Effects of Pseudomonas TCd-1 inoculation on Cd uptake, rhizosphere soils enzyme activities and Cd bioavailability in rice (Oryza sativa) varieties with different Cd tolerance
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摘要: 为探究假单胞菌TCd-1降低水稻镉吸收的根际生态机制, 以高镉耐性水稻品种‘特优671’和低镉耐性水稻品种‘百香139’为材料, 通过盆栽土培试验, 研究了接种TCd-1菌株对10 mg∙kg−1镉处理水稻镉吸收、根际土壤镉形态及酶活性的影响。结果表明: 接种菌株后高、低镉耐性水稻品种各部位的镉含量显著降低(P<0.05), 镉富集系数分别降低35.14%和47.79%, 转移系数无显著变化; 根际土壤可交换态镉含量分别显著降低15.89%和23.81% (P<0.05) , 铁锰氧化结合态镉含量显著提高39.58%和28.81% (P<0.05), 有机态镉含量显著提高36.11%和25.00% (P<0.05); 低镉耐性水稻品种根际土壤酸性磷酸酶、脲酶、蔗糖酶、纤维素酶和过氧化氢酶活性依次提高26.74%、12.07%、62.50%、81.17%和5.13%, 多酚氧化酶活性降低12.40%, 高镉耐性水稻的酸性磷酸酶、脲酶、蔗糖酶、纤维素酶和多酚氧化酶活性依次降低7.19%、9.39%、25.53%、16.20%和11.44%, 过氧化氢酶活性提高5.13%。可见, 假单胞菌TCd-1主要通过降低根际土壤镉的生物有效性、恢复土壤酶活性, 进而提高水稻耐镉能力并抑制水稻对镉的吸收与积累, 对高、低不同镉耐性水稻品种, 接种TCd-1菌株后其镉富集特性、根际土壤酶活性及不同镉形态含量占比均表现出显著差异。Abstract: Heavy metal contamination in rice (Oryza sativa) is a serious problem. Microbial remediation is a promising technique to reduce Cd accumulation in rice. To explore the rhizosphere-associated ecological mechanism of Pseudomonas TCd-1-induced reduction of Cd uptake in rice, two rice varieties, high Cd-tolerant variety ‘Teyou 671’ and low Cd-tolerant variety ‘Baixiang 139’, were used. A set of soil culture pots treated with 10 mg∙kg−1 Cd were employed to evaluate the effects of Pseudomonas TCd-1 inoculation on rice Cd uptake and enzymes activities in rhizosphere soils. The results showed that the Cd content in different parts of both the high and low Cd-tolerant rice varieties significantly decreased (P<0.05) after inoculation of Pseudomonas TCd-1, and the bioconcentration factor (BCF) of Cd decreased by 35.14% and 47.79% (P<0.05), respectively. However, no significant changes were found in the translocation factor (TF). Meanwhile, in rhizosphere soils of the high and low Cd-tolerant rice varieties, the content of exchangeable Cd decreased by 15.89% and 23.81% (P<0.05), Fe-Mn oxide bound Cd increased by 39.58% and 28.81% (P<0.05), and organic matter Cd increased by 36.11% and 25.00% (P<0.05), respectively. In addition, the activities of acid phosphatase, urease, saccharase, cellulase, and catalase significantly increased by 26.74%, 12.07%, 62.50%, 81.17%, and 5.13%, respectively; while the polyphenol oxidase activity decreased by 12.40% in the rhizosphere soils of low Cd-tolerant rice variety. In rhizosphere soils of high Cd-tolerant rice variety, the activities of acid phosphatase, urease, sucrase, cellulase, and polyphenol oxidase decreased by 7.19%, 9.39%, 25.53%, 16.20%, and 11.44%, respectively; while catalase activity increased by 5.13%. There were significant differences in enrichment characteristics, rhizosphere soil enzymes activities, and the proportions of different chemical forms of Cd in rhizosphere soils of different Cd-tolerant rice varieties after inoculation with the TCd-1 strain. The results indicated that inoculation with the strain may partly remediate the changes in soil enzymes activities caused by Cd pollution. In conclusion, Pseudomonas TCd-1 can improve Cd tolerance and inhibit Cd uptake and accumulation in rice, mainly by reducing the bioavailability of soil Cd and restoring the changes in soil enzymes activities caused by Cd pollution.
