The role of plant eIF5A proteins in multiple biological processes, such as protein synthesis regulation, translation elongation, mRNA turnover, programmed cell death and stress tolerance is well known. Toward using th...The role of plant eIF5A proteins in multiple biological processes, such as protein synthesis regulation, translation elongation, mRNA turnover, programmed cell death and stress tolerance is well known. Toward using these powerful proteins to increase stress tolerance in agricultural plants, in the present study, we cloned and characterized PsneIFSA2 and PsneIFSA4 from young poplar (P. simonii × P. nigra) leaves. The deduced amino acid sequences of PsneIF5A2 and PsneIF5A4 were 98 % similar to each other, and they are orthologs of eIF5A 1 in Arabidopsis. In a subcellular localization analysis, PsneIF5A2 and PsneIF5A4 proteins were localized in the nucleus and cytoplasm, qRT-PCR analysis showed that PsneIF5A2 and PsneIF5A4 were transcribed in poplar flowers, stem, leaves, and roots. In addition, they were also induced by abiotic stresses. Transgenic yeast expressing PsneIF5A2 and PsneIF5A4 had increased salt, heavy metal, osmotic, oxidative tolerance. Our results suggest that PsneIF5A2 and PsneIF5A4 are excellent candidates for genetic engineering to improve salt and heavy metal tolerance in agricultural plants.展开更多
Aldehydes play an important role in atmospheric chemistry and plant direct and indirect defense against environmental stresses.In this study,the emissions of saturated C6-C10 aldehydes from Populus simonii × P.py...Aldehydes play an important role in atmospheric chemistry and plant direct and indirect defense against environmental stresses.In this study,the emissions of saturated C6-C10 aldehydes from Populus simonii × P.pyramidalis 'Opera 8277' cuttings were examined by using a gas chromatography/mass spectrometry(GC/MS) technique at three levels of light intensity(400,800 and 1 200 μmol·m-2·s-1).A positive correlation between the emissions of these aldehydes and light intensity was found.Moreover,nordi-hydroguaiaretic acid(NDGA),a special inhibitor of lipoxygenase(LOX),significantly inhibited the emissions of C6-C9 aldehydes at three levels of light intensity,but did not influence the emission of decanal(C10).The emissions of C6-C10 aldehydes in NDGA treated poplar cuttings,exhibited the same positive correlation with light intensity.The results indicated that LOX pathway contributes to the emissions of C6-C9 aldehydes,whereas some pathways regulated by light intensity might be a universal mechanism for emissions of C6-C10 aldehydes.展开更多
Populus euphratica Oliv. is of high salinity tolerance and used as a model species for investigating molecular mechanisms of trees' responses to salt stress. In the work presented here we found that calli of P. euphr...Populus euphratica Oliv. is of high salinity tolerance and used as a model species for investigating molecular mechanisms of trees' responses to salt stress. In the work presented here we found that calli of P. euphratica grew more rapidly and accumulated less Na+, but more K+, under salt stress than those of salt-sensitive poplar, Populus hopeiensis. Different types of Na+/H+ antiporters (SOS1, NhaD1 and NHX1) were isolated from P. euphratica; all of these genes have been shown to play important roles in plant salt tolerance mechanism in previous studies. Expression profiles of these three genes were compared between P. hopeiensis and P. euphratica in the presence and absence of salt stress by real-time PCR. The three genes were induced in both P. euphratica and P. hopeiensis by salt. Transcript levels of PeNHX1 were lower in P. euphratica than in P. hopeiensis under 150 mM NaCl stress. In addition, transcript levels of PeNhaD1 were lower, while PeSOS1 were higher in P. euphratica than in P. hopeiensis under both stressed and unstressed conditions. The results indicated that P. euphratica up-regulates different genes and consistently maintains both effluxes of Na+ and high K+ levels. Our data suggests that differences in gene expression patterns may contribute to the dif-ference in salt tolerance between these two poplars.展开更多
