The transcriptional events and their relationship to physiological changes during poplar seed germination and post-germination.

BACKGROUND: Seed germination, the foundation of plant propagation, involves a series of changes at the molecular level. Poplar is a model woody plant, but the molecular events occurring during seed germination in this species are unclear. RESULTS: In this study, we investigated changes in gene transcriptional levels during different germination periods in poplar by high-throughput sequencing technology. Analysis of genes expressed at specific germination stages indicated that these genes are distributed in many metabolic pathways. Enrichment analysis of significantly differentially expressed genes based on hypergeometric testing revealed that multiple pathways, such as pathways related to glycolysis, lipid, amino acid, protein and ATP synthesis metabolism, changed significantly at the transcriptional level during seed germination. A comparison of ΣZ values uncovered a series of transcriptional changes in biological processes related to primary metabolism during poplar seed germination. Among these changes, genes related to CHO metabolism were the first to be activated, with subsequent expression of genes involved in lipid metabolism and then those associated with protein metabolism. The pattern of metabolomic and physiological index changes further verified the sequence of some biological events. CONCLUSIONS: Our study revealed molecular events occurring at the transcriptional level during seed germination and determined their order. These events were further verified by patterns of changes of metabolites and physiological indexes. Our findings lay a foundation for the elucidation of the molecular mechanisms responsible for poplar seed germination.

New Intrinsic Ecological Mechanisms of Leaf Nutrient Resorption in Temperate Deciduous Trees.

Leaf nutrient resorption is a critical process in plant nutrient conservation during leaf senescence. However, the ecological mechanisms underlying the large variability in nitrogen (NRE) and phosphorous (PRE) resorption efficiencies among trees remain poorly understood. We conducted a comprehensive study on NRE and PRE variability using 61 tree individuals of 10 temperate broad-leaved tree species. Three potentially interrelated intrinsic ecological mechanisms (i.e., leaf senescence phenology, leaf pigments, and energy residual) were verified. We found that a delayed leaf senescence date, increased degradation of chlorophylls and carotenoids, biosynthesis of anthocyanins, and reduced nonstructural carbohydrates were all positively correlated with NRE and PRE at the individual tree level. The intrinsic factors affecting resorption efficiency were ranked in decreasing order of importance: leaf pigments > energy residual > senescence phenology. These factors explained more variability in NRE than in PRE. Our findings highlight the significance of these three ecological mechanisms in leaf nutrient resorption and have important implications for understanding how nutrient resorption responds to climate change.

Trade-offs between leaf stoichiometric characteristics and photosynthetic traits of Larix gmelinii and its differences among provenances.

Variations and trade-offs between leaf stoichiometric characteristics and photosynthetic traits are indicative of ecological adaptation strategies of plants and their responses to environment changes. In a common garden of Maoershan, we measured leaf stoichiometric characteristics (carbon content (C), nitrogen content (N), phosphorus content (P), C/N, C/P, N/P) and photosynthetic traits (maximum net photosynthetic rate (Amax), maximum electron transport rate (Jmax), maximum carboxylation rate (Vmax)) of Larix gmelinii from 17 geographical provenances. We examined the provenance differences in stoichiometric characteristics and photosynthetic traits, and analyzed their trade-offs and influencing factors. The results showed leaf stoichiometric characteristics and photosynthetic traits significantly differed among provenances. The climatic factors of seed-source sites explained 54.8% and 67.2% of the variation in stoichiometric characteristics and photosynthetic traits, respectively. Aridity index (AI) of seed-source sites was positively correlated with C, N, P, Amax, Jmax, Vmax, but negatively with C/N, C/P, and N/P. Results of redundancy analysis showed that stoichiometric characteristics accounted for 75.0% of the variation in photosynthetic traits. Amax, Jmax, Vmax were positively correlated with C, N, P, and negatively correlated with C/N, C/P, N/P. The provenance differences in stoichiometric characteristics, photosynthetic traits, and their synergistic relationship suggested the long-term adaptation of trees to the climate of seed-source sites. These findings were of great significance for understanding ecological adaptation strategies of trees in response to climate change.

Provenance variation of root C, N, P, and K stoichiometric characteristics under different diameter classes of Larix gmelinii.