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Key words:
- Rice /
- Cd pollution /
- Pseudomonas TCd-1 /
- Soil Cd form /
- Bioavailability /
- Soil enzyme activity /
- Microbial remediation
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图 1 高镉耐性(上三角矩阵)和低镉耐性(下三角矩阵)水稻接种假单胞菌TCd-1前(A)后(B)的镉含量、镉富集量、土壤酶活性及镉形态的相关性
Whole plant Cd: 全株镉含量; Root Cd: 根镉含量; Aerial part Cd: 地上部镉含量; Grain Cd: 籽粒镉含量; Root Cd accumulation: 根部镉富集量; Grain Cd accumulation: 籽粒镉富集量; Aerial part Cd accumulation: 地上部镉富集量; Whole plant Cd accumulation: 全株镉富集量; PPO: 多酚氧化酶活性; CAT: 过氧化氢酶活性; URE: 脲酶活性; ACP: 酸性磷酸酶活性; SUC: 纤维素酶活性; CUE: 蔗糖酶活性; EXC-Cd: 可交换态镉含量; Crab-Cd: 碳酸盐结合态镉含量; FeMnO2-Cd: 铁锰氧化物结合态镉含量; OM-Cd: 有机结合态镉含量; RES-Cd: 残渣态镉含量。蓝色表示负相关, 红色表示正相关, 颜色越深, 相关性越强。右边的数值代表Pearson相关性指数。
Figure 1. Correlation among Cd content, Cd accumulation, soil enzyme activity and soil Cd forms contents for high Cd-tolerant rice variety ‘Teyou 671’ (upper triangular matrix) and low Cd-tolerant rice variety ‘Baixiang 139’ (lower triangular matrix) before (A) and after (B) Pseudomonas TCd-1 inoculation
PPO: polyphenol oxidase activity; CAT: catalase activity; URE: urease activity; ACP: acid phosphatase activity; SUC: cellulase activity; CUE: invertase activity; EXC-Cd: exchangeable Cd content; Crab-Cd: carbonate bound Cd content; FeMnO2-Cd: Fe-Mn oxides bound Cd content; OM-Cd: organic matter Cd content; RES-Cd: residual Cd content. Blue indicates a negative correlation, red indicates a positive correlation, the darker the color, the stronger the correlation. The numbers on the right represent the Pearson index.
表 1 菌和Cd互作处理对不同镉耐性水稻各部位镉含量、富集系数(BCF)、转移系数(TF)及镉富集量的影响
Table 1. Effects of Pseudomonas TCd-1, Cd and their interaction on Cd contents, bioconcentration factor (BCF), translocation factor (TF) and Cd accumulation of different Cd-tolerant varieties of rice
处理
Treatment镉含量 Cadmium content (mg∙kg−1) 富集系数
BCF转移系数
TF镉富集量 Cadmium accumulation (μg∙plant−1) 根部
Root地上部
Aerial part籽粒
Grain全株
Whole plant根部
Root地上部
Aerial part籽粒
Grain全株
Whole plantTY-CK 0.28±0.04g 0.05±0.02f 0.03±0.00d 0.09±0.02e 0.52±0.10de 0.18±0.03d 4.05±0.31f 3.51±1.06f 0.99±0.24d 7.56±1.38f TY-CKB 0.33±0.03g 0.09±0.03f 0.06±0.01d 0.12±0.03e 0.67±0.18cde 0.26±0.08d 4.07±0.22f 6.99±2.58f 2.25±0.38d 11.06±2.88f TY-Cd 35.72±0.21a 3.47±0.21c 0.84±0.01b 7.53±0.42b 0.74±0.04cd 0.10±0.01e 342.60±13.33a 231.57±5.70b 28.94±2.62a 574.17±16.71a TY-CdB 24.83±0.10b 2.07±0.10d 0.60±0.03c 4.93±0.16c 0.48±0.02e 0.08±0.00e 227.06±5.92b 131.69±6.66c 20.37±1.46c 358.75±12.58b BX-CK 0.55±0.03f 1.50±0.06e 0.02±0.01d 1.39±0.05d 7.74±0.28b 2.75±0.08d 3.84±0.33f 81.81±3.74c 0.67±0.31d 85.64±4.01e BX-CKB 1.69±0.07e 1.46±0.02e 0.08±0.02d 1.49±0.02d 8.26±0.10a 0.86±0.04b 16.13±0.57e 100.47±5.83d 2.69±0.67d 116.60±6.40d BX-Cd 13.18±0.02c 8.47±0.43a 1.33±0.01a 9.07±0.40a 0.89±0.39c 0.64±0.03c 65.34±2.39c 289.05±8.20a 22.96±2.22b 354.38±10.56b BX-CdB 6.30±0.06d 4.52±0.25b 0.81±0.08b 4.74±0.23c 0.47±0.02e 0.72±0.03c 45.87±1.03d 235.69±1.55b 22.98±1.09b 281.56±2.33c 显著性 Significance 水稻品种