Populus euphratica and Populuspruinosa, sister species in the Turanga Section (Salicaceae), growing in semi-arid saline areas are known for their high salinity tolerance. In this study, by combining growth level wit...Populus euphratica and Populuspruinosa, sister species in the Turanga Section (Salicaceae), growing in semi-arid saline areas are known for their high salinity tolerance. In this study, by combining growth level with Na+ and K+ contents, the expression level of vacuolar Na+/H+ antiporters was investigated for NaCl-induced changes in P. euphratica and t3. pru- inosa calli. Compared to R euphratica, P. pruinosa calli grew well in 200 mM NaC1 stress from 14. to 21 days. Increasing the stressed time caused an increase in Na+ content concomitant with a decrease in K+ content in P. euphratica calli, whereas, with the presence of 200 mM NaCI, K+ content has a less increase in 14 and 21 days than in 7 days which was detected in R pruinosa calli. The transcript levels of six genes coding for NHX-type Na+/H+ antiporters suggest that vacuolar NHX1-NHX6 antiporters play important roles in responding to salt stress in R pruinosa. Our data suggest that there exists a higher salt tolerance for P. pruinosa than P. euphratica at the cellular level, Na+ avoidance or accumulation is observed in cellular compartments, and that expression of NHX antiporters is linked to the accumulator phenotype.展开更多
[Objectives]This study was conducted to investigate the photosynthetic physiological characteristics of Populus euphratica trees at different forest ages in the desert area of Minqin in response to drought stress.[Met...[Objectives]This study was conducted to investigate the photosynthetic physiological characteristics of Populus euphratica trees at different forest ages in the desert area of Minqin in response to drought stress.[Methods]With P.euphratica trees of different ages in Minqin as the research object,the water characteristics and photosynthetic physiological indexes(chlorophyll,soluble sugar,POD,SOD and MDA)were compared under different conditions.[Results]On the time gradient,the soil water contents of P.euphratica of different ages decreased continuously with the extension of irrigation stop time,but the decreases were small.Under the same stress treatment,the soil water contents of P.euphratica of different ages increased with the deepening of soil layer,but the differences were not significant(P>0.05).Under normal condition,the chlorophyll contents of P.euphratica at three ages gradually increased with the increase of age.The chlorophyll contents in leaves of P.euphratica at different ages were all lower under normal condition and higher under water stress.The changes of POD and SOD activity in leaves of P.euphratica at different ages under different conditions were basically the same,showing that the enzyme activity was higher under water stress than under normal under.The MDA contents in leaves of the young and middle P.euphratica forests were higher under water stress at 7 and 21 d than under normal condition,and the differences were not significant(P>0.05);and the MDA content in leaves of the mature P.euphratica forest was higher under water stress at 21 and 35 d than under normal condition.Drought stress has a certain effect on the photosynthetic physiological characteristics of P.euphratica.In summary,under drought stress,the chlorophyll content,SOD and POD activity,and MDA molar concentration in the leaves of P.euphratica were basically higher than under normal condition,indicating that P.euphratica could resist drought environment through osmotic adjustment and showed strong drought resistance.[Conclusions]This study provides a theoretical reference for the restoration,protection and reconstruction of natural P.euphratica forests in the desert area of Minqin.展开更多
Abstract The effects of water potential, NaCl and Na2SO4 on germination and radicle growth of two riparian tree species, Populus euphratica Oliv. and P. pruinosa Schrenk (Salicaceae), were tested. Growth chamber stu...Abstract The effects of water potential, NaCl and Na2SO4 on germination and radicle growth of two riparian tree species, Populus euphratica Oliv. and P. pruinosa Schrenk (Salicaceae), were tested. Growth chamber studies revealed an optimum temperature range for seed germination of both species between 15-35℃. The final germination percentage of both species decreases with decreasing water potential in all types of solution applied in the experiments. P. pruinosa was less tolerant to low ψw stress than P. euphratica, especially in salt solutions. Germination percentages fell below 20% for P. pruinosa at -0.6 MPa (NaCl) or -0.4 MPa (Na2SO4) and for P. euphratica at -1.2 MPa (NaCl) or -0.6 MPa (Na2SO4). Radicle growth of both species was inhibited by high concentrations of PEG1 NaCl