植物生态学报 ›› 2012, Vol. 36 ›› Issue (2): 159-168.DOI: 10.3724/SP.J.1258.2012.00159 cstr: 32100.14.SP.J.1258.2012.00159
收稿日期:2011-09-21
接受日期:2011-11-15
出版日期:2012-09-21
发布日期:2012-02-22
作者简介:* E-mail: sunqingye@ahu.edu.cn
SHEN Zhang-Jun1,2, SUN Qing-Ye1,*(
), TIAN Sheng-Ni3
Received:2011-09-21
Accepted:2011-11-15
Online:2012-09-21
Published:2012-02-22
摘要:
白茅(Imperata cylindrica)为铜尾矿废弃地的自然定居优势植物。以农田正常生长的白茅居群为对照, 通过对不同生长时期的白茅体内氮(N)、磷(P)营养浓度, 叶片硝酸还原酶和酸性磷酸酶活性变化等的研究, 探讨铜尾矿胁迫下白茅在营养利用上的生理适应特征。研究结果表明, 在不同的生长时期, 白茅各器官内N、P分配存在一定的差异。在萌芽期, 两居群白茅体内的N、P均主要集中于根状茎。在花蕾期、成熟期时, 两居群白茅体内的N、P均向成熟叶片中迁移, 叶片中N、P浓度均达到最高。但在衰败期时, 铜尾矿居群白茅体内的N主要迁移到根状茎中, P在根状茎中的浓度也达到生长期中的最高值, 而农田居群白茅成熟叶片内N、P浓度依然最高。铜尾矿白茅叶片N、P的再吸收效率分别为49.54%-65.22%和74.71%-98.71%, 而在农田系统中分别为18.18%-52.81%和71.39%-84.07%; 铜尾矿白茅衰老叶片中P达到完全再吸收的程度。铜尾矿白茅叶片硝酸还原酶活性在生长旺盛期显著高于农田居群(p < 0.05), 是白茅加强对自身氮养分代谢活动调节作用的表现; 同一生长时期白茅叶片酸性磷酸酶活性在两种生境间差异性并不明显(p > 0.05), 但随着生长期的延长, 白茅叶片酸性磷酸酶活性表现出不断升高的趋势, 这有利于生长后期衰老叶片中有机P的水解再吸收。可见, 铜尾矿中生长的白茅通过对N、P养分的适时分配, 提高营养成分的再吸收效率, 调节N、P代谢相关调节酶活性的变化方式来减轻铜尾矿生境的营养胁迫。
沈章军, 孙庆业, 田胜尼. 铜尾矿自然定居白茅对体内氮磷的适时分配及叶片氮磷代谢调节酶活性动态. 植物生态学报, 2012, 36(2): 159-168. DOI: 10.3724/SP.J.1258.2012.00159
SHEN Zhang-Jun, SUN Qing-Ye, TIAN Sheng-Ni. Dynamics of nitrogen and phosphorus concentrations and nitrate reductase and acidic phosphatase activities in Imperata cylindrica on copper mine tailings. Chinese Journal of Plant Ecology, 2012, 36(2): 159-168. DOI: 10.3724/SP.J.1258.2012.00159
| 样地 Sample site | 有机质 Organic matter (g·kg-1) | 总氮 Total nitrogen (g·kg-1) | 硝态氮 Nitrate nitrogen (mg·kg-1) | 有效磷 Available phosphorus (mg·kg-1) | pH | 电导率 Electric conducti- vity (dS·m-1) |
|---|---|---|---|---|---|---|
| 铜尾矿 Copper tailings | 7.85 ± 0.14 | 0.41 ± 0.075 | 5.87 ± 1.49 | 4.25 ± 1.63 | 8.12 ± 0.32 | 1.03 ± 0.14 |
| 农田 Farmland | 13.75 ± 1.23 | 0.75 ± 0.091 | 13.67 ± 6.23 | 15.81 ± 8.42 | 6.89 ± 0.13 | 0.44 ± 0.32 |
表1 两种生境基本养分含量、pH以及电导率(平均值±标准偏差, n = 6)
Table 1 Contents of basic nutrients, pH value and electric conductivity in two habitat (mean ± SD, n = 6)
| 样地 Sample site | 有机质 Organic matter (g·kg-1) | 总氮 Total nitrogen (g·kg-1) | 硝态氮 Nitrate nitrogen (mg·kg-1) | 有效磷 Available phosphorus (mg·kg-1) | pH | 电导率 Electric conducti- vity (dS·m-1) |
|---|---|---|---|---|---|---|
| 铜尾矿 Copper tailings | 7.85 ± 0.14 | 0.41 ± 0.075 | 5.87 ± 1.49 | 4.25 ± 1.63 | 8.12 ± 0.32 | 1.03 ± 0.14 |
| 农田 Farmland | 13.75 ± 1.23 | 0.75 ± 0.091 | 13.67 ± 6.23 | 15.81 ± 8.42 | 6.89 ± 0.13 | 0.44 ± 0.32 |
| 样地 Sample site | 生长期 Growth period | 根 Root | 根状茎 Rhizome | 成熟叶 Mature leaf | 衰老叶 Senescent leaf |
|---|---|---|---|---|---|
