Effects of Fungal Addition and Fertilizer Usage on Physiology and Biochemistry of Rhododendron simsii Seedlings
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摘要:
目的 探索杜鹃苗对不同混合接菌组合及营养元素的生理响应机制。 方法 以2年生杜鹃(Rhododendron simsii)苗为研究对象,采用J1(杜鹃花类菌根真菌混合接菌)、J2(杜鹃花类菌根真菌和马尾松外生菌混合接菌)、J3(不接菌,对照)3种接菌组合与不同氮磷钾施肥水平,研究菌根真菌与氮磷钾肥对杜鹃苗生理生化特性的影响,测定杜鹃幼苗生物量、叶绿素含量(chl)、超氧化物歧化酶(SOD)活性、过氧化物酶(POD)活性、过氧化氢酶(CAT)活性、丙二醛(MDA)含量、可溶性蛋白(SP)含量、可溶性糖(SS)含量、生长素(IAA)含量、细胞分裂素(ZR)含量、赤霉素(GA3)和脱落酸(ABA)含量的变化情况,并采用主成分分析法对各指标进行筛选。 结果 混合接菌和施肥处理都能提高杜鹃叶片的叶绿素a含量、叶绿素b含量、叶绿素总含量、SOD活性、POD活性、CAT活性、SP含量、IAA含量、ZR含量和GA3含量,降低杜鹃幼苗的MDA含量,同时接菌处理较不接菌处理只需要施正常氮磷钾肥量的1/4~1/2水平即可最大程度增强杜鹃苗的生理水平,特别是接J2处理。菌根共生体可能是通过促进养分吸收,增强光合作用能力,刺激激素合成,降低膜脂过氧化程度,提高保护酶活性,增强杜鹃苗的抗逆能力,促进杜鹃苗生长。根据相关性分析可得,各指标均具有相关性。 结论 生理生化指标总体表现为接菌处理优于不接菌处理,J2处理施养分1/2水平(即施用尿素1.1 g·株−1、钙镁磷1 g·株−1、氯化钾0.7 g·株−1)对杜鹃苗生长及生理生化指标的影响最为明显,叶绿素含量(Chl)、超氧化物歧化酶活性(SOD)、可溶性蛋白(SP)、生长素(IAA)、赤霉素(GA3)可以作为反映杜鹃苗生长评价的主要指标。 Abstract:Objective Responses of azalea (Rhododendron simsii) seedlings cultivated under the applications of fungi and NPK fertilizer were studied. Method Various fungal additions and NPK fertilizations were applied in cultivating 2-year-old azalea seedlings. The fungal treatments included Rhododendron mycorrhizae (J1), mixture of R. mycorrhizae and exophytic Pinus massoniana (J2), and blank control (J3). The fertilization was varied on the rate of application. Physiology and biochemistry of the azalea seedlings grown under the treatments were determined. Seedling biomass, chlorophyll (Chl) content, and activities of superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT), as well as contents of malondialdehyde (MDA), soluble protein (SP), soluble sugar (SS), auxin (IAA), cytokinin (ZR), gibberellic acid (GA3), and abscisic acid (ABA) of the plants were measured and subjected to a principal component analysis. Results The fungal additions increased the contents of Chla, Chlb, total Chl, SP, IAA, ZR, and GA3 as well as the activities of SOD, POD, and CAT, but reduced the MDA content in the seedlings. The addition, especially J2 treatment, allowed the seedlings to performed well physiologically with merely 1/4 to 1/2 of the normally applied amount of NPK fertilizer. It was speculated that the mycorrhizal symbiosis as observed might be the result of the improved nutrient absorption, photosynthesis, hormone synthesis, protective enzyme activity, stress resistance, and growth of, as well as the reduced membrane lipid peroxidation in the plants. It appeared that all indicators selected for this study were correlated. Conclusion