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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.

Self-germinated seedlings of Citrus sinensis and C. grandis were supplied with nutrient solution with 0 mM AlCl(3)·6H(2)O (control, −Al) or 1 mM AlCl(3)·6H(2)O (+Al) for 18 weeks. The DW (Dry weights) of leaf, stem, shoot and the whole plant of C. grandis were decreased and the ratio of root DW to shoot DW in C. grandis were increased by Al, whereas these parameters of C. sinensis were not changed by Al. Al treatment dramatically decreased the sulfur (S) content in C. grandis roots and the phosphorus (P) content in both C. sinensis and C. grandis roots. More Al was transported to shoots and leaves in C. grandis than in C. sinensis under Al treatment. Al treatment has more adverse effects on C. grandis than on C. sinensis, as revealed by the higher production of superoxide anion (O(2)(·−)), H(2)O(2) and thiobarbituric acid reactive substace (TBARS) content in C. grandis roots. Via the Illumina sequencing technique, we successfully identified and quantified 12 and 16 differentially expressed miRNAs responding to Al stress in C. sinensis and C. grandis roots, respectively. The possible mechanism underlying different Al tolerance of C. sinensis and C. grandis were summarized as having following aspects: (a) enhancement of adventitious and lateral root development (miR160); (b) up-regulation of stress and signaling transduction related genes, such as SGT1, PLC and AAO (miR477, miR397 and miR398); (c) enhancement of citrate secretion (miR3627); (d) more flexible control of alternative glycolysis pathway and TCA cycle (miR3627 and miR482); (e) up-regulation of S-metabolism (miR172); (f) more flexible control of miRNA metabolism. For the first time, we showed that root development (miR160) and cell wall components (cas-miR5139, csi-miR12105) may play crucial roles in Al tolerance in citrus plants. In conclusion, our study provided a comprehensive profile of differentially expressed miRNAs in response to Al stress between two citrus plants differing in Al tolerance which further enriched our understanding of the molecular mechanism underlying Al tolerance in plants. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (10.1007/s12298-020-00895-y) contains supplementary material, which is available to authorized users.

Sustainable land management is considered an effective measure to ensure national food security by matching improved soil quality with enhanced crop productivity. The objectives of the present study were to evaluate soil quality status among paddy fields, vegetable fields, and pomelo orchards managed by continuous cropping systems, to establish a minimum data set (MDS) and recommend fertilization strategies to enhance productivity in acidic red soil hilly regions. The soil quality index (SQI) was calculated using the total data set (TDS) and MDS as indicator selection methods. A total of 51, 32, 44 soil samples (0–20 cm) were collected from different cropping systems in paddy fields, vegetable fields, and pomelo orchards, respectively. Eleven soil properties (geographical indicator (altitude) and 10 chemical parameters) were measured to evaluate soil quality of acidic red soil hilly regions in Pinghe County, Fujian province, southern China. The results showed significant differences in soil quality indicators between paddy fields, vegetable fields, and pomelo orchards, and altitude, total N, hydrolysable nitrogen (Avail-N), exchangeable calcium (Avail-Ca), and available zinc (Avail-Zn) were adopted as indicators in the MDS using principal component analysis (PCA). After scoring and weighting the selected indicators, the mean SQI scores of paddy field, vegetable field, and pomelo orchard soils were 0.267, 0.282, and 0.236 for TDS, respectively, and the SQI scores were 0.233, 0.256, and 0.166 for MDS. A significant positive correlation was observed between SQI and crop yield using both the TDS and MDS methods, indicating that the MDS method adequately represents the TDS method (R2 = 0.6839) for evaluation of the effects of farming production practices on soil quality. Overall, the unbalanced soil quality indicators, including rich in total N, organic matter and Avail-P, and deficient in Avail-N, Avail-K, Avail-Ca, Avail-Mg, Avail-Zn, and Avail-B, were implicated as the main limiting factors for crop production in the studied regions. Compared with paddy and vegetable fields, the lower total N and Avail-N status at higher altitudes were considered as the major factors limiting pomelo productivity. Therefore, there is a great potential for increasing pomelo productivity by improving soil quality with integrated nutrient management in acidic red soil hilly regions.

