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Over-application of copper (Cu) is prevalent in citrus orchards of South China. Therefore, the evaluation of citrus Cu tolerance is of significance for better nutrient management and in the breeding of Cu-tolerant citrus species. In this study, seedlings of ‘Xuegan’ [Citrus sinensis (L.) Osbeck] and ‘Shatian pummelo’ [Citrus grandis (L.) Osbeck] were treated with 0.5 µM Cu (Control) or 300 µM Cu (Cu toxicity) for 18 weeks in sandy culture medium. The results demonstrated that excess Cu decreased seedling biomass, plant height, and root length and significantly increased Cu accumulation of C. sinensis and C. grandis. The Cu distribution in citrus tissues was following the order: lateral roots > primary roots > stems > leaves. Compared to C. sinensis, C. grandis had a 22.4% higher Cu concentration in the lateral roots under Cu toxicity. According to the stomata limiting theory, the inhibition of the net photosynthetic rate of C. sinensis leaves by Cu toxicity was mainly attributed to non-stomata limitation. The chlorophyll a transient indicated much more severe damage to the structure and function of leaf photosynthetic system II (PS II) in C. sinensis than C. grandis by Cu toxicity. Conclusively, a higher Cu retention in the lateral roots, the maintenance of a stable PS II structure, and higher electron transferring rate from PS II to PS I implied the relatively higher Cu tolerance of C. grandis seedlings. These results provide essential information for further investigation on the Cu-tolerant mechanisms of citrus species and the future breeding of Cu-tolerant citrus species.

Background In this study, we aimed to test the hypothesis that magnesium (Mg) remobilization in citrus plants is regulated by Mg supply and contributes to differences in the growth of the parent and branch organs. Citrus seedlings were grown in sand under Mg deficient (0 mmol Mg2+ L−1, -Mg) and Mg sufficient (2 mmol Mg2+ L−1, + Mg) conditions. The effects on biomass, Mg uptake and transport, gas exchange and chlorophyll fluorescence, as well as related morphological and physiological parameters were evaluated in different organs. Results Mg deficiency significantly decreased plant biomass, with a decrease in total plant biomass of 39.6%, and a greater than twofold decrease in the branch organs compared with that of the parent organs. Reduced photosynthesis capacity was caused by a decreased in pigment levels and photosynthetic electron transport chain disruption, thus affecting non-structural carbohydrate accumulation and plant growth. However, the adaptive responses of branch leaves to Mg deficiency were greater than those in parent leaves. Mg deficiency inhibited plant Mg uptake but enhanced Mg remobilization from parent to branch organs, thus changing related growth variables and physiological parameters, including protein synthesis and antioxidant enzyme activity. Moreover, in the principal components analysis, these variations were highly clustered in both the upper and lower parent leaves, but highly separated in branch leaves under the different Mg conditions. Conclusions Mg deficiency inhibits the growth of the parent and branch organs of citrus plants, with high Mg mobility contributing to differences in physiological metabolism. These findings suggest that Mg management should be optimized for sustainable citrus production.

Phosphorus (P) is an essential macronutrient for plant growth, development and production. However, little is known about the effects of P deficiency on nutrient absorption, photosynthetic apparatus performance and antioxidant metabolism in citrus. Seedlings of ‘sour pummelo’ (Citrus grandis) were irrigated with a nutrient solution containing 0.2 mM (Control) or 0 mM (P deficiency) KH2PO4 until saturated every other day for 16 weeks. P deficiency significantly decreased the dry weight (DW) of leaves and stems, and increased the root/shoot ratio in C. grandis but did not affect the DW of roots. The decreased DW of leaves and stems might be induced by the decreased chlorophyll (Chl) contents and CO2 assimilation in P deficient seedlings. P deficiency heterogeneously affected the nutrient contents of leaves, stems and roots. The analysis of Chl a fluorescence transients showed that P deficiency impaired electron transport from the donor side of photosystem II (PSII) to the end acceptor side of PSI, which showed a greater impact on the performance of the donor side of PSII than that of the acceptor side of PSII and photosystem I (PSI). P deficiency increased the contents of ascorbate (ASC), H2O2 and malondialdehyde (MDA) as well as the activities of superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR) and glutathione reductase (GR) in leaves. In contrast, P deficiency increased the ASC content, reduced the glutathione (GSH) content and the activities of SOD, CAT, APX and monodehydroascorbate reductase (MDHAR), but did not increase H2O2 production, anthocyanins and MDA content in roots. Taking these results together, we conclude that P deficiency affects nutrient absorption and lowers photosynthetic performance, leading to ROS production, which might be a crucial cause of the inhibited growth of C. grandis.

