• Energy Research

Dioecious plants, which comprise more than 14,620 species, account for an important component of terrestrial ecosystems. Hence, understanding the sexually dimorphic responses in balancing carbon (C) supply and demand under elevated CO(2) is important for understanding leaf sink-to-source transitions. Here we investigate sex-related responses of the dioecious Populus cathayana Rehd. to elevated CO(2) and elevated temperature. The plants were grown in environmentally controlled growth chambers at two CO(2) enrichment regimes (350 ± 20 and 700 ± 20 μmol mol(-1)) with two temperature levels, elevated by 0 and 2 ± 0.2 °C (compared with the out-of-chamber environment). Plant growth characteristics, carbohydrate accumulation, C and nitrogen (N) allocation, photosynthetic capacity, N use efficiency and the morphology of mesophyll cells were investigated in the developing leaves (DLs) and expanded leaves (ELs) of both males and females. Elevated CO(2) enhanced plant growth and photosynthetic capacity in DLs of both males and females, and induced the male ELs to have a greater leaf mass production, net photosynthesis rate (P(n)), chlorophyll a/b ratio (Chl a/b), soluble protein level (SP), photosynthetic N use efficiency and soluble sugar level compared with females at the same leaf stage. Elevated temperature enhanced source activities and N uptake status during CO(2) enrichment, and the combined treatment induced males to be more responsive than females in sink capacities, especially in ELs, probably due to greater N acquisition from other plant parts. Our findings showed that elevated CO(2) increases the sink capacities of P. cathayana seedlings, and elevated temperature enhances the stimulation effect of elevated CO(2) on plant growth. Male ELs were found to play an important role in N acquisition from roots and stems under decreasing N in total leaves under elevated CO(2). Knowledge of the sex-specific leaf adaptability to warming climate can help us to understand sex-related source-to-sink transitions in dioecious plant species.

Abstract Dioecious trees have evolved sex-specific adaptation strategies to cope with inorganic phosphorus (Pi) limitation. Yet, little is known about the effects of Pi limitation on plant metabolism, particularly in dioecious woody plants. To identify potential gender-specific metabolites appearing in response to Pi limitation in poplars, we studied the metabolic and ionomic responses in the roots and leaves of Populus cathayana Rehd males and females exposed to a 60-day period of Pi deficiency. Besides significant decreases in phosphorus contents in both Pi-deficient roots and leaves, the calcium level decreased significantly and the sulfur content increased significantly in Pi-deficient male roots, while the zinc and ferrum contents increased significantly in Pi-deficient female roots. Inorganic P deficiency caused a smaller change in the abscisic acid content, but a significant increase in the jasmonic acid content was detected in both leaves and roots. Salicylic acid significantly decreased under Pi deficiency in male leaves and female roots. Changes were found in phospholipids and phosphorylated metabolites (e.g., fructose-6-phosphate, glycerol-3-phosphate, glucose-6-phosphate, phosphoric acid and inositol-1-phosphate) in roots and leaves. Both P. cathayana males and females relied on inorganic pyrophosphate-dependent but not on Pi-dependent glycolysis under Pi-deficient conditions. Sex-specific metabolites in leaves were primarily in the category of primary metabolites (e.g., amino acids), while in roots primarily in the category of secondary metabolites (e.g., organic acids) and sugars. The metabolome analysis revealed that sexually different pathways occurred mainly in amino acid metabolism, and the tissue-related differences were in the shikimate pathway and glycolysis. We observed changes in carbon flow, reduced root biomass and increased amino acid contents in P. cathayana males but not in females, which indicated that males have adopted an energy-saving strategy to adapt to Pi deficiency. Thus, this study provides new insights into sex-specific metabolic responses to Pi deficiency.