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.

Mg deficiency affected only the differentiation zone in Citrus sinensis roots. Phloem impairment in Mg-deficient leaves resulted in cell wall lignifications of both vascular cambium and spongy parenchyma cells. Decrease of Mg content in the roots hampered secondary lignification of endodermal and protoxylem cell walls. Magnesium (Mg) is a vital nutrient for plant photosynthesis and biochemical reactions. Mg deficiency is a widespread problem in citrus production, directly affecting yield and quality. With the objective of investigating the relationship among root and leaf anatomy, photosynthetic efficiency and mineral nutrient uptake and transport in Mg-deficient citrus, the effects of Mg deficiency in ‘Xuegan’ (Citrus sinensis) seedlings on plant biomass, leaf gas exchange and chlorophyll, nutrient contents, root and leaf anatomy and lignin biosynthesis were studied. Mg deficiency decreased root and shoot dry weight, leaf CO2 assimilation, chlorophyll and Mg, phosphorus (P), boron (B), cupper (Cu) and iron (Fe) concentration in roots, stems and leaves, and increased potassium (K) and calcium (Ca) concentration. Mg deficiency in the leaves resulted in phloem disorder that blocked phloem transport, leading to carbohydrate accumulation in chloroplasts and secondary lignifications of both vascular cambium and spongy parenchyma cell walls. In contrast, decrease of Mg content in the roots hampered lignin deposition on the protoxylem and endodermal cell walls at the root differentiation zone. Poorly lignified Casparian strip and xylem cell walls in Mg-deficient root tips might decrease xylem loading efficiency of nutrients, lowering their contents in the above ground parts, which ultimately inhibited whole plant growth.

We identified more Mn-toxicity-responsive genes from Mn-intolerant Citrus grandis than from Mn-tolerant Citrus sinensis roots. These findings increased our understanding of the molecular mechanisms on plant Mn toxicity and Mn tolerance. Manganese (Mn) toxicity is the most important factor limiting crop production after aluminum toxicity in acidic soils. However, little is known about Mn-toxicity-induced alterations of gene expression profiles in woody plants. Using cDNA-AFLP, we identified 87 and 63 Mn-toxicity-responsive genes from Mn-intolerant ‘Sour pummelo’ (Citrus grandis) and Mn-tolerant ‘Xuegan’ (Citrus sinensis) roots. Among these genes, only 22 genes with the same accession number were shared by both. Protein phosphorylation/dephosphorylation-related genes were upregulated in C. sinensis roots, and downregulated in C. grandis roots except for one differentially expressed gene. Sulfur metabolism-related genes were repressed only in Mn-toxic C. grandis roots. Obviously, great differences existed in Mn-toxicity-induced alterations of gene expression profiles between C. sinensis and C. grandis roots. Genes related to protein phosphorylation/dephosphorylation (i.e., cyclin-dependent kinase-activating kinase assembly factor-related protein and PP2A regulatory subunit TAP46), cellular transport (i.e., Ca-transporting ATPase 1), and nucleic acid (i.e., ethylene-responsive transcription factor ERF109-like, structural maintenance of chromosomes protein 4-like, RNA-binding protein and DEAD-box ATP-dependent RNA helicase 21), cell wall (i.e., pectin methylesterase 1 and invertase/pectin methylesterase inhibitor family protein) and fatty acid (i.e., carboxylesterase 20) metabolisms might play a role in C. sinensis Mn tolerance. In addition, cell wall materials were increased in Mn-toxic C. sinensis and C. grandis roots, especially in the former. Interestingly, lignin content was increased in Mn-toxic C. sinensis roots, while a reverse trend was displayed in Mn-toxic C. grandis roots. In conclusion, our results provided novel clues to the molecular mechanisms on Mn toxicity and Mn tolerance in higher plants.

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.