This present study examined excess copper (Cu) effects on seedling growth, leaf Cu concentration, gas exchange, and protein profiles identified by a two-dimensional electrophoresis (2-DE) based mass spectrometry (MS) approach after Citrus sinensis and Citrus grandis seedlings were treated for six months with 0.5 (control), 200, 300, or 400 &mu, M CuCl2. Forty-one and 37 differentially abundant protein (DAP) spots were identified in Cu-treated C. grandis and C. sinensis leaves, respectively, including some novel DAPs that were not reported in leaves and/or roots. Most of these DAPs were identified only in C. grandis or C. sinensis leaves. More DAPs increased in abundances than DAPs decreased in abundances were observed in Cu-treated C. grandis leaves, but the opposite was true in Cu-treated C. sinensis leaves. Over 50% of DAPs were associated with photosynthesis, carbohydrate, and energy metabolism. Cu-toxicity-induced reduction in leaf CO2 assimilation might be caused by decreased abundances of proteins related to photosynthetic electron transport chain (PETC) and CO2 assimilation. Cu-effects on PETC were more pronounced in C. sinensis leaves than in C. grandis leaves. DAPs related to antioxidation and detoxification, protein folding and assembly (viz., chaperones and folding catalysts), and signal transduction might be involved in Citrus Cu-toxicity and Cu-tolerance.

Seedlings of ‘Shatian pummelo’ (Citrus grandis) and ‘Xuegan’ (Citrus sinensis) were supplied daily with nutrient solution at a concentration of 0.5 (control), 100, 200, 300, 400, or 500 μM CuCl2 for 6 months. Thereafter, seedling growth; leaf, root, and stem levels of nutrients; leaf gas exchange; levels of pigments; chlorophyll a fluorescence (OJIP) transients and related parameters; leaf and root relative water content; levels of nonstructural carbohydrates; H2O2 production rate; and electrolyte leakage were comprehensively examined (a) to test the hypothesis that Cu directly damages root growth and function, thus impairing water and nutrient uptake and hence inhibiting shoot growth; (b) to establish whether the Cu-induced preferential accumulation of Cu in the roots is involved in Cu tolerance of Citrus; and (c) to elucidate the possible causes for the Cu-induced decrease in photosynthesis. Most of the growth and physiological parameters were greatly altered only at 300–500 μM (excess) Cu-treated seedlings. Cu supply increased the level of Cu in the roots, stems, and leaves, with a greater increase in the roots than that in the stems and leaves. Many of the fibrous roots became rotten and died under excess Cu. These findings support the hypothesis that Cu directly damages root growth and function, thus impairing water and nutrient uptake and hence inhibiting shoot growth, and the conclusion that the preferential accumulation of Cu in the roots under excess Cu is involved in the tolerance of Citrus to Cu toxicity. The lower CO2 assimilation in excess Cu-treated leaves was caused mainly by nonstomatal factors, including structural damage to thylakoids, feedback inhibition due to increased accumulation of nonstructural carbohydrates, decreased uptake of water and nutrients, increased production of reactive oxygen species, and impaired photosynthetic electron transport chain. Also, we discussed the possible causes for the excess Cu-induced decrease in leaf pigments and accumulation of nonstructural carbohydrates in the roots and leaves.