| 铜尾矿 Copper tailings | 3月 March | 15.59 ± 0.91cd | 26.65 ± 2.40d | - | - |
| 4月 April | 14.51 ± 0.93c | 19.27 ± 2.89bc | 27.02 ± 3.64bc | - | |
| 6月 June | 9.71 ± 1.85a | 18.11 ± 0.68b | 20.25 ± 1.44a | 7.65 ± 0.65a | |
| 10月 October | 15.49 ± 0.77cd | 25.81 ± 7.84d | 19.88 ± 0.64a | 9.19 ± 0.80a | |
| 农田 Farmland | 3月 March | 16.29 ± 0.59d | 22.77 ± 0.72cd | - | - |
| 4月 April | 10.87 ± 0.84ab | 11.39 ± 0.58a | 29.59 ± 2.78c | - | |
| 6月 June | 11.71 ± 1.49b | 21.42 ± 1.36bc | 29.49 ± 1.25c | 16.10 ± 1.96b | |
| 10月 October | 15.49 ± 0.63cd | 21.61 ± 1.27bc | 25.11 ± 0.99b | 18.53 ± 1.79b |
表2 不同生长时期白茅不同器官总氮浓度分配(平均值±标准偏差, n = 6)
Table 2 Concentrations of total nitrogen in different organs of Imperata cylindrica (g·kg-1) (mean ± SD, n = 6)
| 样地 Sample site | 生长期 Growth period | 根 Root | 根状茎 Rhizome | 成熟叶 Mature leaf | 衰老叶 Senescent leaf |
|---|---|---|---|---|---|
| 铜尾矿 Copper tailings | 3月 March | 15.59 ± 0.91cd | 26.65 ± 2.40d | - | - |
| 4月 April | 14.51 ± 0.93c | 19.27 ± 2.89bc | 27.02 ± 3.64bc | - | |
| 6月 June | 9.71 ± 1.85a | 18.11 ± 0.68b | 20.25 ± 1.44a | 7.65 ± 0.65a | |
| 10月 October | 15.49 ± 0.77cd | 25.81 ± 7.84d | 19.88 ± 0.64a | 9.19 ± 0.80a | |
| 农田 Farmland | 3月 March | 16.29 ± 0.59d | 22.77 ± 0.72cd | - | - |
| 4月 April | 10.87 ± 0.84ab | 11.39 ± 0.58a | 29.59 ± 2.78c | - | |
| 6月 June | 11.71 ± 1.49b | 21.42 ± 1.36bc | 29.49 ± 1.25c | 16.10 ± 1.96b | |
| 10月 October | 15.49 ± 0.63cd | 21.61 ± 1.27bc | 25.11 ± 0.99b | 18.53 ± 1.79b |
| 样地 Sample site | 生长期 Growth period | 根 Root | 根状茎 Rhizome | 成熟叶 Mature leaf | 衰老叶 Senescent leaf |
|---|---|---|---|---|---|
| 铜尾矿 Copper tailings | 3月 March | 0.19 ± 0.05ab | 0.31 ± 0.03ab | - | - |
| 4月 April | 0.57 ± 0.19d | 0.27 ± 0.03ab | 1.07 ± 0.23a | - | |
| 6月 June | 0.27 ± 0.06bc | 0.35 ± 0.05b | 1.73 ± 0.57c | 0.15 ± 0.12a | |
| 10月 October | 0.34 ± 0.04c | 0.52 ± 0.11c | 0.73 ± 0.22ab | 0.05 ± 0.03a | |
| 农田 Farmland | 3月 March | 0.06 ± 0.02a | 0.36 ± 0.05bc | - | - |
| 4月 April | 0.16 ± 0.02ab | 0.15 ± 0.07a | 0.30 ± 0.03a | - | |
| 6月 June | 0.89 ± 0.08e | 1.57 ± 0.02d | 2.25 ± 0.14c | 0.39 ± 0.03b | |
| 10月 October | 0.61 ± 0.15d | 1.80 ± 0.22e | 2.09 ± 0.20c | 0.49 ± 0.06b |
表3 不同生长时期白茅不同器官总磷浓度(平均值±标准偏差, n = 6)
Table 3 Concentrations of total phosphorus in different organs of Imperata cylindrica (g·kg-1) (mean ± SD, n = 6)
| 样地 Sample site | 生长期 Growth period | 根 Root | 根状茎 Rhizome | 成熟叶 Mature leaf | 衰老叶 Senescent leaf |
|---|---|---|---|---|---|
| 铜尾矿 Copper tailings | 3月 March | 0.19 ± 0.05ab | 0.31 ± 0.03ab | - | - |