Eighty-six differentially abundant proteins were identified in Citrus grandis roots in response to boron-deficiency using the iTRAQ technique and possible mechanism underlying boron-deficiency tolerance of citrus plants was identified. Boron (B) is an essential element for plant growth and development and adequate B supply is an important determinant of good quality and high yield of crops. B-deficiency is a worldwide problem in agricultural production including citrus. However, little is known about the molecular mechanism of plant tolerance to B-deficiency. Using the iTRAQ technique, 86 differentially abundant proteins were identified from B-deficient Citrus grandis roots. The adaptive strategy of C. grandis roots under B-deficiency was summarized as follows: (1) enhancement of alternative splicing of mRNA and DNA methylation; (2) up-regulation of post-translation modification (PTM) and turnover of proteins; (3) reinforcement of cellular transport; (4) enhancement of antioxidant system and signal transduction. In general, these results increase our understanding of molecular mechanisms underlining the resistance of citrus plant under B-deficiency. Further studies should focus on how do roots perceive B deficiency in the rhizosphere and which pathway or proteins react to this adverse condition in the first place and then stimulates the downstream responses in Citrus plants.

Typical toxic symptom only occurred in B-toxic C. grandis leaves. B-toxicity induced PCD of C. grandis leaf phloem tissue. The lower leaf free B might contribute to the higher B-tolerance of C. sinensis. Seedlings of ‘Xuegan’ (Citrus sinensis) and ‘Sour pummelo’ (Citrus grandis) differing in boron (B)-tolerance were irrigated with nutrient solution containing 10 (control) or 400 (B-toxic) μM H3BO3 for 15 weeks. Thereafter, the effects of B-toxicity on leaf photosynthesis, chlorophyll, plant B absorption and distribution, root and leaf anatomy were investigated to elucidate the possible B-tolerant mechanisms of Citrus plants. Typical toxic symptom only occurred in B-toxic C. grandis leaves. Similarly, B-toxicity only affected C. grandis photosynthesis and chlorophyll. Although total B concentration in B-toxic roots and leaves was similar between the two species, leaves from B-toxic C. grandis plant middle had higher free B and lower bound B as compared with those from C. sinensis. Effects of B-toxicity on leaf structure were mainly limited to the mesophyll cells and the phloem of leaf veins. Although irregular cell wall thickening was observed in leaf cortex cells and phloem tissue of B-toxic C. grandis and C. sinensis leaves, exocytosis only occurred in the companion cells and the parenchyma cells of B-toxic C. sinensis leaf phloem. Also, B-toxicity induced cell death of phloem tissue through autophagy in C. grandis leaf veins. B-toxicity caused death of root epidermal cells of the two Citrus species. B-toxicity restrained degradation of middle lamella, but did not alter ultrastructure of Golgi apparatus and mitochondria in root elongating zone cells. In conclusion, C. sinensis was more tolerant to B-toxicity than C. grandis. The lower leaf free B and higher bound B might contribute to the higher B-tolerance of C. sinensis.

Fifty-three and thirty-nine differentially expressed protein spots were isolated from Mn-toxic Citrus sinensis and Citrus grandis roots, respectively. Mn-toxicity-induced changes in protein profiles greatly differed between the two species. Limited information is available on the manganese (Mn)-toxicity-responsive proteins in plant roots. ‘Sour pummelo’ (Citrus grandis) and ‘Xuegan’ (Citrus sinensis) seedlings were irrigated for 17 weeks with 2 (control) or 600 μM (Mn-toxic) MnSO4. C. sinensis displayed more tolerance to Mn-toxicity than C. grandis, which may be related to more Mn accumulation in roots and less Mn distribution in shoots. Using two-dimensional electrophoresis (2-DE), we isolated 11 up-regulated and 42 down-regulated protein spots from Mn-toxic C. sinensis roots, and 25 up-regulated and 14 down-regulated protein spots from Mn-toxic C. grandis roots. This indicated more metabolic flexibility in C. sinensis roots, thus contributing to the Mn-tolerance of C. sinensis. According to the biological functional properties, these differentially expressed proteins in the two species were classified into the following categories: protein metabolism, nucleic acid metabolism, carbohydrate and energy metabolism, stress responses, cell wall and cytoskeleton, cell transport, signal transduction and fatty acid metabolism. Under Mn-toxicity, proteins involved in nucleic acid metabolism, glycolysis and cell transport were up-regulated in nontolerant C. grandis roots, and down-regulated in tolerant C. sinensis roots. The notable down-regulation of proteins in Mn-toxic C. sinensis roots with less accumulation of carbohydrates may provide an advantage to the net carbon balance by lowering related metabolic processes, and enhancing the Mn-tolerance of C. sinensis. To conclude, there are many important differences in Mn-toxicity-induced changes in protein profiles and metabolic responses between the two species.