Background Magnesium (Mg)-deficiency is one of the most prevalent physiological disorders causing a reduction in Citrus yield and quality. ‘Xuegan’ (Citrus sinensis) seedlings were irrigated for 16 weeks with nutrient solution containing 2 mM (Mg-sufficiency) or 0 mM (Mg-deficiency) Mg(NO3)2. Thereafter, we investigated the Mg-deficient effects on gas exchange and chlorophyll a fluorescence in the upper and lower leaves, and Mg, reactive oxygen species (ROS) and methylglyoxal (MG) metabolisms in the roots, lower and upper leaves. The specific objectives were to corroborate the hypothesis that the responses of ROS and MG metabolisms to Mg-deficiency were greater in the lower leaves than those in the upper leaves, and different between the leaves and roots. Results Mg level was higher in the Mg-deficient upper leaves than that in the Mg-deficient lower leaves. This might be responsible for the Mg-deficiency-induced larger alterations of all the measured parameters in the lower leaves than those in the upper leaves, but they showed similar change patterns between the Mg-deficient lower and upper leaves. Accordingly, Mg-deficiency increased greatly their differences between the lower and upper leaves. Most of parameters involved in ROS and MG metabolisms had similar variation trends and degrees between the Mg-deficient lower leaves and roots, but several parameters (namely glutathione S-transferase, sulfite reductase, ascorbate and dehydroascorbate) displayed the opposite variation trends. Obviously, differences existed in the Mg-deficiency-induced alterations of ROS and MG metabolisms between the lower leaves and roots. Although the activities of most antioxidant and sulfur metabolism-related enzymes and glyoxalase I and the level of reduced glutathione in the Mg-deficient leaves and roots and the level of ascorbate in the leaves were kept in higher levels, the levels of malonaldehyde and MG and/or electrolyte leakage were increased in the Mg-deficient lower and upper leaves and roots, especially in the Mg-deficient lower leaves and roots. Conclusions The ROS and MG detoxification systems as a whole did not provide sufficient detoxification capacity to prevent the Mg-deficiency-induced production and accumulation of ROS and MG, thus leading to lipid peroxidation and the loss of plasma membrane integrity, especially in the lower leaves and roots. Electronic supplementary material The online version of this article (10.1186/s12870-019-1683-4) contains supplementary material, which is available to authorized users.

In south China, Magnesium (Mg) deficiency is commonly found in Citrus orchard due to soil acidification. However, less information is available discussing the global metabolic profile of Mg-deficient leaves in Citrus species. In the study, seedlings of ‘Xuegan’ [Citrus sinensis (L.) Osbeck] were treated with Mg-plus (2 mM, as Control) or Mg-deficiency (0 mM) for 16 weeks by sandy culture, respectively. The untargeted metabolomics based on UHPLC-Q-TOF/MS were investigated to reveal the metabolic responses to Mg-deficiency in leaves of C. sinensis. The results indicated severe growth inhibition and obvious leaf chlorosis in Mg-deficient seedlings of C. sinensis compared to control. The principal component analysis (PCA) and orthogonal partial least squares-discriminate analysis (OPLS-DA) on the LC/GC data demonstrated a clear discrimination between the profiles of metabolites in control leaves and Mg-deficient leaves of C.sinensis. Besides, the identification and classification of those differential expressed metabolites (DEMs) revealed Mg-deficiency upregulated relative abundances of 78.42 % lipids and lipid-like molecules. In contrast, Mg-deficiency downregulated 89.42 % phenylpropanoids and polyketides in leaves of C.sinensis. Remarkably, relative abundances of most monosaccharides and amino acids increased while relative abundances of most leaf pigment compounds and organic acids decreased in response to Mg-deficiency in C. sinensis leaves. Those results support both primary and secondary metabolic modifications in C. sinensis leaves exposed to Mg-deficiency. Overall, the findings of present study contribute to a better understanding on the metabolic changes of Mg-deficient Citrus species, which will provide references for nutrient management and quality control in Citrus production.