| 4月 April | 0.57 ± 0.19d | 0.27 ± 0.03ab | 1.07 ± 0.23a | - | |
| 6月 June | 0.27 ± 0.06bc | 0.35 ± 0.05b | 1.73 ± 0.57c | 0.15 ± 0.12a | |
| 10月 October | 0.34 ± 0.04c | 0.52 ± 0.11c | 0.73 ± 0.22ab | 0.05 ± 0.03a | |
| 农田 Farmland | 3月 March | 0.06 ± 0.02a | 0.36 ± 0.05bc | - | - |
| 4月 April | 0.16 ± 0.02ab | 0.15 ± 0.07a | 0.30 ± 0.03a | - | |
| 6月 June | 0.89 ± 0.08e | 1.57 ± 0.02d | 2.25 ± 0.14c | 0.39 ± 0.03b | |
| 10月 October | 0.61 ± 0.15d | 1.80 ± 0.22e | 2.09 ± 0.20c | 0.49 ± 0.06b |
| 样地 Sample site | 生长期 Growth period | 组织 Organs | N : P | REN | REP |
|---|---|---|---|---|---|
| 铜尾矿 Copper tailings | 4月 April | 成熟叶 Mature leaves | 26.73 ± 2.33ab | ||
| 6月 June | 成熟叶 Mature leaves | 12.96 ± 6.43a | 62.14 ± 3.08d | 88.17 ± 13.46ab | |
| 衰老叶 Senescent leaves | 68.56 ± 38.00bc | ||||
| 10月 October | 成熟叶 Mature leaves | 29.30 ± 7.96ab | 53.73 ± 4.19c | 91.36 ± 7.35b | |
| 衰老叶 Senescent leaves | 166.10 ± 50.43d | ||||
| 农田 Farmland | 4月 April | 成熟叶 Mature leaves | 97.14 ± 8.38c | ||
| 6月 June | 成熟叶 Mature leaves | 12.89 ± 0.55a | 45.28 ± 7.53b | 82.86 ± 1.21ab | |
| 衰老叶 Senescent leaves | 46.34 ± 3.58ab | ||||
| 10月 October | 成熟叶 Mature leaves | 12.21 ± 0.85a | 26.09 ± 7.91a | 76.08 ± 4.69a | |
| 衰老叶 Senescent leaves | 37.34 ± 6.41ab |
表4 不同生长时期白茅成熟叶片和衰老叶片内氮磷比和氮磷再吸收率(平均值±标准偏差, n = 6)
Table 4 N and P resorption efficiency, ratios of N:P in mature and senescent leaves of Imperata cylindrica during the different growth periods (mean ± SD, n = 6)
| 样地 Sample site | 生长期 Growth period | 组织 Organs | N : P | REN | REP |
|---|---|---|---|---|---|
| 铜尾矿 Copper tailings | 4月 April | 成熟叶 Mature leaves | 26.73 ± 2.33ab | ||
| 6月 June | 成熟叶 Mature leaves | 12.96 ± 6.43a | 62.14 ± 3.08d | 88.17 ± 13.46ab | |
| 衰老叶 Senescent leaves | 68.56 ± 38.00bc | ||||
| 10月 October | 成熟叶 Mature leaves | 29.30 ± 7.96ab | 53.73 ± 4.19c | 91.36 ± 7.35b | |
| 衰老叶 Senescent leaves | 166.10 ± 50.43d | ||||
| 农田 Farmland | 4月 April | 成熟叶 Mature leaves | 97.14 ± 8.38c | ||
| 6月 June | 成熟叶 Mature leaves | 12.89 ± 0.55a | 45.28 ± 7.53b | 82.86 ± 1.21ab | |
| 衰老叶 Senescent leaves | 46.34 ± 3.58ab | ||||
| 10月 October | 成熟叶 Mature leaves | 12.21 ± 0.85a | 26.09 ± 7.91a | 76.08 ± 4.69a | |
| 衰老叶 Senescent leaves | 37.34 ± 6.41ab |
图1 不同生长期白茅成熟叶片内硝酸还原酶活性(NRA) (A)和酸性磷酸酶活性(APA) (B)的动态。图中不同小写字母表示差异显著(p < 0.05)。
Fig. 1 Dynamics of nitrate reductase (NR) (A) and acid phosphatase (AP) (B) activity in the mature leaves of Imperata cylindrica during the different growth periods. Different small letters in the figure mean significant difference (p < 0.05).