ABSTRACTDrought stress responses and sensitivity of dioecious plants, such as Populus cathayana Rehd., are determined by different mechanisms in each sex. In general, males tend to be more resistant while females are more sensitive. Here, we used reciprocal grafting between males and females to determine the relative importance of roots and shoots when plants are exposed to drought stress. Total dry matter accumulation (DMA), photosynthetic capacity, long‐term water‐use efficiency (Δ), water potential and ultrastructure of mesophyll cells were evaluated to determine the different roles of root and shoot in sex‐related drought responses. Plants with male roots were found to be more resistant and less sensitive to water stress than those with female roots under drought conditions. On the contrary, plants with female shoots grew better than those with male shoots under well‐watered conditions. These results indicated that the sensitivity of males and females to water stress is primarily influenced by root processes, while under well‐watered conditions sexual differences in growth are primarily driven by shoot processes. Furthermore, grafting female shoot scion onto male rootstock was proved to be an effective mean to improve resistance to water stress in P. cathayana females.

The degree to which branches are autonomous in their acclimation responses to alteration in light environment is still poorly understood. We investigated the effects of shading of the sapling crown of Cunninghamia lanceolata (Lamb.) Hook on the whole-tree and mid-crown branch growth and current-year foliage structure and physiology. Four treatments providing 0, 50, 75 and 90% shading compared with full daylight (denoted as Treatment(0), Treatment(50%), Treatment(75%) and Treatment(90%), and Shaded(0), Shaded(50%), Shaded(75%) and Shaded(90%) for the shaded branches and Sunlit(0), Sunlit(50%), Sunlit(75%) and Sunlit(90%) for the opposite sunlit branches under natural light conditions, respectively), were applied over two consecutive growing seasons. Shading treatments decreased the growth of basal stem diameter, leaf dry mass per unit leaf area, stomatal conductance, transpiration rate, the ratio of water-soluble to structural leaf nitrogen content, photosynthetic nitrogen-use efficiency and instantaneous and long-term (estimated from carbon isotope composition) water-use efficiency in shaded branches. Differences between shaded and sunlit branches increased with increasing severity and duration of shading. A non-autonomous, partly compensatory behavior of non-shaded branches was observed for most traits, thus reflecting the dependence between the traits of sunlit branches and the severity of shading of the opposite crown half. The results collectively indicated that tree growth and branch and leaf acclimation responses of C. lanceolata are not only affected by the local light environment, but also by relative within-crown light conditions. We argue that such a non-autonomous branch response to changes in light conditions can improve whole-tree resource optimization. These results contribute to better understanding of tree growth and utilization of water and nitrogen under heterogeneous light conditions within tree canopies.

The functional balance between leaves and roots is believed to be mediated by the specific location of shoots and roots, i.e. differences in transport distances and degrees of organ connectivity. However, it remains unknown whether the adaptive responses of trees to biomass removal depend on the relative orientation of leaf and root pruning. Here, we applied five pruning treatments to saplings of Cunninghamia lanceolata (Chinese fir) under field and glasshouse conditions, namely no pruning (control), half of lateral branches pruned, half of lateral roots pruned, half of the branches and roots pruned on the same side of the plant, and half of the branches and roots pruned on opposite sides of the plant. The effects of pruning on the growth, carbon storage and allocation, and physiology of leaves and fine roots on the same and opposite sides of the plant were investigated. Compared with the effect of root-pruning on leaves, fine roots were more limited by carbon availability and their physiological activity was more strongly reduced by shoot pruning, especially when branches on the same side of the plant were removed. Pruning of branches and roots on the opposite side of the plant resulted in the lowest carbon assimilation rates and growth among all treatments. The results of a stable-isotope labeling indicated that less C was distributed to fine roots from the leaves on the opposite side of the plant compared to those on the same side, but N allocation from roots to leaves depended less on the relative root and leaf orientation. The results collectively indicate that the functional responses of C. lanceolata to pruning are not only determined by the source-sink balance model but are also related to interactions between leaves and fine roots. We argue that the connectivity among lateral branches and roots depends on their relative orientation, which is therefore critical for the functional balance between leaves and fine roots.