and Na2SO4. However, growth was enhanced at -0.13 and -0.29 MPa in PEG or at -0.13 MPa in NaCl solutions compared to distilled water. Radicle growth of P. euphratica was higher than that of P pruinosa. Germination and radicle growth of both species exhibited ion toxicity. Na2SO4 was more toxic than iso-osmotic solutions of NaCl. Radicle growth proved to be more sensitive than seed germination. Thus, flooding does not only yield the necessary soil moisture for germination but also favors seedling establishment of both species through leaching of salts from the soil surface. The different sensitivity of the species during their early growth stages might, moreover, contribute to the observed differences in their distribution in the Talim Basin (northwest China).展开更多
We used a salt-resistant poplar genotype Populus euphratica and two salt-sensitive genotypes, Populus 'popularis 35-44' (P. popularis) and the hybrid P. talassica Kom x (P. euphratica + Salix alba L.) to exami...We used a salt-resistant poplar genotype Populus euphratica and two salt-sensitive genotypes, Populus 'popularis 35-44' (P. popularis) and the hybrid P. talassica Kom x (P. euphratica + Salix alba L.) to examine genotypic differences in nutrient selectivity under NaCl stress. One-year-old seedlings of P. euphratica and one-year-old hardwood cuttings of P. popularis were used in a short-term study (24 hours), while in a long-term study, up to 4 weeks, two-year-old seedlings of P. euphratica and the hybrid P. talassica Kom x (P. euphratica + Salix alba L.) were compared. In the short-term study, K+ concentration in the xylem sap ([K+]xylem) of P. euphratica significantly increased after salt stress was initiated, and maintained 1-2 fold higher than control levels during the period of salt stress (24 hours). Xylem Ca2+ and Mg2+ concentrations ([Ca2+]xylem, [Mg2+]xylem) in P. euphratica resembled the pattern of K+ despite a lesser magnitude in elevation. However, [K+]xylem, [Ca2+]xylem and [Mg2+]xylem in P. popularis exhibited a transient increase at the beginning of salt treatment, thereafter, they all returned to control levels at 4 hours and no further rise was observed in the following hours. Xylem Na+/K+, Na+/Ca2+ and Na+/Mg2+ in P. popularis increased sharply upon NaCl stress and steadily reached the maximum at 24 hours. In contrast, xylem Na+/K+, Na+/Ca2+ and Na+/Mg2+ in NaCl-treated plants of P. euphratica did not significantly increase during the period of salt stress (24 hours). Noteworthy, Na+/K+ markedly declined after the onset of stress. These results suggest that P. euphratica had a higher nutrient selectivity in face of salinity. A same trend was observed in a 4-week study. Xylem Na+/K+, Na+/Ca2+ and Na+/Mg2+ in salinised plants of the hybrid abruptly increased after 4 days of stress, and then continuously increased to reach the highest level at day 8 or day 15. In comparison, the magnitude of Na+/K+, Na+/Ca2+ and Na+/Mg2+ elevation in the xylem of P. euphratica was much lower during the observation period. In conclusion, salt-tolerant genotype P. euphratica maintained a higher nutrient selectivity under saline stress, as compared to the two salt-sensitive genotypes. The high capacity for nutrient uptake and transport presumably contributes to the salt tolerance of P. euphratica in a longer term.展开更多
The aim of this study was to investigate the role of rhizosphere organic phosphorus (P) in soil P supply in semiarid forests and the effects of tree species on rihizosphere organic P. We examined organic P fractions...The aim of this study was to investigate the role of rhizosphere organic phosphorus (P) in soil P supply in semiarid forests and the effects of tree species on rihizosphere organic P. We examined organic P fractions in rhizosphere and bulk soils of mono-specific Simon poplar (Populus simonii) and Mongolian pine (Pinus sylvestris var. mongolica) plantations in a semiarid sandy soil of Horqin Sandy Land in Northeast China. Total organic P (TPo) accounted for 76% of total P across the two stands. The concentration of organic P (Po) fractions decreased in the order of NaOH-Po〉Res-Po〉HCI-Po〉NaHCO3-Po in both plantations. The concentration of NaHCO3-Po was 38% and 43% lower in rhizosphere soil than in bulk soil in Simon poplar and Mongolian pine plantations, respectively. In contrast, total P, TPo and NaOH-Po significantly accumulated in rhizosphere soil in Simon poplar plantations, but no change in Mongolian pine plantations. Soil recalcitrant organic P fractions were positively correlated with soil organic carbon. The results suggest that rhizosphere labile organic P was an important source of plant-available P in this semiarid region, but the dynamic of rhizosphere recalcitrant organic P fractions varied with tree species and was correlated to organic carbon dynamics.展开更多