| Y-X | 公式 Equation | n | r | p | Y-X | 公式 Equation | n | r | p |
|---|---|---|---|---|---|---|---|---|---|
| Nm-Pm | - | 12 | -0.361 | 0.352 | NRA-Nm | - | 12 | 0.342 | 0.732 |
| Nm-Ns | y = 19.186 + 0.104x | 12 | 0.109 | 0.009 | NRA-Pm | - | 12 | 0.692 | 0.069 |
| Pm-Ps | - | 12 | 0.386 | 0.067 | NRA-PEN | - | 12 | 0.683 | 0.389 |
| REN-Nm | - | 12 | 0.307 | 0.159 | NRA-MN︰P | y = 209.041 - 3.047x | 12 | 0.819 | 0.031 |
| REN-Ns | y = 94.473 - 4.356x | 12 | -0.819 | 0.002 | NRA-APA | y = 273.746 - 2.308x | 12 | -0.922 | 0.001 |
| REN-MN︰P | y = 64.190 - 0.303x | 12 | -0.620 | 0.000 | APA-Nm | - | 12 | -0.123 | 0.594 |
| REN-REP | y = 89.159 - 0.349x | 12 | -0.607 | 0.012 | APA-Pm | y = -22.547 + 84.522x | 12 | 0.827 | 0.001 |
| REP-Pm | y = 99.666 - 7.815x | 12 | -0.502 | 0.000 | APA-REP | v | 12 | 0.255 | 0.846 |
| REP-Ps | y = 84.283 + 54.819x | 12 | 0.531 | 0.000 | APA-MN︰P | v | 12 | 0.735 | 0.051 |
| REP-MN︰P | y = 87.882 + 0.089x | 12 | 0.105 | 0.001 |
表5 铜尾矿白茅叶片内氮磷浓度、氮磷再吸收率、氮磷比值、硝酸还原酶和酸性磷酸酶活性之间的相关性
Table 5 Correlation among N and P resorption efficiency, NRA, APA, nitrogen, phosphorus and N : P in the leaves of Imperata cylindrica in the tailings
| Y-X | 公式 Equation | n | r | p | Y-X | 公式 Equation | n | r | p |
|---|---|---|---|---|---|---|---|---|---|
| Nm-Pm | - | 12 | -0.361 | 0.352 | NRA-Nm | - | 12 | 0.342 | 0.732 |
| Nm-Ns | y = 19.186 + 0.104x | 12 | 0.109 | 0.009 | NRA-Pm | - | 12 | 0.692 | 0.069 |
| Pm-Ps | - | 12 | 0.386 | 0.067 | NRA-PEN | - | 12 | 0.683 | 0.389 |
| REN-Nm | - | 12 | 0.307 | 0.159 | NRA-MN︰P | y = 209.041 - 3.047x | 12 | 0.819 | 0.031 |
| REN-Ns | y = 94.473 - 4.356x | 12 | -0.819 | 0.002 | NRA-APA | y = 273.746 - 2.308x | 12 | -0.922 | 0.001 |
| REN-MN︰P | y = 64.190 - 0.303x | 12 | -0.620 | 0.000 | APA-Nm | - | 12 | -0.123 | 0.594 |
| REN-REP | y = 89.159 - 0.349x | 12 | -0.607 | 0.012 | APA-Pm | y = -22.547 + 84.522x | 12 | 0.827 | 0.001 |
| REP-Pm | y = 99.666 - 7.815x | 12 | -0.502 | 0.000 | APA-REP | v | 12 | 0.255 | 0.846 |
| REP-Ps | y = 84.283 + 54.819x | 12 | 0.531 | 0.000 | APA-MN︰P | v | 12 | 0.735 | 0.051 |
| REP-MN︰P | y = 87.882 + 0.089x | 12 | 0.105 | 0.001 |
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