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  Background To investigate the salt tolerance of Populus talassica  ×  Populus euphratica , morphological and physiological parameters were measured on the second day after the 15th, 30th and 45th days of NaCl treatment, revealing significant effects of NaCl on growth. To further elucidate the mechanisms underlying salt tolerance, transcriptomic and metabolomic analysis were conducted under different NaCl treatments. Results The results of morphological and physiological indexes showed that under low salt treatment, P. talassica  ×  P. euphratica was able to coordinate the growth of aboveground and belowground parts. Under high salt concentration, the growth and water balance of P. talassica  ×  P. euphratica were markedly inhibited. The most significant differences between treatments were observed on the second day after the 45th day of NaCl treatment. Transcriptomic analysis showed that the pathways of gene enrichment in the roots and stems of P. talassica  ×  P. euphratica were different in the salt resistance response. And it involves several core pathways such as plant hormone signal transduction, phenylpropanoid biosynthesis, MAPK signaling pathway—plant, plant- pathogen interaction, carbon metabolism, biosynthesis of amino acids, and several key Transcription factors (TFs) such as AP2/ERF, NAC, WRKY and bZIP. Metabolomic analysis revealed that KEGG pathway enrichment analysis showed unique metabolic pathways were enriched in P. talassica  ×  P. euphratica under both 200 mM and 400 mM NaCl treatments. Additionally, while there were some differences in the metabolic pathways enriched in the roots and stems, both tissues commonly enriched pathways related to the biosynthesis of secondary metabolites, biosynthesis of cofactors, biosynthesis of amino acids, flavonoid biosynthesis, and ABC transporters. Association analysis further indicated that biosynthesis of amino acids and plant hormone signal transduction pathway play key roles in the response of P. talassica  ×  P. euphratica to salt stress. The interactions between the differentially expressed genes (DEGs) and several differentially accumulated metabolites (DAMs), especially the strong association between LOC105124002 and Jasmonoyl-L-Isoleucine (pme2074), were again revealed by the interactions analysis. Conclusions In this study, we resolved the changes of metabolic pathways in roots and stems of P. talassica  ×  P. euphratica under different NaCl treatments and explored the associations between characteristic DEGs and DAMs, which provided insights into the mechanisms of P. talassica  ×  P. euphratica in response to salt stress.

Although a substantial proportion of plant biomass originates from the activity of vascular cambium, the molecular basis of radial plant growth is still largely unknown. To address whether cytokinins are required for cambial activity, we studied cytokinin signaling across the cambial zones of 2 tree species, poplar (Populus trichocarpa) and birch (Betula pendula). We observed an expression peak for genes encoding cytokinin receptors in the dividing cambial cells. We reduced cytokinin levels endogenously by engineering transgenic poplar trees (P. tremula x tremuloides) to express a cytokinin catabolic gene, Arabidopsis CYTOKININ OXIDASE 2, under the promoter of a birch CYTOKININ RECEPTOR 1 gene. Transgenic trees showed reduced concentration of a biologically active cytokinin, correlating with impaired cytokinin responsiveness. In these trees, both apical and radial growth was compromised. However, radial growth was more affected, as illustrated by a thinner stem diameter than in WT at same height. To dissect radial from apical growth inhibition, we performed a reciprocal grafting experiment. WT scion outgrew the diameter of transgenic stock, implicating cytokinin activity as a direct determinant of radial growth. The reduced radial growth correlated with a reduced number of cambial cell layers. Moreover, expression of a cytokinin primary response gene was dramatically reduced in the thin-stemmed transgenic trees. Thus, a reduced level of cytokinin signaling is the primary basis for the impaired cambial growth observed. Together, our results show that cytokinins are major hormonal regulators required for cambial development.

Wood development is strictly regulated by various phytohormones and auxin plays a central regulatory role in this process. However, how the auxin signaling is transducted in developing secondary xylem during wood formation in tree species remains unclear. Here, we identified an Aux/INDOLE-3-ACETIC ACID 9 (IAA9)-AUXIN RESPONSE FACTOR 5 (ARF5) module in Populus tomentosa as a key mediator of auxin signaling to control early developing xylem development. PtoIAA9, a canonical Aux/IAA gene, is predominantly expressed in vascular cambium and developing secondary xylem and induced by exogenous auxin. Overexpression of PtoIAA9m encoding a stabilized IAA9 protein significantly represses secondary xylem development in transgenic poplar. We further showed that PtoIAA9 interacts with PtoARF5 homologs via the C-terminal III/IV domains. The truncated PtoARF5.1 protein without the III/IV domains rescued defective phenotypes caused by PtoIAA9m. Expression analysis showed that the PtoIAA9-PtoARF5 module regulated the expression of genes associated with secondary vascular development in PtoIAA9m- and PtoARF5.1-overexpressing plants. Furthermore, PtoARF5.1 could bind to the promoters of two Class III homeodomain-leucine zipper (HD-ZIP III) genes, PtoHB7 and PtoHB8, to modulate secondary xylem formation. Taken together, our results suggest that the Aux/IAA9-ARF5 module is required for auxin signaling to regulate wood formation via orchestrating the expression of HD-ZIP III transcription factors in poplar.