Self-rooted, 10-month-old, uniform tea [Camellia sinensis (L.) O. Kuntze cv. Huangguanyin] trees were supplied for 17 weeks with 0, 40, 80, 160, 400 or 1,000 μM phosphorus (P). Thereafter, plant growth, leaf malondialdehyde (MDA), antioxidant enzymes and metabolites were measured to determine how antioxidant system functions to protect P-deficient leaves from photo-oxidative damage. Whole plant dry weight increased as P supply increased from 0 to 160 μM, and then kept unchanged with further increasing P supply. Phosphorus-deficient leaves displayed lower or similar activities of superoxide dismutase, ascorbate peroxidase, monodehydroascorbate reductase, dehydroascorbate reductase, glutathione reductase and catalase and contents of ascorbate and reduced glutathione depending on the degree of P-deficiency and enzyme species, but P supply did not significantly affect leaf MDA content. In conclusion, the antioxidant system in P-deficient leaves provides considerable protection to them against photo-oxidative damage.

Magnesium (Mg)-deficiency affects productivity and quality in agriculture, yet at a physiological level it is not well understood. Citrus grandis and Citrus sinensis seedlings were irrigated for 12 weeks with 0, 50, 500 or 2,000 μM MgSO4. Thereafter, Mg-deficiency-induced changes in photosynthesis, antioxidant system and carbohydrates were investigated. Mg-deficiency affected CO2 assimilation more in C. grandis leaves than in C. sinensis ones, but Mg-deficiency-induced accumulation of sugars was not higher in the former except for sucrose. Mg-deficiency-induced photoinhibitory impairment occurring on the whole photosynthetic electron transport chain was more severe in C. grandis leaves than in C. sinensis ones. Mg-deficient leaves had higher or similar activities of antioxidant enzymes and contents of antioxidant metabolites except for catalase (CAT) activity and reduced glutathione (GSH) content. However, Mg-deficiency increased leaf malondialdehyde (MDA) content. In conclusion, the greater decrease in CO2 assimilation in Mg-deficient C. grandis leaves may be caused by the greater decrease in the photosynthetic electron transport capacity. Mg-deficiency-induced up-regulation in leaf antioxidant system does not provide enough protection to Mg-deficient leaves against the oxidative damage.

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To investigate the photoinhibition of photosynthesis in ‘Honeycrisp’ apple (Malus domestica Borkh. cv. Gala) leaves with zonal chlorosis, we compared pigments, CO2 assimilation and chlorophyll (Chl) a fluorescence (OJIP) transient between chlorotic leaves and normal ones. Chl and carotenoids (Car) contents, Chl a/b ratio, and absorptance were lower in chlorotic leaves than in normal ones, whereas Car/Chl ratio was higher in the former. Although CO2 assimilation and stomatal conductance were lower in chlorotic leaves, intercellular CO2 concentration did not differ significantly between the two leaf types. Compared with normal leaves, chlorotic ones had increased deactivation of oxygen-evolving complexes (OEC), minimum fluorescence (F o), dissipated energy, relative variable fluorescence at L-, W-, J- and I-steps, and decreased maximum fluorescence (F m), maximum quantum yield for primary photochemistry (F v /F m or TRo/ABS), quantum yield for electron transport (ETo/ABS), quantum yield for the reduction of end acceptors of photosystem I (PSI) (φRo and REo/ABS), maximum amplitude of IP phase, amount of active photosystem II (PSII) reaction centers (RCs) per cross section (CS) and total performance index (PItot,abs). In conclusion, photoinhibition occurs at both the donor (i.e., the OEC) and the acceptor sides of PSII in chlorotic leaves. The acceptor side is damaged more severely than the donor side, which possibly is the consequence of over-reduction of PSII due to the slowdown of Calvin cycle. In addition to decreasing light absorptance by lowering Chl level, energy dissipation is enhanced to protect chlorotic leaves from photo-oxidative damage.