The overall physiological and biochemical indicators on the azalea seedlings cultivated with the fungal applications were better than those of control. In particular, the treatment of J2 not only reduced the NPK fertilization by half (i.e., urea 1.1 g ·plant−1, CaMgP 1 g·plant−1, and KCl 0.7 g KCl·pllant−1) but also elevated the levels of Chl, SOD, SP, IAA, and GA3 on the azalea seedlings. -
表 1 施肥具体方案
Table 1. Experimental design
肥料
Fertilizer施肥水平
Fertilization level处理组合
Treatment码值方案 Coding value 实施方案 Aetualizing N P K N/
(g·株−1)P2O5/
(g·株−1)K2O/
(g·株−1)N肥 N fertilizer 0水平 0 level N0P2K2 0 2 2 0.0 1.0 0.7 1/4水平 1/4 level N3P2K2 3 2 2 0.6 1.0 0.7 1/2水平 1/2 level N2P2K2 2 2 2 1.1 1.0 0.7 1水平 1 level N1P2K2 1 2 2 2.2 1.0 0.7 P肥 P fertilizer 0水平 0 level N2P0K2 2 0 2 1.1 0.0 0.7 1/4水平 1/4 level N2P3K2 2 3 2 1.1 0.5 0.7 1/2水平 1/2 level N2P2K2 2 2 2 1.1 1.0 0.7 1水平 1 level N2P1K2 2 1 2 1.1 1.9 0.7 K肥 K fertilizer 0水平 0 level N2P2K0 2 2 0 1.1 1.0 0.0 1/4水平 1/4 level N2P2K3 2 2 3 1.1 1.0 0.4 1/2水平 1/2 level N2P2K2 2 2 2 1.1 1.0 0.7 1水平 1 level N2P2K1 2 2 1 1.1 1.0 1.4 表 2 不同接菌处理杜鹃菌根真菌侵染率比较
Table 2. Infection rates of mycorrhizal fungi on azalea seedlings under varied treatments
不同接菌处理
Different inoculation treatments菌根侵染率
Mycorrhizal infection rate/%J1处理 J1 treatment 58.18±1.579 b J2处理 J2 treatment 62.25±1.675 a J3处理 J3 treatment 8.85±0.696 c 注:表中数据为平均值±标准误差,数值后不同字母表示差异显著(P<0.05,Duncan`s检测),表3~7同。
Note: The data in the table are mean ± standard error,different letters after the value indicate significant difference(P<0.05, Duncan’s test), the same as Table 3–7.表 3 混合接菌及营养元素对杜鹃苗生物量的影响
Table 3. Effect of fungal addition and fertilizer usage on biomass of azalea seedlings
肥料
Fertilizer施肥水平
Fertilization level茎粗增幅 Stem thickness increase/cm 根冠增幅 Root crown increase/cm 鲜重 Fresh weight/g J1 J2 J3 J1 J2 J3 J1 J2 J3 N肥 N fertilizer 0水平 0 level 0.14 c 0.16 c 0.09 b 10.63 d 10.90 d 10.60 d 17.67 d 22.30 c 13.17 d 1/4水平 1/4 level 0.24 b 0.31 b 0.19 a 10.93 c 11.27 c 10.73 c 31.53 c 46.17 b 22.27 c 1/2水平 1/2 level 0.28 a 0.34 a 0.20 a 11.27 b 11.87 b 11.33 b 38.23 a 55.63 a 25.53 b 1水平 1 level 0.28 a 0.32 ab 0.20 a 11.63 a 12.07 a 11.53 a 35.07 b 58.33 a 28.73 a P肥 P fertilizer 0水平 0 level 0.15 d 0.19 b 0.09 b 10.70 d 10.93 c 10.77 c 19.47 c 25.47 c 15.00 d 1/4水平 1/4 level 0.21 c 0.23 b 0.19 a 11.13 c 11.47 b 10.93 b 23.30 b 37.27 b 21.40 c 1/2水平 1/2 level 0.28 a 0.34 a 0.20 a 11.27 b 11.87 a 11.33 a 38.23 a 55.63 a 25.53 b 1水平 1 level 0.25 b 0.30 a 0.18 a 11.57 a 11.77 a 11.40 a 40.30 a 57.57 a 29.57 a K肥 K fertilizer 0水平 0 level 0.17 c 0.18 c 0.12 c 10.77 c 11.03 c 10.83 c 20.30 c 28.07 c 16.27 d 1/4水平 1/4 level 0.22 b 0.24 b 0.15 b 11.37 ab 11.50 b 11.03 b 24.23 b 40.77 b 20.10 c 1/2水平 1/2 level 0.28 a 0.34 a 0.20 a 11.27 b 11.87 a 11.33 a 38.23 a 55.63 a 25.53 b 1水平 1 level 0.17 c 0.21 b 0.14 bc 11.53 a 11.60 b 11.37 a 37.70 a 56.17 a 29.60 a 表 4 混合接菌及营养元素对杜鹃苗叶绿素含量的影响