For exploring the difference of root stoichiometric characteristics among diameter classes and provenances, we examined the contents and stoichiometric ratios of carbon (C), nitrogen (N), phosphorus (P), and potassium (K) in three diameter classes of roots (0-1, 1-2 and 2-5 mm, respectively) of 39-year-old Larix gmelinii grown in a common garden. The results showed that root element contents and their stoichiometric ratios had significant difference among three diameter classes of roots. C content, C:N, C:P, C:K were the lowest, and N, P, K contents, N:P, and N:K were the highest in 0-1 mm diameter class roots. Compared with the 1-2 and 2-5 mm diameter class roots, 0-1 mm diameter class roots had different seasonal dynamics, which might be caused by the fact that 0-1 mm diameter class roots are absorptive roots and the other diameter class roots are transport roots. There was no provenance difference in C content among all diameter class roots, while significant provenance differences were found in N, K contents, C:N, and C:K in 0-1 mm diameter class roots, and great provenance differences for in P content, C:P, N:P, and N:K in 0-1 and 1-2 mm diameter class roots. N content, K content, C:P, N:P, and N:K in 0-1 mm diameter class roots had positive correlation with the aridity index of seed-source sites, while the P content, C:N and C:K had negative correlations. The stoichiometric characteristics were related with the diameter (or function) of roots, and had significant provenance differences in 0-1 mm (absorptive root) and 1-2 mm diameter class roots, which might be attributed to their genotypic adaptation to the environment of seed-source sites.

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To reveal molecular mechanisms underlying photosynthesis responses of Dahurian larch (Larix gmelinii) to environmental changes, we used the high-throughput sequencing technology to sequence the transcriptome of larch leaves from four latitudinal sites with different environmental conditions, and compared differential expression genes (DEGs). The four sites from high- to low-latitude were Tahe (52°52' N), Songling (50°72' N), Heihe (49°22' N), and Dailing (47°08' N). A total of 282428811 clean reads were sequenced out, among which the abundace of DEGs were 16915, 18812, 28536, 20635, 29957 and 23617 for the Tahe-Songling, Tahe-Heihe, Tahe-Dailing, Songling-Heihe, Songling-Dailing, and Heihe-Dailing comparisons, respectively. The expression of nine Psb genes family (i.e., PsbB, PsbK, PsbO, PsbP, PsbQ, PsbS, PsbW, Psb27, and Psb28) encoding Photosystem Ⅱ and that of three genes (ATPF1A,atpA, ATPF1G, atpG, and ATPF1D, atpH) encoding F-type ATPase, which were involved in photosynthesis pathway, were significantly up-regulated with increasing environmental differences among the sites. A similar up-regulation pattern occurred for the expression of genes encoding glutamine synthetase (glnA, GLUL), nitrate reductase (NR), and carbonic anhydrase (cynT, can) that were involved in nitrogen metabolism pathway. The numbers of DEGs and up-regulated genes increased with the increases in environmental changes among the sites, resulting in inter-site divergence of photosynthetic capacity of larch trees.

Transplanting larch trees into warmer areas increases the photosynthesis and its temperature sensitivity.

To investigate the effects of climate warming on photosynthesis, Dahurian larch (Larix gmelinii Rupr.) trees from four sites (spanning ~ 5.5° in latitude and ~4 °C of warming) within the geographic range in China were transplanted into a common garden close to the warmer border in 2004. Throughout the growing season of 2018, the CO2- and temperature-response curves of the photosynthesis in the common garden and at the original sites were measured. It was discovered that warming treatment considerably increased the maximum net photosynthetic rate (Amax) by 23.4-35.3% depending on the sites, signifying that warming upregulated Amax with respect to the degree of warming. At 25 °C, warming enhanced the maximum Rubisco carboxylation rate (Vcmax), maximum electron transport rate (Jmax), and mass-based leaf nitrogen concentration (Nmass). The climate warming effect (CWE) on Amax was positively associated with the CWEs on Vcmax, Jmax and Nmass, which indicated that warming promoted Amax primarily via increasing carboxylation and photosynthetic electron transport rates and leaf nitrogen supply. The CWE in optimal photosynthetic temperature (Topt) was significant for the trees from the northern sites rather than the southern sites; however, the effect vanished for the trees transplanted to the common garden; this implied that Topt exhibited limited local thermal acclimation. Nevertheless, warming narrowed the temperature-response curve, the effect of which was positively associated with the warming magnitude. These findings implied that trees transplanted into warmer areas changed the photosynthetic optimum temperature and sensitivity. In summary, our results deepen the understanding of the underlying mechanisms of intraspecific responses of photosynthesis to temperature changes, including which of the modeling would improve the prediction of tree growth and forest carbon cycling under climate warming.