Rice variety (V)** ** ** ** ** ** ** ** NS ** 镉处理
Cadmium treatment (Cd)** ** ** ** ** ** ** ** ** ** 菌处理
Bacterial treatment (B)** ** ** ** NS ** ** ** * ** V×Cd ** ** ** ** ** ** ** NS NS ** V×B ** ** ** ** NS ** ** ** ** ** Cd×B ** ** ** ** ** ** ** ** ** ** V×Cd×B ** ** ** ** * ** ** ** ** ** 表中同列不同小写字母表示处理间在 P<0.05水平差异显著; *和**分别表示在 P<0.05和 P<0.01水平差异显著; NS表示无显著性差异。TY: 高镉耐性水稻品种‘特优671’; BX: 低镉耐性品种‘百香139’; CK: 不加镉且不接种菌株; CKB: 不加镉但接种菌株; Cd: 加镉但不接种菌株; CdB: 加镉且接种菌株。In the table, different lowercase letters in the same column mean significant differences among treatments at P<0.05. * and ** mean significant differences at P<0.05 and P<0.01, respectively. NS means no significant difference. TY: high Cd-tolerant rice variety ‘Teyou 671’; BX: low Cd-tolerant rice variety ‘Baixiang 139; CK: control treatment without Cd and Pseudomonas TCd-1; CKB: control treatment without Cd but with Pseudomonas TCd-1 inoculation; Cd: treatment of 10 mg·kg−1 Cd; CdB: treatment with inoculation of Pseudomonas TCd-1 and 10 mg·kg−1 Cd. 表 2 菌和Cd互作处理对不同镉耐性水稻品种根际土壤酶活性的影响
Table 2. Effect of Pseudomonas TCd-1, Cd and their interaction on enzymes activities in rhizosphere soil of different Cd-tolerant varieties of rice
处理
Treatment酸性磷酸酶
Acid phosphatase (ACP)
[mg∙g−1(FW)∙(24 h)−1]脲酶
Urease (URE)
[mg∙g−1(FW)∙
(24 h)−1]蔗糖酶
Sucrase (SUC)
[mg∙g−1(FW)∙
(24 h)−1]纤维素酶
Cellulase
(CUE)
[mg∙g−1(FW)∙(24 h)−1]过氧化氢酶
Catalase
(CAT)
[mL∙g−1(FW)∙(20 min−1]多酚氧化酶
Polyphenol oxidase (PPO)
[mg∙g−1(FW)∙
(24 h)−1]TY-CK 1.06±0.01d 1.47±0.02f 0.46±0.01a 13.02±0.86b 0.46±0.01a 16.16±1.00a TY-CKB 1.20±0.01c 1.49±0.00f 0.46±0.03a 14.09±0.11a 0.45±0.00a 17.07±0.30a TY-Cd 1.39±0.01a 1.81±0.02c 0.47±0.02a 14.44±0.34a 0.39±0.00c 16.08±0.48a TY-CdB 1.29±0.01b 1.64±0.00e 0.35±0.00c 12.10±0.50c 0.41±0.01b 14.24±0.77b BX-CK 0.98±0.01e 2.16±0.01a 0.29±0.01d 8.88±0.030d 0.36±0.00d 14.59±0.46b BX-CKB 0.96±0.01e 1.33±0.02g 0.29±0.00d 9.18±0.13d 0.39±0.01c 16.01±0.74a BX-Cd 0.86±0.00f 1.74±0.02d 0.24±0.01e 7.70±0.030e 0.39±0.00c 16.37±0.75a BX-CdB 1.09±0.12d 1.95±0.03b 0.39±0.02b 13.95±0.04a 0.41±0.01b 14.34±0.24b 显著性 Significance 水稻品种
Rice variety (V)** ** ** ** ** * 镉处理
Cd treatment (Cd)** ** NS ** ** * 菌处理