Table 4. Effect of fungal addition and fertilizer usage on Chl content of azalea seedlings
肥料
Fertilizer施肥水平
Fertilization level叶绿素a含量
Chlorophyll a content/(mg·g−1)叶绿素b含量
Chlorophyll b content/(mg·g−1)叶绿素总含量
Total chlorophyll content/(mg·g−1)J1 J2 J3 J1 J2 J3 J1 J2 J3 N肥 N fertilizer 0水平 0 level 2.06 b 2.45 b 1.74 b 1.05 c 1.32 b 0.83 b 3.36 b 4.14 c 2.84 b 1/4水平 1/4 level 2.22 b 2.56 b 1.81 b 1.12 c 1.49 b 0.75 b 3.66 b 4.37 c 3.00 b 1/2水平 1/2 level 2.58 a 2.66 ab 2.03 b 1.50 b 2.66 a 0.96 b 4.43 a 5.29 a 3.34 b 1水平 1 level 2.68 a 2.83 a 2.70 a 1.92 a 1.50 b 2.01 a 4.85 a 4.78 b 4.96 a P肥 P fertilizer 0水平 0 level 2.10 d 2.54 c 1.83 b 0.97 d 1.41 d 0.74 b 3.44 c 4.31 d 2.91 b 1/4水平 1/4 level 2.26 c 2.59 b 1.89 b 1.14 c 2.12 b 0.77 b 3.72 c 4.95 b 3.09 b 1/2水平 1/2 level 2.58 a 2.66 a 2.03 ab 1.50 a 2.66 a 0.96 ab 4.43 a 5.29 a 3.34 ab 1水平 1 level 2.51 b 2.65 a 2.32 a 1.45 b 1.70 c 1.12 a 4.26 b 4.63 c 3.85 a K肥 K fertilizer 0水平 0 level 2.39 c 2.62 c 1.88 a 1.35 b 1.67 c 0.82 a 4.07 d 4.56 c 3.09 a 1/4水平 1/4 level 2.48 b 2.64 b 1.88 a 1.29 c 1.59 c 0.96 a 4.14 c 4.50 c 3.13 a 1/2水平 1/2 level 2.58 a 2.66 a 2.03 a 1.50 a 2.66 a 0.96 a 4.43 a 5.29 a 3.34 a 1水平 1 level 2.49 b 2.64 b 2.14 a 1.49 a 2.34 b 0.97 a 4.30 b 5.07 b 3.56 a 表 5 混合接菌及营养元素对杜鹃苗酶活性的影响
Table 5. Effect of fungal addition and fertilizer usage on enzyme activities of azalea seedlings
肥料
fertilizer施肥水平
Fertilization level超氧化物歧化酶SOD/(U·g−1) 过氧化物酶POD/(U·g−1) 过氧化氢酶CAT/(U·g−1) J1 J2 J3 J1 J2 J3 J1 J2 J3 N肥 N fertilizer 0水平 0 level 430.44 b 539.28 d 419.53 b 143.53 c 288.03 c 106.41 c 37.29 c 57.63 c 33.43 c 1/4水平 1/4 level 432.75 b 886.84 a 443.44 a 326.36 b 786.14 a 220.10 b 50.85 bc 88.14 a 39.93 bc 1/2水平 1/2 level 479.48 a 755.51 b 444.04 a 500.33 ab 642.99 b 252.82 b 77.97 a 84.75 ab 50.85 ab 1水平 1 level 478.27 a 661.18 c 451.33 a 569.39 a 605.25 b 319.85 a 64.41 ab 68.34 bc 61.13 a P肥 P fertilizer 0水平 0 level 431.14 c 602.05 d 420.52 b 144.26 b 321.38 b 181.89 b 47.46 a 61.16 b 30.51 c 1/4水平 1/4 level 448.18 bc 819.72 a 437.91 ab 358.85 a 748.97 a 222.05 ab 62.03 a 85.22 a 37.29 bc 1/2水平 1/2 level 479.48 a 755.51 b 444.04 a 500.33 a 642.99 a 252.82 ab 77.97 a 84.75 a 50.85 ab 1水平 1 level 460.07 ab 