混合接菌及营养元素对杜鹃苗生理生化特性的影响

杨舒婷; 欧静; 李林盼; 何跃军

doi:10.19303/j.issn.1008-0384.2020.05.007

混合接菌及营养元素对杜鹃苗生理生化特性的影响

doi: 10.19303/j.issn.1008-0384.2020.05.007

贵州大学林学院，贵州　贵阳　550025

基金项目: 国家自然科学基金项目（31560223）；贵州省科技计划项目（黔科合支撑[2016]2522号）

详细信息

作者简介:
杨舒婷（1993−），女，硕士研究生，主要从事园林植物栽培方面研究（E-mail：1007964839@qq.com）

通讯作者:
欧静（1974−），女，博士，教授，主要从事园林植物栽培与应用研究（E-mail：coloroj@126.com）

中图分类号: S 685.21
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出版历程
- 收稿日期: 2019-12-21
- 修回日期: 2020-03-20
- 刊出日期: 2020-05-01

Effects of Fungal Addition and Fertilizer Usage on Physiology and Biochemistry of Rhododendron simsii Seedlings

College of Forestry, Guizhou University, Guiyang, Guizhou　550025, China

摘要

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。
- 菌根真菌 /
- 混合接菌 /
- 营养元素 /
- 杜鹃 /
- 生理生化特性
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.
- Mycorrhizal fungi /
- fungal addition /
- fertilizer /
- azalea (Rhododendron simsii) /
- physiological and biochemical characteristics

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图 1 主成分排序

Figure 1. Ranking of principal components

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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
肥料 Fertilizer	施肥水平 Fertilization level	处理组合 Treatment	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

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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.

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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
肥料 Fertilizer	施肥水平 Fertilization level	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

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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）
肥料 Fertilizer	施肥水平 Fertilization level	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

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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）
肥料 fertilizer	施肥水平 Fertilization level	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

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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）
肥料 Fertilizer	施肥水平 Fertilization level	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

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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）
肥料 Fertilizer	施肥水平 Fertilization level	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

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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.

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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

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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 2
Chl	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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