Background The plant microbiome represents one of the key determinants of plant health and productivity by providing a plethora of functional capacities such as access to low-abundance nutrients, suppression of phytopathogens, and resistance to biotic and/or abiotic stressors. However, a robust understanding of the structural composition of the bacterial microbiome present in different plant microenvironments and especially the relationship between below-ground and above-ground communities has remained elusive. In this work, we addressed hypotheses regarding microbiome niche differentiation and structural stability of the bacterial communities within different ecological plant niches. Methods We sampled the rhizosphere soil, root, stem, and leaf endosphere of field-grown poplar trees ( Populus tremula × Populus alba ) and applied 16S rRNA amplicon pyrosequencing to unravel the bacterial communities associated with the different plant habitats. Results We found that the structural variability of rhizosphere microbiomes in field-grown poplar trees ( P. tremula × P. alba ) is much lower than that of the endosphere microbiomes. Furthermore, our data not only confirm microbiome niche differentiation reports at the rhizosphere soil–root interface but also clearly show additional fine-tuning and adaptation of the endosphere microbiome in the stem and leaf compartment. Each plant compartment represents an unique ecological niche for the bacterial communities . Finally, we identified the core bacterial microbiome associated with the different ecological niches of Populus. Conclusions Understanding the complex host–microbe interactions of Populus could provide the basis for the exploitation of the eukaryote–prokaryote associations in phytoremediation applications, sustainable crop production (bio-energy efficiency), and/or the production of secondary metabolites.

Forest trees display a perennial growth behavior characterized by a multiple-year delay in flowering and, in temperate regions, an annual cycling between growth and dormancy. We show here that the CO/FT regulatory module, which controls flowering time in response to variations in daylength in annual plants, controls flowering in aspen trees. Unexpectedly, however, it also controls the short-day-induced growth cessation and bud set occurring in the fall. This regulatory mechanism can explain the ecogenetic variation in a highly adaptive trait: the critical daylength for growth cessation displayed by aspen trees sampled across a latitudinal gradient spanning northern Europe.

Intercellular communication mediated by CLAVATA3/EMBRYO SURROUNDING REGION-RELATED (CLE) peptides and their receptors is crucial for plant development and environmental adaptation. In this study, 45 and 89 CLEs were identified in Populus tomentosa and Populus alba × Populus glandulosa, respectively, and, together with the 52 CLEs in Populus trichocarpa, the chromosome localization, gene and protein characteristics, collinearity and gene duplication events, cis-acting regulatory elements in promoters and evolutionary relationships of CLEs in these three poplar species were analyzed. The CLEs of three poplar species were divided into four subfamilies. Among them, the CLEs in subfamilies I, II and IV were A-type CLEs, while those in subfamily III were B-type CLEs. During the evolutionary process of poplar, the selection pressure faced by whole-genome duplication or segmental duplication was purifying selection, and the duplication events led to the expansion of the CLE family in poplar. The exogenous addition of a certain concentration of poplar CLE13 peptides inhibits the root growth of Arabidopsis thaliana and poplar and simultaneously reduces the expression levels of ARFs and LBDs in the roots. In addition, drought stress induces the expression of PtrCLE13A. The overexpression of preCLE13A significantly enhances the osmotic and drought tolerance in Populus tomentosa. These results have provided valuable information for further research on the molecular mechanisms of CLE peptide signaling pathways in the woody model plant poplar regarding plant growth and stress resistance.

Abscisic acid (ABA), a widely known phytohormone involved in the plant response to abiotic stress, plays a vital role in mitigating Cd super(2+) toxicity in herbaceous species. However, the role of ABA in ameliorating Cd super(2+) toxicity in woody species is largely unknown. In the present study, we investigated ABA restriction on Cd super(2+) uptake and the relevance to Cd super(2+) stress alleviation in Cd super(2+)-hypersensitive Populus euphratica. ABA (5 mu M) markedly improved cell viability and growth but reduced membrane permeability in CdCl sub(2) (100 mu M)-stressed P. euphratica cells. Moreover, ABA significantly increased the activity of the antioxidant enzymes catalase (CAT), superoxide dismutase (SOD), and ascorbate peroxidase (APX), contributing to the scavenging of Cd super(2+)-elicited H sub(2)O sub(2) within P. euphratica cells during the period of CdCl sub(2) exposure (100 mu M, 24-72 h). ABA alleviation of Cd super(2+) toxicity was mainly the result of ABA restriction of Cd super(2+) uptake under Cd super(2+) stress. Steady-state and transient flux recordings showed that ABA inhibited Cd super(2+) entry into Cd super(2+)-shocked (100 mu M, 30 min) and short-term-stressed P. euphratica cells (100 mu M, 24-72 h). Non-invasive micro-test technique data showed that H sub(2)O sub(2) (3 mM) stimulated the Cd super(2+)-elicited Cd super(2+) influx but that the plasma membrane (PM) Ca super(2+) channel inhibitor LaCl sub(3) blocked it, suggesting that the Cd super(2+) influx was through PM Ca super(2+)-permeable channels. These results suggested that ABA up-regulated antioxidant enzyme activity in Cd super(2+)-stressed P. euphratica and that these enzymes scavenged the Cd super(2+)-elicited H sub(2)O sub(2) within cells. The entry of Cd super(2+) through the H sub(2)O sub(2)-mediated Ca super(2+)-permeable channels was subsequently restricted; thus, Cd super(2+) buildup and toxicity were reduced in the Cd super(2+)-hypersensitive species, P. euphratica.