Background Magnesium (Mg)-deficiency is frequently observed in Citrus plantations and is responsible for the loss of productivity and poor fruit quality. Knowledge on the effects of Mg-deficiency on upstream targets is scarce. Seedlings of ‘Xuegan’ [Citrus sinensis (L.) Osbeck] were irrigated with Mg-deficient (0 mM MgSO4) or Mg-sufficient (1 mM MgSO4) nutrient solution for 16 weeks. Thereafter, we first investigated the proteomic responses of C. sinensis roots and leaves to Mg-deficiency using two-dimensional electrophoresis (2-DE) in order to (a) enrich our understanding of the molecular mechanisms of plants to deal with Mg-deficiency and (b) understand the molecular mechanisms by which Mg-deficiency lead to a decrease in photosynthesis. Results Fifty-nine upregulated and 31 downregulated protein spots were isolated in Mg-deficient leaves, while only 19 upregulated and 12 downregulated protein spots in Mg-deficient roots. Many Mg-deficiency-responsive proteins were involved in carbohydrate and energy metabolism, followed by protein metabolism, stress responses, nucleic acid metabolism, cell wall and cytoskeleton metabolism, lipid metabolism and cell transport. The larger changes in leaf proteome versus root one in response to Mg-deficiency was further supported by our observation that total soluble protein concentration was decreased by Mg-deficiency in leaves, but unaffected in roots. Mg-deficiency had decreased levels of proteins [i.e. ribulose-1,5-bisphosphate carboxylase (Rubisco), rubisco activase, oxygen evolving enhancer protein 1, photosynthetic electron transfer-like protein, ferredoxin-NADP reductase (FNR), aldolase] involved in photosynthesis, thus decreasing leaf photosynthesis. To cope with Mg-deficiency, C. sinensis leaves and roots might respond adaptively to Mg-deficiency through: improving leaf respiration and lowering root respiration, but increasing (decreasing) the levels of proteins related to ATP synthase in roots (leaves); enhancing the levels of proteins involved in reactive oxygen species (ROS) scavenging and other stress-responsive proteins; accelerating proteolytic cleavage of proteins by proteases, protein transport and amino acid metabolism; and upregulating the levels of proteins involved in cell wall and cytoskeleton metabolism. Conclusions Our results demonstrated that proteomics were more affected by long-term Mg-deficiency in leaves than in roots, and that the adaptive responses differed between roots and leaves when exposed to long-term Mg-deficiency. Mg-deficiency decreased the levels of many proteins involved in photosynthesis, thus decreasing leaf photosynthesis. Electronic supplementary material The online version of this article (doi:10.1186/s12864-015-1462-z) contains supplementary material, which is available to authorized users.

Three hundred and nineteen ,Guanximiyou, pummelo (Citrus grandis) orchards from Pinghe county, the southern region of Fujian province, China, were selected for this study. The objectives were to determine (i) the soil and leafnutrient status, (ii) the relationships between leaf mineral elements and the corresponding soil elements, and (iii) the relationships between fruit quality and mineral nutrients. The results showed that soil acidification was a major problem in these orchards, with an average pH of 4.34. Soil acidification affected the availability of soil N, P, Ca, Mg, S, B, Cu and Zn and the levels of organic matter (OM) and cation exchange capacity (CEC), thus inducing soil and leaf nutrient imbalance. Indeed, severe nutrient imbalance existed in these orchard soils. 77.4% and 65.8% of soils were sub-optimum in exchangeable Mg and Ca, while 96.6% and 82.1% of soils were super-optimum in available S and P, respectively. Besides, severe nutrient deficiencies and excesses co-existed in leaves. 46.8% and 35.6% of leaves were deficient in N and Mg, while 74.8% and 70.4% of leaves were excess in B and Cu, respectively. Regressive analysis showed that leaf content of mineral elements was poorly related with the available content of the corresponding soil elements, respectively. In some orchards, severe juice sac granulation, an important factor affecting fruit quality, was observed. Regressive analysis indicated that Mg, S, Cu and Mn played a role in juice sac granulation of fruits. In conclusion, soil acidification might lead to severe soil nutrient imbalance, thus inducing leaf nutrient imbalance, eventually impairing fruit quality parameters such sac granulation.

