys edulis) are tropical or subtropical plants that greatly contribute to human well-being. Low temperature is one of the main environmental factors restricting bamboo growth and geographic distribution. Our knowledge of the molecular changes during bamboo adaption to cold stress remains limited. Here, we provided a general overview of the cold-responsive transcriptional profiles in Moso bamboo by systematically analyzing its transcriptomic response under cold stress. Our results showed that low temperature induced strong morphological and biochemical alternations in Moso bamboo. To examine the global gene expression changes in response to cold, 12 libraries (non-treated, cold-treated 0.5, 1 and 24 h at −2 °C) were sequenced using an Illumina sequencing platform. Only a few differentially expressed genes (DEGs) were identified at early stage, while a large number of DEGs were identified at late stage in this study, suggesting that the majority of cold response genes in bamboo are late-responsive genes. A total of 222 transcription factors from 24 different families were differentially expressed during 24-h cold treatment, and the expressions of several well-known C-repeat/dehydration responsive element-binding factor negative regulators were significantly upregulated in response to cold, indicating the existence of special cold response networks. Our data also revealed that the expression of genes related to cell wall and the biosynthesis of fatty acids were altered in response to cold stress, indicating their potential roles in the acquisition of bamboo cold tolerance. In summary, our studies showed that both plant kingdom-conserved and species-specific cold response pathways exist in Moso bamboo, which lays the foundation for studying the regulatory mechanisms underlying bamboo cold stress response and provides useful gene resources for the construction of cold-tolerant bamboo through genetic engineering in the future.

Description: Trees of the genus Prunus produce some of the most widely consumed fruits globally. The combination of climate change-related warming and increased drought stress, scarcity of freshwater resources for irrigation, and increasing demands due to population growth creates a need for increased drought tolerance in these tree species. Recently, we have shown in the field that a native wild pear species performs better under drought than two cultivated pear species. Here, a comparative field study was conducted in Israel to investigate traits associated with drought tolerance in almond (cultivated Prunus dulcis (Mill.) D. A. Webb vs wild Prunus ramonensis Danin) and plum (cultivated Prunus domestica L. vs wild Prunus ursina Kotschy). Measurements of xylem embolism and shoot and root carbon reserves were done along a year, including seasonal drought in the wild and a 35-day drought experiment in the orchards. Synchronous measurements of native xylem embolism and shoot water potential showed that cultivated and wild almond trees lost ~50% of hydraulic conductivity at −2.3 and −3.2 MPa, respectively. Micro-CT images confirmed the higher embolism ratio in cultivated versus wild almond, whereas the two plum species were similar. Dynamics of tissue concentrations of nonstructural carbohydrates were mostly similar across species, with higher levels in cultivated versus wild plum. Our results indicate an advantage for the wild almond over its cultivated relative in terms of xylem resistance to embolism, a major risk factor for trees under drought stress. This result is in line with our previous experiment on pear species. However, the opposite trends observed among the studied plum species mean that these trends cannot be generalized. It is possible that the potential for superior drought tolerance in wild tree species, relative to their cultivated relatives, is limited to wild species from dry and hot habitats.

Description: WRKY transcription factors, one of the largest transcription factor families, play important roles in regulating the synthesis of secondary metabolites. In sweet osmanthus (Osmanthus fragrans), the monoterpenes have been demonstrated as the most important volatile compounds, and the W-box, which is the cognate binding site of WRKY transcription factors, could be identified in most of the terpene-synthesis-related genes’ promoters. However, the role of the WRKY family in terpene synthesis in sweet osmanthus has rarely been examined. In this study, 154 WRKY genes with conserved WRKY domain were identified and classified into three groups. The group II was further divided into five subgroups, and almost all members of IId contained a plant zinc cluster domain. Eight OfWRKYs (OfWRKY7/19/36/38/42/84/95/139) were screened from 20 OfWRKYs for their flower-specific expression patterns in different tissues. Simultaneously, the expression patterns of OfWRKYs and emission patterns of volatile compounds during the flowering process were determined and gas chromatography-mass spectrometry results showed that monoterpenes, such as linalool and ocimene, accounted for the highest proportion, contributing to the floral scent of sweet osmanthus in two cultivars. In addition, correlation analysis revealed the expression patterns of OfWRKYs (OfWRKY7/19/36/139) were each correlated with distinct monoterpenes (linalool, linalool derivatives, ocimene and ocimene derivatives). Subcellular localization analysis showed that p35S::GFP–OfWRKY7/38/95/139 were localized in the nucleus and OfWRKY139 had very strong transactivation activity. Collectively, the results indicated potential roles of OfWRKY139 and OfWRKYs with plant zinc cluster domain in regulating synthesis of aromatic compounds in sweet osmanthus, laying the foundation for use of OfWRKYs to improve the aroma of ornamental plants.

Description: Drought and nitrogen (N) addition have been shown to affect tree hydraulic traits, but few studies have been made on their interactions across species with different wood types or leaf forms. We examined the responses of hydraulic conductance and xylem anatomical traits of Quercus mongolica (ring porous with simple leaves), Fraxinus mandshurica (ring porous with compound leaves) and Tilia amurensis (diffuse porous with simple leaves) to drought, N addition and their interactions. Drought stress decreased current-year xylem-specific conductivity in stems (Ksx) and leaf hydraulic conductance (Kleaf ), but N addition affected Ksx and Kleaf differently among species and watering regimes. These divergent effects were associated with different responses of anatomical traits and leaf forms. Higher mean vessel diameter in stems and lower vessel density in leaves were observed with N addition. The three-way interactive effects of drought, N addition and tree species were significant for most values of anatomical traits. These results were also reflected in large differences in vessel diameter and density among species with different wood types or leaf forms. The two-way interactive effects of drought and N addition were significant on Kleaf and predawn water potential, but not Ksx, indicating that leaves were more sensitive than stems to a combination of drought stress and N addition. Our results provide mechanistic insight into the variable responses of xylem water transport to the interactions of drought and N availability.

Description: A critical process that allows multiple, similar species to coexist in an ecological community is their ability to partition local habitat gradients. The mechanisms that underlie this separation at local scales may include niche differences associated with their biogeographic history, differences in ecological function associated with the degree of shared ancestry and trait-based performance differences, which may be related to spatial or temporal variation in habitat. In this study we measured traits related to water-use, growth and stress tolerance in mature trees and seedlings of three oak species (Quercus alba L., Quercus falcata Michx. and Quercus palustris Münchh). which co-occur in temperate forests across the eastern USA but tend to be found in contrasting hydrologic environments. The three species showed significant differences in their local distributions along a hydrologic gradient. We tested three possible mechanisms that influence their contrasting local environmental distributions and promote their long-term co-existence: (i) differences in their climatic distributions across a broad geographic range, (ii) differences in functional traits related to water use, drought tolerance and growth and (iii) contrasting responses to temporal variation in water availability. We identified key differences between the species in both their range-wide climatic distributions (especially aridity index and mean annual temperature) and physiological traits in mature trees and seedlings, including daily water loss, hydraulic conductance, stress responses, growth rate and biomass allocation. Taken together, these differences explain the habitat partitioning that allows three closely related species to co-occur locally.