657.18 c 448.58 a 464.16 a 537.70 ab 285.59 a 74.58 a 67.05 ab 61.77 a K肥 K fertilizer 0水平 0 level 438.66 b 629.36 d 423.88 b 213.19 a 465.93 a 177.40 a 47.68 b 61.62 b 31.02 c 1/4水平 1/4 level 439.52 b 801.15 a 438.75 a 366.60 a 680.13 a 250.19 a 57.63 ab 91.53 a 40.69 bc 1/2水平 1/2 level 479.48 a 755.51 b 444.04 a 500.33 a 642.99 a 252.82 a 77.97 a 84.75 ab 50.85 ab 1水平 1 level 451.39 b 647.38 c 447.13 a 429.00 a 500.88 a 254.81 a 54.24 b 61.02 b 57.63 a 表 6 混合接菌及营养元素对杜鹃苗MDA及渗透调节物质的影响
Table 6. Effects of fungal addition and fertilizer usage on MDA and osmolytes of azalea seedlings
肥料
Fertilizer施肥水平
Fertilization level丙二醛含量 MDA content/(nmol·g−1) 可溶性蛋白含量 Soluble protein content/(mg·g−1) 可溶性糖含量 Soluble sugar content/(mg·g−1) J1 J2 J3 J1 J2 J3 J1 J2 J3 N肥 N fertilizer 0水平 0 level 0.11 b 0.11 a 0.27 a 2.61 b 2.50 c 2.44 b 1.88 b 2.03 a 2.42 c 1/4水平 1/4 level 0.07 d 0.07 d 0.14 b 2.63 b 2.68 b 2.40 b 1.40 d 1.91 c 2.38 d 1/2水平 1/2 level 0.07 c 0.07 c 0.14 c 2.76 a 2.89 a 2.42 b 1.65 c 1.95 bc 2.47 b 1水平 1 level 0.12 a 0.11 b 0.14 c 2.53 c 2.67 b 2.55 a 2.31 a 1.99 ab 2.51 a P肥 P fertilizer 0水平 0 level 0.10 c 0.09 c 0.28 a 2.79 a 2.69 b 2.38 b 1.94 c 2.02 b 2.22 d 1/4水平 1/4 level 0.12 b 0.11 b 0.15 b 2.60 b 2.57 c 2.43 b 2.02 b 2.03 b 2.32 c 1/2水平 1/2 level 0.07 d 0.07 d 0.14 c 2.76 a 2.89 a 2.42 b 1.65 d 1.95 c 2.47 b 1水平 1 level 0.19 a 0.13 a 0.10 d 2.55 b 2.73 b 2.50 a 2.15 a 2.11 a 2.51 a K肥 K fertilizer 0水平 0 level 0.18 a 0.10 a 0.27 a 2.58 b 2.61 c 2.40 b 2.00 b 2.01 b 2.20 c 1/4水平 1/4 level 0.11 b 0.09 b 0.15 b 2.63 b 2.68 b 2.45 b 2.38 a 2.04 b 2.22 c 1/2水平 1/2 level 0.07 d 0.07 d 0.14 c 2.76 a 2.89 a 2.42 b 1.65 c 1.95 c 2.47 b 1水平 1 level 0.08 c 0.09 c 0.14 c 2.58 b 2.71 b 2.57 a 1.97 b 2.28 a 2.79 a 表 7 混合接菌及营养元素对杜鹃苗内源激素的影响
Table 7. Effect of fungal addition and fertilizer usage on endogenous hormones of azalea seedlings
肥料
Fertilizer施肥水平
Fertilization level细胞分裂素ZR/(ng·g−1) 生长素IAA/(ng·g−1) 赤霉素GA3/(ng·g−1) 脱落酸ABA/(ng·g−1) J1 J2 J3 J1 J2 J3 J1 J2 J3 J1 J2 J3 N肥 N fertilizer 0水平 0 level 8.47 b 9.38 c 7.23 a 48.63 a 53.17 c 41.34 b 6.43 c 7.15 c 5.74 b 100.64 a 101.03 a 79.24 a 1/4水平 1/4 level 9.09 b 11.66 a 7.75 a 50.58 a 79.30 a 45.49 ab 7.73 b 9.32 a 5.93 b 62.53 c 75.72 b 69.86 ab 1/2水平 1/2 level 10.36 a 10.62 ab 7.78 a 55.06 a 68.91 b 49.55 a 9.13 a 8.36 b 6.87 a 63.89 c 78.33 b 68.19 ab 1水平 1 level 9.25 b 10.26 bc 7.83 a 55.07 a 58.64 c 51.56 a 7.39 b 8.04 b 7.14 a 81.58 b 96.71 a 63.66 b P肥 P fertilizer 0水平 0 level 8.62 b 9.30 b 7.59 a 49.35 a 53.23 c 46.20 a 5.78 c 6.86 c 5.75 c 92.32 a 102.32 a 81.16 