Thermal acclimation of leaf dark respiration of Larix gmelinii: A latitudinal transplant experiment.

The response of tree leaf dark respiration (Rd) to temperature change is important in modeling and predicting forest carbon (C) cycling under climate change, but it has rarely been investigated in nature. We conducted a field experiment by transplanting the trees of Larix gmelinii - the dominant tree species in Chinese boreal forests from four latitudinal sites to a common garden near the warm border of its range. Our objective was to explore thermal acclimation of Rd and the underlying mechanisms by comparing the temperature-response curves of Rd and related leaf traits both in the common garden and at the original sites. We found that warming significantly decreased Rd and its temperature sensitivity (Q10), which changed across the growing season and were correlated with the mean annual temperature of the original sites, reflecting a combination of both short- and long-term respiratory acclimation to warming. The trees from the southern sites tended to have higher thermal acclimation of Rd and lower Q10 than that from the northern sites. Rd and Q10 were highly correlated with the concentrations of leaf nitrogen and soluble sugars, which may be used as proxies for assessing thermal acclimation of respiration. Considering both short- and long-term thermal acclimation of Rd likely improves the prediction of forest C cycling in response to climate change.

[Effects of nitrogen fertilization on fine root lifespan of Fraxinus mandshurica and Larix gmelinii].

Root observation tubes (minirhizotrons) were used to study the effects of nitrogen addition on the fine root growth of Fraxinus mandshurica and Larix gmelinii, with the correlations between the fine root lifespan and nitrogen availability analyzed. After the nitrogen addition, the fine root number of F. mandshurica and L. gmelinii had a decreasing trend, but the fine root diameter became larger and the branching ratio decreased. The survival rate of F. mandshurica fine roots increased, and the median root lifespan prolonged 105 days, compared with the control. No significant responses to the nitrogen addition were observed in the survival rate of L. gmelinii fine roots. The first-order fine roots with smaller diameter, the roots in surface soil (0-15 cm), and the fine roots newly born in spring and summer were vulnerable to extend their lifespan by nitrogen addition, suggesting that the fine roots with higher physiological activity were easily to be affected by nitrogen fertilization.

[Carbon and nitrogen storages and allocation in tree layers of Fraxinus mandshurica and Larix gmelinii plantations].

By the methods of wood analysis and sequential soil core, the biomass and productivity of the tree layers in 20-year old Fraxiuns mandshurica and Larix gmelinii plantations, as well as the carbon (C) and nitrogen (N) storages in the above- and below-ground organs of the stands, were estimated. The biomass of F. mandshurica and L. gmelinii was 6815.10 g x m(-2) and 9295.95 g x m(-2), in which, stem occupied 57.32% and 58.01%, and fine roots occupied 2.67% and 1.80%, respectively. The annual productivity of F. mandshurica and L. gmelinii was 1618.16 and 2102.45 g x m(-2) x a(-1), in which, stem accounted for 39.34% and 46.70%, and fine roots accounted for 12.06% and 5.25%, respectively. The C content in the organs of F. mandshurica was lower than that of L. gmelinii, while the N content was in adverse. The C storage of F. mandshurica was lower than that of L. gmelinii, while the N storage had no significant difference between the two tree species. The biomass, productivity, and C and N storages of aboveground organs were lower for F. mandshurica than for L. gmelinii, indicating the higher construction efficiency of the aboveground part of L. gmelinii. Due to the significant differences in the C and N contents between tree species and between the organs of same tree species, the measurement should be made on different tree species and different organs to have an accurate estimation of forest C and N storages.

[Effects of nitrogen fertilization on ectomycorrhizal infection of first order roots and root morphology of Larix gmelinii plantation].

In this paper, the first order roots of Larix gmelinii plantation under N fertilization were sampled from different soil depths in different seasons to study their morphology under effects of ectomycorrhizal fungi. The results showed that the infection rate of ectomycorrhizal fungi on the first order roots was significantly affected by soil N availability, soil depth, and season. N fertilization induced a decrease of the infection rate, and the decrement varied with soil depth and season. In comparing with the control, the infected first order roots had an obvious variation of their morphology, e. g., averagely, root diameter increased by 18.7%, root length decreased by 23.7%, and specific root length decreased by 16.3%, which differed significantly with N application rate, soil depth, and season. The infection of ectomycorrhizal fungi changed the first order root morphology of L. gmelinii, which might substantially affect the physiological and ecological processes of host plant fine roots.