基金supported by the Program for New Century Excellent Talents in University(No.NCET-12-0808)National Natural Science Foundation of China(No.31370661)the Fundamental Research Funds for the Central Universities(No.2572014AA26)
文摘The role of plant eIF5A proteins in multiple biological processes, such as protein synthesis regulation, translation elongation, mRNA turnover, programmed cell death and stress tolerance is well known. Toward using these powerful proteins to increase stress tolerance in agricultural plants, in the present study, we cloned and characterized PsneIFSA2 and PsneIFSA4 from young poplar (P. simonii × P. nigra) leaves. The deduced amino acid sequences of PsneIF5A2 and PsneIF5A4 were 98 % similar to each other, and they are orthologs of eIF5A 1 in Arabidopsis. In a subcellular localization analysis, PsneIF5A2 and PsneIF5A4 proteins were localized in the nucleus and cytoplasm, qRT-PCR analysis showed that PsneIF5A2 and PsneIF5A4 were transcribed in poplar flowers, stem, leaves, and roots. In addition, they were also induced by abiotic stresses. Transgenic yeast expressing PsneIF5A2 and PsneIF5A4 had increased salt, heavy metal, osmotic, oxidative tolerance. Our results suggest that PsneIF5A2 and PsneIF5A4 are excellent candidates for genetic engineering to improve salt and heavy metal tolerance in agricultural plants.
基金supported by National Natural Science Foundation of China (30871727 31071817)+1 种基金the Funding Project for Academic Human Resources Development in Institutions of Higher Learning under the Jurisdiction of Beijing Municipality (PXM20090142076309)the Beijing Science and Technology Plan Program (Z080005032508017)
文摘Aldehydes play an important role in atmospheric chemistry and plant direct and indirect defense against environmental stresses.In this study,the emissions of saturated C6-C10 aldehydes from Populus simonii × P.pyramidalis 'Opera 8277' cuttings were examined by using a gas chromatography/mass spectrometry(GC/MS) technique at three levels of light intensity(400,800 and 1 200 μmol·m-2·s-1).A positive correlation between the emissions of these aldehydes and light intensity was found.Moreover,nordi-hydroguaiaretic acid(NDGA),a special inhibitor of lipoxygenase(LOX),significantly inhibited the emissions of C6-C9 aldehydes at three levels of light intensity,but did not influence the emission of decanal(C10).The emissions of C6-C10 aldehydes in NDGA treated poplar cuttings,exhibited the same positive correlation with light intensity.The results indicated that LOX pathway contributes to the emissions of C6-C9 aldehydes,whereas some pathways regulated by light intensity might be a universal mechanism for emissions of C6-C10 aldehydes.
基金NSFC (30800865, 40801001)the Education Ministry of China (NCET-05-0886)China Postdoctoral Science Foundation (20070420758)
文摘Populus euphratica Oliv. is of high salinity tolerance and used as a model species for investigating molecular mechanisms of trees' responses to salt stress. In the work presented here we found that calli of P. euphratica grew more rapidly and accumulated less Na+, but more K+, under salt stress than those of salt-sensitive poplar, Populus hopeiensis. Different types of Na+/H+ antiporters (SOS1, NhaD1 and NHX1) were isolated from P. euphratica; all of these genes have been shown to play important roles in plant salt tolerance mechanism in previous studies. Expression profiles of these three genes were compared between P. hopeiensis and P. euphratica in the presence and absence of salt stress by real-time PCR. The three genes were induced in both P. euphratica and P. hopeiensis by salt. Transcript levels of PeNHX1 were lower in P. euphratica than in P. hopeiensis under 150 mM NaCl stress. In addition, transcript levels of PeNhaD1 were lower, while PeSOS1 were higher in P. euphratica than in P. hopeiensis under both stressed and unstressed conditions. The results indicated that P. euphratica up-regulates different genes and consistently maintains both effluxes of Na+ and high K+ levels. Our data suggests that differences in gene expression patterns may contribute to the dif-ference in salt tolerance between these two poplars.