Nitrogen (N) is an essential mineral nutrient for plant growth. Appropriate N fertilization may enhance the efficiency of N use and improve crop yield. However, the effects of N form and level on citrus plants are not well understood. We hypothesized that citrus plants are ammonium (as NH4+-N or AN) partial or insensitive based on the characteristics of preferential growth in the acidic red soil of southern China, where AN is the dominant N form in the soil. Citrus (Citrus reticulata Blanco cv. Ponkan) seedlings were cultivated in sand in the presence of three N levels (1, 4 and 8 mmol L−1) supplied in two N forms (AN; nitrate, as NO3--N or NN). Biomass, root morphology, leaf characteristics, N uptake, and assimilation were determined. Compared with NN treatment, plant growth characteristics, including biomass, growth rate, root and leaf characteristics were all markedly inhibited under AN conditions, resulting in malondialdehyde (MDA) accumulation throughout the plants, except the specific leaf area. Under AN conditions, higher N concentration and free amino acid (FAA) content were observed, while N accumulation, soluble protein content, and the activities of nitrate reductase (NR), glutamine synthetase (GS) and glutamate synthase (GOGAT) in root and leaf were reduced, except for GOGAT in root. Moreover, the trend in these changes was amplified by increasing N supply, and significant interactions between N form and level were also found. Taken together, our results demonstrate that citrus plants are highly ammonium sensitive, the findings provide a theoretical basis for the optimized N management in the high-quality production of citrus.

Citrus are mainly grown in low pH soils with high active aluminum (Al). 'Xuegan' (Citrus sinensis (L.) Osbeck) and 'Shatian pummelo' (Citrus grandis (L.) Osbeck) seedlings were fertilized for 18 weeks with nutrient solution containing either 0 mM (control) or 1 mM (Al toxicity) AlCl3·6H2O. Aluminum induced decreases of biomass, leaf photosynthesis, relative water content and total soluble protein levels, and increases of methylglyoxal levels only occurred in C. grandis roots and leaves. Besides, the Al-induced decreases of pigments and alterations of chlorophyll a fluorescence transients and fluorescence parameters were greater in C. grandis leaves than those in C. sinensis leaves. Aluminum-treated C. grandis had higher stem and leaf Al levels and similar root Al levels relative to Al-treated C. sinensis, but lower Al distribution in roots and Al uptake per plant. Aluminum toxicity decreased nitrogen, phosphorus, potassium, calcium, magnesium and sulfur uptake per plant in C. grandis and C. sinensis seedlings, with the exception of Al-treated C. sinensis seedlings exhibiting increased sulfur uptake per plant and unaltered magnesium uptake per plant. Under Al-stress, macroelement uptake per plant was higher in C. sinensis than that in C. grandis. Aluminum toxicity decreased the ratios of reduced glutathione/(reduced + oxidized glutathione) and of ascorbate/(ascorbate + dehydroascorbate) only in C. grandis roots and leaves. The activities of most antioxidant enzymes, sulfur metabolism-related enzymes and glyoxalases and the levels of S-containing compounds were higher in Al-treated C. sinensis roots and leaves than those in Al-treated C. grandis ones. Thus, C. sinensis displayed higher Al tolerance than C. grandis did. The higher Al tolerance of C. sinensis might involve: (i) more Al accumulation in roots and less transport of Al from roots to shoots; (ii) efficient maintenance of nutrient homeostasis; and (iii) efficient maintenance of redox homeostasis via detoxification systems of reactive oxygen species and methylglyoxal.