The early phase of the interaction between tree roots and ectomycorrhizal fungi, prior to symbiosis establishment, is accompanied by a stimulation of lateral root (LR) development. We aimed to identify gene networks that regulate LR development during the early signal exchanges between poplar (Populus tremula 3 Populus alba) and the ectomycorrhizal fungus Laccaria bicolor with a focus on auxin transport and signaling pathways. Our data demonstrated that increased LR development in poplar and Arabidopsis (Arabidopsis thaliana) interacting with L. bicolor is not dependent on the ability of the plant to form ectomycorrhizae. LR stimulation paralleled an increase in auxin accumulation at root apices. Blocking plant polar auxin transport with 1-naphthylphthalamic acid inhibited LR development and auxin accumulation. An oligoarray-based transcript profile of poplar roots exposed to molecules released by L. bicolor revealed the differential expression of 2,945 genes, including several components of polar auxin transport (PtaPIN and PtaAUX genes), auxin conjugation (PtaGH3 genes), and auxin signaling (PtaIAA genes). Transcripts of PtaPIN9, the homolog of Arabidopsis AtPIN2, and several PtaIAAs accumulated specifically during the early interaction phase. Expression of these rapidly induced genes was repressed by 1-naphthylphthalamic acid. Accordingly, LR stimulation upon contact with L. bicolor in Arabidopsis transgenic plants defective in homologs of these genes was decreased or absent. Furthermore, in Arabidopsis pin2, the root apical auxin increase during contact with the fungus was modified.We propose a model in which fungus-induced auxin accumulation at the root apex stimulates LR formation through a mechanism involving PtaPIN9-dependent auxin redistribution together with PtaIAA-based auxin signaling.

Background Growth-regulating factors (GRFs) are plant-specific transcription factors that control organ size. Nineteen GRF genes were identified in the Populus trichocarpa genome and one was reported to control leaf size mainly by regulating cell expansion. In this study, we further characterize the roles of the other poplar GRFs in leaf size control in a similar manner. Results The 19 poplar GRF genes were clustered into six groups according to their phylogenetic relationship with Arabidopsis GRFs. Bioinformatic analysis, degradome, and transient transcription assays showed that 18 poplar GRFs were regulated by miR396, with GRF12b the only exception. The functions of PagGRF6b ( Pag , Populus alba × P. glandulosa ), PagGRF7a , PagGRF12a , and PagGRF12b , representing three different groups, were investigated. The results show that PagGRF6b may have no function on leaf size control, while PagGRF7a functions as a negative regulator of leaf size by regulating cell expansion. By contrast, PagGRF12a and PagGRF12b may function as positive regulators of leaf size control by regulating both cell proliferation and expansion, primarily cell proliferation. Conclusions The diversity of poplar GRFs in leaf size control may facilitate the specific, coordinated regulation of poplar leaf development through fine adjustment of cell proliferation and expansion.

Recent findings were able to show significant variability of stomatal dynamics between species, but not much is known about factors influencing stomatal dynamics and its consequences on biomass production, transpiration and water-use efficiency (WUE). We assessed the dynamics of stomatal conductance (g(s)) to a change of irradiance or vapour-pressure deficit (VPD) in two Populus euramericana and two Populus nigra genotypes grown under control and drought conditions. Our objectives were to determine the diversity of stomatal dynamics among poplar genotypes, and if soil water deficit can alter it. Physiological and morphological factors were investigated to find their potential links with stomatal morphology, WUE and its components at the whole-plant level. We found significant genotypic variability of g(s) dynamics to both irradiance and VPD. Genotypes with faster stomatal dynamics were correlated with higher stomatal density and smaller stomata, and the implications of these correlations are discussed. Drought slowed g(s) dynamics, depending on genotype and especially during stomatal closing. This finding is contrary to previous research on more drought-tolerant species. Independently of the treatment, faster stomatal dynamics were negatively correlated with daily whole-plant transpiration, presenting new evidence of a previously hypothesized contribution of stomatal dynamics to whole-plant water use.