Lead (Pb) and drought frequently coexist in China's forests and seriously affect their biomass, net primary productivity and biodiversity, particularly among dioecious trees with different sex-related reproduction costs. Here, the effects of Pb, deposited into soil (Pbsoil) and leaves (Pb(leaf)), on the traits related directly or indirectly to photosynthetic activity were evaluated in the dioecious tree, Populus cathayana Rehd. In addition, we analysed the potential of P. cathayana males and females for phytoremediation based on the photosynthetic activity, cellular ultrastructure and phytoremediation-related parameters. The Pb level was 2.7 mmol Pb(NO3)2 kg(-1) dry soil in the Pb(soil) treatment and 1.8 mmol Pb(NO3)2 per plant in the Pb(leaf) treatment. In parallel experiments, two watering regimes, 100 and 50% of the field capacity, were applied. The stress duration was 2 months. Dry mass production, net photosynthetic rate increased in both sexes, particularly in females, when exposed to Pb(leaf) alone, Pb(soil) with drought and Pb(leaf) with drought. The study revealed that males exhibit greater plasticity in the photosynthetic capacity than females. Severe damage to cellular ultrastructure was also observed in the leaves of males and females exposed to Pb(leaf), but more strongly in females. However, the Pb(soil) treatment alone did not affect these traits as significantly compared with other treatments. Moreover, drought significantly increased the sensitivity to Pb stress in both sexes, but more so in females. In addition, changes in the photosynthetic capacity and cellular ultrastructure combined with the analysis of tolerance index, translocation factor, bioconcentration factor and Pb concentration showed that males and females could be employed for different purposes in phytoremediation: females are promising candidates for phytoextraction, whereas males are promising candidates for phytostabilization.

Abstract It remains unclear how global climate change affects dioecious plants that may be especially vulnerable to climate drivers, because they often exhibit skewed sex ratios and eco-physiological specialization in certain microhabitats. In this study, female and male saplings of Populus cathayana were employed to explore sex-specific responses and the effects of sexual competition under elevated temperature (ET), elevated CO2 (EC) and combination of elevated temperature and CO2 (ETC). The results demonstrated that elevated temperature and CO2 interactively modulated sexual competition and responses of P. cathayana. Moreover, competition patterns affected the eco-physiological responses of P. cathayana to climate change treatments. Under both intra- and inter-sexual competition, biomass components, photosynthetic parameters and carbon-related metabolites of females were most strongly affected by ET, while males exhibited a higher photosynthesis and resource use efficiency, and a better biomass accumulation and carbon balance mechanism when compared to females when experiencing intra-sexual competition under EC. The competitive pressure of females on males in inter-sexual competition was intensified by ET, while it was alleviated by ETC. We conclude that climate change drivers and competition patterns differently regulate the sex-specific responses and competitive intensity of males and females, which may have a crucial effect on sex ratios, spatial sexual segregation, biomass production and carbon sequestration in dioecious species in the future.

Abstract In this study, intergeneric grafting was employed between Populus cathayana and Salix rehderiana to investigate the grafting compatibility of the two Salicaceae plants and to reveal whether grafting can improve their drought resistance. Under different grafting combinations (P. cathayana scion with P. cathayana rootstock, P/P; P. cathayana scion with S. rehderiana rootstock, P/S; S. rehderiana scion with S. rehderiana rootstock, S/S; and S. rehderiana scion with P. cathayana rootstock, S/P), the survival and growth rate, biomass accumulation and allocation, photosynthetic traits, carbon isotope composition (δ13C), relative water content (RWC) and non-structural carbohydrates (NSCs) were measured. The results showed that the grafting compatibility between P. cathayana and S. rehderiana was very high, as the survival rates ranged from 76% to 100% under different grafting combinations. Drought significantly decreased growth, biomass accumulation, photosynthetic pigment contents, net photosynthesis rates (Pn) and RWC, and increased δ13C in all grafting combinations. Under drought stress, biomass accumulation, total chlorophyll, transpiration rate (E) and Pn were higher in P/P and P/S than in S/S and S/P. Compared with P/P, the growth rate, biomass accumulation, root/aboveground ratio (R/A ratio), carotenoid, RWC, starch and total soluble sugar (TSS) of P/S were less affected by drought. The height growth rate (GRH), R/A ratio, carotenoid, chlorophyll a, total chlorophyll, WUEi and TSS of S/P were lower than those of S/S under water-limited conditions. Moreover, a principal component analysis indicated that P/S and S/S had higher drought resistance than P/P and S/P under water deficits. The used method allows combining specific advantageous traits from P. cathayana and S. rehderiana, which may be a highly useful tool to enhance drought resistance in the cultivation of Salicaceae plants.