In order to evaluate differential growth, photosynthesis and H + -ATPase activity responses to salt-induced stress, two Jerusalem artichoke ( Helianthus tuberosus L.) genotypes (Nanyu No. 1 and Qingyu No. 2) were used in sand-culture experiment with different concentrations of NaCl (0, 30, 60, 90, 120 and 150 mM). After 20 days of growth, the NaCl stress resulted in a decrease of biomass accumulation, relative leaf expansion rate and photosynthetic rate, but an increase of proline content in both genotypes. Compared with Qingyu No. 2, Nanyu No. 1 had lower biomass, photosynthetic rate, gas exchange and transpiration rate, but higher proline content, activities of plasma membrane H + -ATPase (PM H + -ATPase) and vacuolar membrane H + -ATPase (VM H + -ATPase). Hence, the NaCl adaptation strategy in Nanyu No. 1 was by lowering photosynthetic rate, stomatal conductance and transpiration rate while maintaining high H + -ATPase activities, whereas the adjustment of Qingyu No. 2 was by keeping much higher rate of proline accumulation and concentration of chlorophyll. The differences in salt tolerance showed that different adaptation mechanisms existed between cultivars of Jerusalem artichoke. The findings offered the possibility of selecting salt-tolerant genotypes of Jerusalem artichoke.

To explore genetic variability for two Jerusalem artichoke ( Helianthus tuberosus ) cultivars, N1 (the sixth-generation cultivated with 75% seawater irrigation for six years) and N7 (a general variety), a experiment was conducted to study the changes in physiological attributes under different concentrations (0%, 10% and 25% of seawater concentration in greenhouse and 0%, 30% and 50% of seawater concentration in the field) of seawater salinity stress. In the greenhouse experiment, decreases of dry growth rate, but increases of dry matter percentage and membrane injury occurred in both the genotypes at 10% and 25% seawater treatments, although lesser cell membrane damage was observed in N1 than N7. N1 accumulated greater contents of Na + , Cl − , soluble sugar and proline in leaves compared with N7. In the field experiment, the yields of shoot, root and tuber, and the contents of total-sugar and inulin in tubers of N1 were higher than those of N7. Lesser degree of salt injury in N1 indicated that the relatively salt-tolerant cultivar had higher K + /Na + ratio, lower Na + /Ca 2+ ratio, and the salt-induced enhancement of osmotic adjustment.

Two Jerusalem artichoke (Helianthus tuberosus L.) genotypes, NY-1 and NY-7, were subjected to different seawater concentrations (0, 10, 20, 30, 40, and 50%) for various periods of time to determine the effects on seedling growth, ion content, and photosynthetic productivity in a greenhouse. Under different seawater concentrations, sprouting rates varied greatly among the genotypes. The differences in relative growth rate (RGR), leaf chlorophyll content, total leaf area (TLA), plant dry weight (PDW), photosynthetic rate (A), stomatal conductance (gs), and efficiency of the light harvesting of photosystem II (Fv′/Fm′) were significant between NY-1 and NY-7 after 12 days of stress at 40 and 50% seawater. Seawater treatments resulted in the reduction of almost all the growth parameters and coincident increases of Na+ and Ca2+ concentrations in plant tissues. Our results indicate that there is great variability for seawater tolerance among H. tuberosus varieties, and that greater photosynthesis capacity, higher RGR, and relatively higher tissue Na+ accumulation at high seawater concentrations appears to be associated with seawater tolerance in H. tuberosus varieties.

Proline accumulation is an important mechanism for osmotic regulation under salt stress. In this study, we evaluated proline accumulation profiles in roots, stems and leaves of Jerusalem artichoke (Helianthus tuberosus L.) plantlets under NaCl stress. We also examined HtP5CS, HtOAT and HtPDH enzyme activities and gene expression patterns of putative HtP5CS1, HtP5CS2, HtOAT, HtPDH1, and HtPDH2 genes. The objective of our study was to characterize the proline regulation mechanisms of Jerusalem artichoke, a moderately salt tolerant species, under NaCl stress. Jerusalem artichoke plantlets were observed to accumulate proline in roots, stems and leaves during salt stress. HtP5CS enzyme activities were increased under NaCl stress, while HtOAT and HtPDH activities generally repressed. Transcript levels of HtP5CS2 increased while transcript levels of HtOAT, HtPDH1 and HtPDH2 generally decreased in response to NaCl stress. Our results supports that for Jerusalem artichoke, proline synthesis under salt stress is mainly through the Glu pathway, and HtP5CS2 is predominant in this process while HtOAT plays a less important role. Both HtPDH genes may function in proline degradation.