Four scion-rootstock combination [i.e., X/X and X/SP, ‘Xuegan’ (Citrus sinensis) grafted on ‘Xugan’ and ‘Sour pummelo’ (Citrus grandis), respectively, and SP/X and SP/SP, ‘Sour pummelo’ grafted on ‘Xuegan’ and ‘Sour pummelo’, respectively] plants were treated for 18 weeks with 0 (−Al) or 1.2 mM AlCl3·6H2O (+Al). Thereafter, leaf, stem and root concentrations of phosphorus and aluminum (Al), leaf and root levels of organic acids (OAs), Al-induced release of OA anions (i.e., malate and citrate), photosynthesis and chlorophyll a fluorescence (OJIP) transients were measured. Al-induced decrease of photosynthesis and damage of photosynthetic electron transport chain were less pronounced in X/X and X/SP leaves than in SP/SP and SP/X leaves, which might be related with the higher Al-induced root efflux of OA anions and leaf P concentration. C. sinensis rootstock alleviated the influences of Al-toxicity on leaf photosynthetic electron transport chain by enhancing Al-induced release of root OA anions, hence lessening Al-induced photosynthesis inhibition in SP/X plants, while the reverse was the case for C. grandis rootstock in X/SP plants. In conclusion, the tolerance of grafted Citrus plants to Al depends on the scion as well as rootstock genotype, and the scion-rootstock interaction.

Background Boron (B)-toxicity is an important disorder in agricultural regions across the world. Seedlings of ‘Sour pummelo’ (Citrus grandis) and ‘Xuegan’ (Citrus sinensis) were fertigated every other day until drip with 10 μM (control) or 400 μM (B-toxic) H3BO3 in a complete nutrient solution for 15 weeks. The aims of this study were to elucidate the adaptive mechanisms of citrus plants to B-toxicity and to identify B-tolerant genes. Results B-toxicity-induced changes in seedlings growth, leaf CO2 assimilation, pigments, total soluble protein, malondialdehyde (MDA) and phosphorus were less pronounced in C. sinensis than in C. grandis. B concentration was higher in B-toxic C. sinensis leaves than in B-toxic C. grandis ones. Here we successfully used cDNA-AFLP to isolate 67 up-regulated and 65 down-regulated transcript-derived fragments (TDFs) from B-toxic C. grandis leaves, whilst only 31 up-regulated and 37 down-regulated TDFs from B-toxic C. sinensis ones, demonstrating that gene expression is less affected in B-toxic C. sinensis leaves than in B-toxic C. grandis ones. These differentially expressed TDFs were related to signal transduction, carbohydrate and energy metabolism, nucleic acid metabolism, protein and amino acid metabolism, lipid metabolism, cell wall and cytoskeleton modification, stress responses and cell transport. The higher B-tolerance of C. sinensis might be related to the findings that B-toxic C. sinensis leaves had higher expression levels of genes involved in photosynthesis, which might contribute to the higher photosyntheis and light utilization and less excess light energy, and in reactive oxygen species (ROS) scavenging compared to B-toxic C. grandis leaves, thus preventing them from photo-oxidative damage. In addition, B-toxicity-induced alteration in the expression levels of genes encoding inorganic pyrophosphatase 1, AT4G01850 and methionine synthase differed between the two species, which might play a role in the B-tolerance of C. sinensis. Conclusions C. sinensis leaves could tolerate higher level of B than C. grandis ones, thus improving the B-tolerance of C. sinensis plants. Our findings reveal some novel mechanisms on the tolerance of plants to B-toxicity at the gene expression level. Electronic supplementary material The online version of this article (doi:10.1186/s12870-014-0284-5) contains supplementary material, which is available to authorized users.