基金Financial support for this research was provided by the Program for New Century Excellent Talents in the Ministry of Education in China(NCET-09-0446),NSFC(31370396,30800865) and lzujbky-2012-k22 to YuXia Wu
文摘Populus euphratica and Populuspruinosa, sister species in the Turanga Section (Salicaceae), growing in semi-arid saline areas are known for their high salinity tolerance. In this study, by combining growth level with Na+ and K+ contents, the expression level of vacuolar Na+/H+ antiporters was investigated for NaCl-induced changes in P. euphratica and t3. pru- inosa calli. Compared to R euphratica, P. pruinosa calli grew well in 200 mM NaC1 stress from 14. to 21 days. Increasing the stressed time caused an increase in Na+ content concomitant with a decrease in K+ content in P. euphratica calli, whereas, with the presence of 200 mM NaCI, K+ content has a less increase in 14 and 21 days than in 7 days which was detected in R pruinosa calli. The transcript levels of six genes coding for NHX-type Na+/H+ antiporters suggest that vacuolar NHX1-NHX6 antiporters play important roles in responding to salt stress in R pruinosa. Our data suggest that there exists a higher salt tolerance for P. pruinosa than P. euphratica at the cellular level, Na+ avoidance or accumulation is observed in cellular compartments, and that expression of NHX antiporters is linked to the accumulator phenotype.
基金Supported by Research and Development Program of Gansu Province(17YF1FA113)The National Natural Science Foundation of China(31560128).
文摘[Objectives]This study was conducted to investigate the photosynthetic physiological characteristics of Populus euphratica trees at different forest ages in the desert area of Minqin in response to drought stress.[Methods]With P.euphratica trees of different ages in Minqin as the research object,the water characteristics and photosynthetic physiological indexes(chlorophyll,soluble sugar,POD,SOD and MDA)were compared under different conditions.[Results]On the time gradient,the soil water contents of P.euphratica of different ages decreased continuously with the extension of irrigation stop time,but the decreases were small.Under the same stress treatment,the soil water contents of P.euphratica of different ages increased with the deepening of soil layer,but the differences were not significant(P>0.05).Under normal condition,the chlorophyll contents of P.euphratica at three ages gradually increased with the increase of age.The chlorophyll contents in leaves of P.euphratica at different ages were all lower under normal condition and higher under water stress.The changes of POD and SOD activity in leaves of P.euphratica at different ages under different conditions were basically the same,showing that the enzyme activity was higher under water stress than under normal under.The MDA contents in leaves of the young and middle P.euphratica forests were higher under water stress at 7 and 21 d than under normal condition,and the differences were not significant(P>0.05);and the MDA content in leaves of the mature P.euphratica forest was higher under water stress at 21 and 35 d than under normal condition.Drought stress has a certain effect on the photosynthetic physiological characteristics of P.euphratica.In summary,under drought stress,the chlorophyll content,SOD and POD activity,and MDA molar concentration in the leaves of P.euphratica were basically higher than under normal condition,indicating that P.euphratica could resist drought environment through osmotic adjustment and showed strong drought resistance.[Conclusions]This study provides a theoretical reference for the restoration,protection and reconstruction of natural P.euphratica forests in the desert area of Minqin.
基金supported by the Pro-gramme for Changjiang Scholars and the Innovative Research Team in Universities of China (PCSIRT0607)by the National Natural Science Foundation of China (30871727+2 种基金 30872037)the National Key Project of Scientific and Technical Supporting Programmes Funded by the Ministry of Science & Technology of China (2006BAD01A15 2006BAD24B04)
文摘Abstract The effects of water potential, NaCl and Na2SO4 on germination and radicle growth of two riparian tree species, Populus euphratica Oliv. and P. pruinosa Schrenk (Salicaceae), were tested. Growth chamber studies revealed an optimum temperature range for seed germination of both species between 15-35℃. The final germination percentage of both species decreases with decreasing water potential in all types of solution applied in the experiments. P. pruinosa was less tolerant to low ψw stress than P. euphratica, especially in salt solutions. Germination percentages fell below 20% for P. pruinosa at -0.6 MPa (NaCl) or -0.4 MPa (Na2SO4) and for P. euphratica at -1.2 MPa (NaCl) or -0.6 MPa (Na2SO4). Radicle growth of both species was inhibited by high concentrations of PEG1 NaCl and Na2SO4. However, growth was enhanced at -0.13 and -0.29 MPa in PEG or at -0.13 MPa in NaCl solutions compared to distilled water. Radicle growth of P. euphratica was higher than that of P pruinosa. Germination and radicle growth of both species exhibited ion toxicity. Na2SO4 was more toxic than iso-osmotic solutions of NaCl. Radicle growth proved to be more sensitive than seed germination. Thus, flooding does not only yield the necessary soil moisture for germination but also favors seedling establishment of both species through leaching of salts from the soil surface. The different sensitivity of the species during their early growth stages might, moreover, contribute to the observed differences in their distribution in the Talim Basin (northwest China).