Bacterial treatment (B)** ** NS ** ** * V×Cd ** ** ** ** ** * V×B * ** ** ** ** NS Cd×B NS ** NS ** NS ** V×Cd×B ** ** ** ** ** NS 表中同列不同小写字母表示处理间在P<0.05水平差异显著; *和**分别表示在P<0.05和P<0.01水平差异显著; NS表示无显著性差异。TY: 高镉耐性水稻品种‘特优671’; BX: 低镉耐性品种‘百香139’; CK: 不加镉且不接种菌株; CKB: 不加镉但接种菌株; Cd: 加镉但不接种菌株; CdB: 加镉且接种菌株。In the table, different lowercase letters in the same column mean significant differences among treatments at P<0.05. * and ** mean significant differences at P<0.05 and P<0.01, respectively. NS means no significant difference. TY: high Cd-tolerant rice variety ‘Teyou 671’; BX: low Cd-tolerant rice variety ‘Baixiang 139; CK: control treatment without Cd and Pseudomonas TCd-1; CKB: control treatment without Cd but with Pseudomonas TCd-1 inoculation; Cd: treatment of 10 mg·kg−1 Cd; CdB: treatment with inoculation of Pseudomonas TCd-1 and 10 mg·kg−1 Cd. 表 3 菌和镉互作处理对不同镉耐性水稻根际土壤中不同形态镉含量的影响
Table 3. Effect of Pseudomonas TCd-1, Cd and their interaction on different forms of cadmium in rhizosphere soil of different Cd-tolerant varieties of rice
处理
Treatment可交换态镉
Exchangeable cadmium
(EXC-Cd)碳酸盐结合态镉
Carbonate bound cadmium
(Crab-Cd)铁锰氧化结合态镉
Fe-Mn oxides bound cadmium
(FeMnO2-Cd)有机态镉
Organic matter cadmium
(OM-Cd)残渣态镉
Residual cadmium
(RES-Cd)mg·kg−1 TY-CK 0.08±0.00e 0.04±0.00c 0.03±0.00d 0.01±0.01c 0.01±0.00c TY-CKB 0.06±0.01e 0.05±0.01c 0.03±0.00d 0.02±0.00c 0.01±0.00c TY-Cd 4.09±0.55a 2.05±0.30a 1.92±0.29b 0.72±0.12b 0.59±0.09b TY-CdB 3.44±0.28b 1.81±0.15a 2.68±0.20a 0.98±0.08a 0.70±0.06b BX-CK 0.04±0.01e 0.02±0.01c 0.02±0.00d 0.01±0.00c 0.07±0.01c BX-CKB 0.05±0.01e 0.02±0.01c 0.01±0.00d 0.01±0.00c 0.08±0.01c BX-Cd 2.73±0.11c 1.39±0.23b 0.59±0.03c 0.80±0.07b 3.94±0.53a BX-CdB 2.08±0.14d 1.85±0.11a 0.76±0.05c 1.00±0.08a 4.07±0.24a 显著性 Significance 水稻品种 Rice variety (V) ** * ** NS ** 镉处理 Cadmium treatment (Cd) ** ** ** ** ** 菌处理 Bacterial treatment (B) * NS ** ** NS V×Cd ** * ** NS ** V×B NS * * NS NS Cd×B * NS ** ** NS V×Cd×B NS * * NS NS 表中同列不同小写字母表示处理间在P<0.05水平差异显著; *和**分别表示在P<0.05和P<0.01水平差异显著; NS表示无显著性差异。TY: 高镉耐性水稻品种‘特优671’; BX: 低镉耐性品种‘百香139’; CK: 不加镉且不接种菌株; CKB: 不加镉但接种菌株; Cd: 加镉但不接种菌株; CdB: 加镉且接种菌株。In the table, different lowercase letters in the same column mean significant differences among treatments at P<0.05. * and ** mean significant differences at P<0.05 and P<0.01, respectively. NS means no significant difference. TY: high Cd-tolerant rice variety ‘Teyou 671’; BX: low Cd-tolerant rice variety ‘Baixiang 139; CK: control treatment without Cd and Pseudomonas TCd-1; CKB: control treatment without Cd but with Pseudomonas TCd-1 inoculation; Cd: treatment of 10 mg·kg−1 Cd; CdB: treatment with inoculation of Pseudomonas TCd-1 and 10 mg·kg−1 Cd. -
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