Background Very little is known about manganese (Mn)-toxicity-responsive genes in citrus plants. Seedlings of ‘Xuegan’ (Citrus sinensis) and ‘Sour pummelo’ (Citrus grandis) were irrigated for 17 weeks with nutrient solution containing 2 μM (control) or 600 μM (Mn-toxicity) MnSO4. The objectives of this study were to understand the mechanisms of citrus Mn-tolerance and to identify differentially expressed genes, which might be involved in Mn-tolerance. Results Under Mn-toxicity, the majority of Mn in seedlings was retained in the roots; C. sinensis seedlings accumulated more Mn in roots and less Mn in shoots (leaves) than C. grandis ones and Mn concentration was lower in Mn-toxicity C. sinensis leaves compared to Mn-toxicity C. grandis ones. Mn-toxicity affected C. grandis seedling growth, leaf CO2 assimilation, total soluble concentration, phosphorus (P) and magenisum (Mg) more than C. sinensis. Using cDNA-AFLP, we isolated 42 up-regulated and 80 down-regulated genes in Mn-toxicity C. grandis leaves. They were grouped into the following functional categories: biological regulation and signal transduction, carbohydrate and energy metabolism, nucleic acid metabolism, protein metabolism, lipid metabolism, cell wall metabolism, stress responses and cell transport. However, only 7 up-regulated and 8 down-regulated genes were identified in Mn-toxicity C. sinensis ones. The responses of C. grandis leaves to Mn-toxicity might include following several aspects: (1) accelerating leaf senescence; (2) activating the metabolic pathway related to ATPase synthesis and reducing power production; (3) decreasing cell transport; (4) inhibiting protein and nucleic acid metabolisms; (5) impairing the formation of cell wall; and (6) triggering multiple signal transduction pathways. We also identified many new Mn-toxicity-responsive genes involved in biological and signal transduction, carbohydrate and protein metabolisms, stress responses and cell transport. Conclusions Our results demonstrated that C. sinensis was more tolerant to Mn-toxicity than C. grandis, and that Mn-toxicity affected gene expression far less in C. sinensis leaves. This might be associated with more Mn accumulation in roots and less Mn accumulation in leaves of Mn-toxicity C. sinensis seedlings than those of C. grandis seedlings. Our findings increase our understanding of the molecular mechanisms involved in the responses of plants to Mn-toxicity.

BackgroundVery little is known about the effects of manganese (Mn)-excess on citrus photosynthesis and antioxidant systems. Seedlings of sour pummelo (Citrus grandis) were irrigated for 17 weeks with nutrient solution containing 2 μM (control) or 500 μM (excess) MnSO4. The objective of this study were to understand the mechanisms by which Mn-excess leads to a decrease in CO2assimilation and to test the hypothesis that Mn-induced changes in antioxidant systems differ between roots and leaves.ResultsMn-excess decreased CO2assimilation and stomatal conductance, increased intercellular CO2concentration, but did not affect chlorophyll (Chl) level. Both initial and total ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) activity in Mn-excess leaves decreased to a lesser extent than CO2assimilation. Contents of glucose, fructose, starch and total nonstructural carbohydrates did not differ between Mn-excess leaves and controls, while sucrose content was higher in the former. Chl a fluorescence (OJIP) transients from Mn-excess leaves showed increased O-step and decreased P-step, accompanied by positive L- and K-bands. Mn-excess decreased maximum quantum yield of primary photochemistry (Fv/Fm) and total performance index (PItot,abs), but increased relative variable fluorescence at I-steps (VI) and energy dissipation. On a protein basis, Mn-excess leaves displayed higher activities of monodehydroascorbate reductase (MDAR), glutathione reductase (GR), superoxide dismutase (SOD), catalase (CAT) and guaiacol peroxidase (GPX) and contents of antioxidants, similar ascorbate peroxidase (APX) activities and lower dehydroascorbate reductase (DHAR) activities; while Mn-excess roots had similar or lower activities of antioxidant enzymes and contents of antioxidants. Mn-excess did not affect malondialdehyde (MDA) content of roots and leaves.ConclusionsMn-excess impaired the whole photosynthetic electron transport chain from the donor side of photosystem II (PSII) up to the reduction of end acceptors of photosystem I (PSI), thus limiting the production of reducing equivalents, and hence the rate of CO2assimilation. Both the energy dissipation and the antioxidant systems were enhanced in Mn-excess leaves, while the antioxidant systems in Mn-excess roots were not up-regulated, but still remained high activity. The antioxidant systems in Mn-excess roots and leaves provided sufficient protection to them against oxidative damage.