a 1/4水平 1/4 level 9.06 b 10.80 a 7.62 a 51.46 a 85.38 a 46.98 a 7.83 b 9.18 a 5.99 bc 70.44 c 71.50 b 71.72 ab 1/2水平 1/2 level 10.36 a 10.62 a 7.78 a 55.06 a 68.91 b 49.55 a 9.13 a 8.36 b 6.87 ab 63.89 c 78.33 b 68.19 b 1水平 1 level 9.38 b 10.33 ab 8.56 a 53.46 a 57.67 Bc 49.65 a 7.36 b 8.09 b 7.26 a 81.12 b 90.94 a 65.89 b K肥 K fertilizer 0水平 0 level 7.75 c 9.14 b 6.98 b 50.16 a 54.23 b 47.48 a 6.07 c 6.98 c 5.41 b 99.27 a 108.06 a 82.53 a 1/4水平 1/4 level 9.04 b 10.69 a 7.50 b 52.81 a 63.39 a 47.78 a 6.75 bc 7.84 ab 5.89 b 73.97 bc 84.91 b 73.56 ab 1/2水平 1/2 level 10.36 a 10.62 a 7.78 b 55.06 a 68.91 a 49.55 a 9.13 a 8.36 a 6.87 a 63.89 c 78.33 b 68.19 b 1水平 1 level 10.20 a 10.36 a 9.55 a 52.58 a 54.64 b 49.10 a 7.11 b 7.32 bc 6.66 a 78.22 b 87.01 b 66.83 b 表 8 杜鹃苗各生理指标相关性分析
Table 8. Correlation Analysis of physiological indexes of Azalea
指标 index Chl SOD POD CAT MDA SP SS IAA ZR GA3 ABA Chl 1.000 SOD 0.680** 1.000 POD 0.826** 0.847** 1.000 CAT 0.821** 0.779** 0.912** 1.000 MDA −0.430* −0.383* −0.437* −0.549** 1.000 SP 0.717** 0.597** 0.649** 0.722** −0.464** 1.000 SS −0.033 −0.122 0.003 −0.020 0.000 −0.187 1.000 IAA 0.735** 0.899** 0.874** 0.825** −0.409* 0.614** −0.146 1.000 ZR 0.778** 0.784** 0.881** 0.889** −0.525** 0.774** −0.129 0.801** 1.000 GA3 0.767** 0.718** 0.886** 0.908** −0.483** 0.679** −0.137 0.820** 0.884** 1.000 ABA −0.029 0.072 −0.230 −0.199 0.034 −0.076 −0.592** −0.095 −0.083 −0.276 1.000 注:*:P<0.05;**:P<0.01;Chl:叶绿素;MDA:丙二醛;SOD:超氧化物歧化酶;POD:过氧化物敏;CAT:过氧化氢酶;SP:可溶性蛋白;SS:可溶性糖;IAA:生长素;ZR:细胞分裂素;GA3:赤霉素:ABA:脱落酸。表10、图1同。
Note: *: P<0.05; * *: P<0.01; Chl: chlorophyll; MDA: malondialdehyde; SOD: superoxide dismutase; POD: peroxidase; CAT: catalase; SP: soluble protein; SS: soluble sugar; IAA: auxin; ZR: cytokinin; GA3: gibberellin: ABA: abscisic acid. The same as Table 10 and Fig.1.表 9 成分特征值分布及累积分析
Table 9. Distribution and accumulation of components
成分
Composition特征值
The eigenvalue方差
Variance/%累积
Cumulative/%1 6.853 62.299 62.299 2 1.648 14.981 77.280 3 0.771 7.010 84.290 4 0.562 5.111 89.401 5 0.449 4.085 93.486 6 0.286 2.599 96.085 7 0.180 1.635 97.720 8 0.088 0.803 98.523 9 0.080 0.726 99.248 10 0.055 0.500 99.748 11 0.028 0.252 100.000 表 10 主成分得分系数分析
Table 10. Coefficients of scores on principal components
指标
Index主成分1
Principal component 1主成分2
Principal component 2Chl 0.127 0.009 SOD 0.127 0.088 POD 0.137 −0.073 CAT 0.139 −0.058 MDA −0.083 0.013 SP 0.117 0.058 SS −0.022 −0.534 IAA 0.132 0.032 ZR 0.138 0.025 GA3 0.135 −0.049 ABA −0.013 0.545 -
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