基金Foundation for the Author of National Excellent Doctoral Dissertation of PRC (Grant No. 200152)the Teaching and Research Award Program for Outstanding Young Teachers in Higher Education Institution of MOE, PRC, the National Nat
文摘We used a salt-resistant poplar genotype Populus euphratica and two salt-sensitive genotypes, Populus 'popularis 35-44' (P. popularis) and the hybrid P. talassica Kom x (P. euphratica + Salix alba L.) to examine genotypic differences in nutrient selectivity under NaCl stress. One-year-old seedlings of P. euphratica and one-year-old hardwood cuttings of P. popularis were used in a short-term study (24 hours), while in a long-term study, up to 4 weeks, two-year-old seedlings of P. euphratica and the hybrid P. talassica Kom x (P. euphratica + Salix alba L.) were compared. In the short-term study, K+ concentration in the xylem sap ([K+]xylem) of P. euphratica significantly increased after salt stress was initiated, and maintained 1-2 fold higher than control levels during the period of salt stress (24 hours). Xylem Ca2+ and Mg2+ concentrations ([Ca2+]xylem, [Mg2+]xylem) in P. euphratica resembled the pattern of K+ despite a lesser magnitude in elevation. However, [K+]xylem, [Ca2+]xylem and [Mg2+]xylem in P. popularis exhibited a transient increase at the beginning of salt treatment, thereafter, they all returned to control levels at 4 hours and no further rise was observed in the following hours. Xylem Na+/K+, Na+/Ca2+ and Na+/Mg2+ in P. popularis increased sharply upon NaCl stress and steadily reached the maximum at 24 hours. In contrast, xylem Na+/K+, Na+/Ca2+ and Na+/Mg2+ in NaCl-treated plants of P. euphratica did not significantly increase during the period of salt stress (24 hours). Noteworthy, Na+/K+ markedly declined after the onset of stress. These results suggest that P. euphratica had a higher nutrient selectivity in face of salinity. A same trend was observed in a 4-week study. Xylem Na+/K+, Na+/Ca2+ and Na+/Mg2+ in salinised plants of the hybrid abruptly increased after 4 days of stress, and then continuously increased to reach the highest level at day 8 or day 15. In comparison, the magnitude of Na+/K+, Na+/Ca2+ and Na+/Mg2+ elevation in the xylem of P. euphratica was much lower during the observation period. In conclusion, salt-tolerant genotype P. euphratica maintained a higher nutrient selectivity under saline stress, as compared to the two salt-sensitive genotypes. The high capacity for nutrient uptake and transport presumably contributes to the salt tolerance of P. euphratica in a longer term.
基金funded by the National Natural Science Foundation of China (41373087, 30800887)the State Key Laboratory of Forest and Soil Ecology (LFSE2013-11)
文摘The aim of this study was to investigate the role of rhizosphere organic phosphorus (P) in soil P supply in semiarid forests and the effects of tree species on rihizosphere organic P. We examined organic P fractions in rhizosphere and bulk soils of mono-specific Simon poplar (Populus simonii) and Mongolian pine (Pinus sylvestris var. mongolica) plantations in a semiarid sandy soil of Horqin Sandy Land in Northeast China. Total organic P (TPo) accounted for 76% of total P across the two stands. The concentration of organic P (Po) fractions decreased in the order of NaOH-Po〉Res-Po〉HCI-Po〉NaHCO3-Po in both plantations. The concentration of NaHCO3-Po was 38% and 43% lower in rhizosphere soil than in bulk soil in Simon poplar and Mongolian pine plantations, respectively. In contrast, total P, TPo and NaOH-Po significantly accumulated in rhizosphere soil in Simon poplar plantations, but no change in Mongolian pine plantations. Soil recalcitrant organic P fractions were positively correlated with soil organic carbon. The results suggest that rhizosphere labile organic P was an important source of plant-available P in this semiarid region, but the dynamic of rhizosphere recalcitrant organic P fractions varied with tree species and was correlated to organic carbon dynamics.