刈割、施肥和浇水对矮嵩草补偿生长的影响

doi:10.3724/SP.J.1258.2011.00641

摘要/Abstract

摘要：

通过对青海海北高寒矮嵩草(Kobresia humilis)草甸进行为期3年的野外控制试验, 研究了刈割(留茬1 cm、3 cm及不刈割)、施肥(2.5 g·m^-2尿素+ 0.6 g·m^-2磷酸二胺、不施肥)和浇水(20.1 kg·m^-2、不浇水)处理对矮嵩草补偿生长(包括分株密度、株高和分株地上生物量)的影响, 及其比叶面积、叶片净光合速率和相对增长率的变化, 探讨矮嵩草补偿生长的机制。研究结果表明: 刈割后, 矮嵩草的补偿生长高度和比叶面积显著降低; 分株密度有增加的趋势, 但会随刈割强度的增加而下降; 株高和生物量的相对增长率随刈割强度的增加而呈上升趋势; 补偿地上生物量在重度刈割处理下最高。施肥能显著增加矮嵩草的补偿高度、分株密度、补偿地上生物量、株高相对增长率、生物量相对增长率、比叶面积和净光合速率; 与不浇水处理相比, 浇水处理对重度刈割处理下的分株地上生物量、密度相对增长率、比叶面积和净光合速率无影响, 而显著降低了中度刈割处理下的补偿高度和株高相对增长率, 提高了不刈割处理下的分株密度和重度刈割处理下的生物量相对增长率。刈割、施肥和浇水处理的交互作用也显示出刈割与施肥对矮嵩草补偿生长具有拮抗效应, 而刈割与浇水具有协同效应。上述结果说明, 矮嵩草在刈割后可通过增加分株密度和相对增长率等途径来提高补偿能力, 弥补在生长高度上出现的低补偿, 而施肥可显著抵消刈割的不利影响, 提高矮嵩草的补偿能力。

关键词: 刈割, 补偿性生长, 施肥, 矮嵩草, 浇水

Abstract:

Aims Kobresia humilis meadow is a disturbance-dependent (disclimax) community resulting from long-term and heavy grazing. Our objectives were to (a) examine how the compensatory growth of K. humilis (including ramet density, compensatory height and compensatory aboveground biomass per ramet) varies with different levels of clipping, fertilizing and watering and (b) examine changes in specific leaf area (SLA), lamina net photosynthetic rate (P_n) and relative growth rate (RGR) in order to reveal the compensatory mechanisms of the species.

Methods Our field experiment was carried out at the Haibei Research Station of the Chinese Academy of Sciences from 2007 to 2009. The experiment used a split-plot design with clipping treatment on the whole plot (stubbled 1 cm, 3 cm and unclipped) and both fertilizer (fertilized and unfertilized) and water (watered and unwatered) treatments in subplots. We analyzed effects of clipping, fertilizing and watering on the response variables by multivariate ANOVA. We used stepwise regression analysis to determine the contribution of SLA, P_n and RGR to compensatory growth and Pearson’s correlation to determine the correlation among ramet density, compensatory height and compensatory aboveground biomass.

Important findings Compensatory height and SLA were reduced after clipping; RGR of ramet height and aboveground biomass, ramet density and compensatory biomass were increased. Fertilizing can increase ramet density, compensatory height, compensatory biomass, RGR of ramet height and aboveground biomass, SLA and P_n. The effect of watering on compensatory height and RGR of ramet height under stubbled 1 cm clipping was not obvious, but reduced them under stubbled 3 cm clipping. The interactions of clipping, fertilizing and watering showed that the relation between clipping and fertilizing is “antagonistic” and the relation between clipping and watering is “cooperative”. Although K. humilis appeared under compensation in plant height, fertilizing could enhance the compensation ability and promote tolerance against defoliation at a certain level of clipping.

Key words: clipping, compensatory growth, fertilizing, Kobresia humilis, watering

张璐璐, 周晓松, 李英年, 袁芙蓉, 樊瑞俭, 朱志红. 刈割、施肥和浇水对矮嵩草补偿生长的影响. 植物生态学报, 2011, 35(6): 641-652. DOI: 10.3724/SP.J.1258.2011.00641

ZHANG Lu-Lu, ZHOU Xiao-Song, LI Ying-Nian, YUAN Fu-Rong, FAN Rui-Jian, ZHU Zhi-Hong. Effects of clipping, fertilizing and watering on compensatory growth of Kobresia humilis. Chinese Journal of Plant Ecology, 2011, 35(6): 641-652. DOI: 10.3724/SP.J.1258.2011.00641

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https://www.plant-ecology.com/CN/Y2011/V35/I6/641

图/表 6

表1 刈割、施肥和浇水在不同年份对矮嵩草补偿高度(GH)、补偿密度(GD)、株高相对增长率(RGRH)和密度相对增长率(RGRD)影响的方差分析

Table 1 ANOVA for the effects of clipping, fertilizing and watering on compensatory height (GH), compensatory density (GD), relative growth rate of height (RGRH) and relative growth rate of density (RGRD) of Kobresia humilis in different years

	变异来源 Source of variance	自由度 df (i, j)	G_H		G_D		RGR_H		RGR_D
	变异来源 Source of variance	自由度 df (i, j)	F-test	p	F-test	p	F-test	p	F-test	p
主区 Whole plot	Y C B Y × C	2, 16 2, 16 2, 16 4, 16	96.611 129.002 7.809 2.974	0.000^ 0.000^ 0.004^** 0.052	1.335 1.613 1.773 2.790	0.291 0.230 0.202 0.062	54.590 152.833 4.476 9.192	0.000^ 0.000^ 0.029^* 0.000^**	24.607 0.489 1.048 11.602	0.000^ 0.622 0.373 0.000^
副区 Subplot	F W Y × F Y × W C × F C × W F × W Y × C × F Y × F × W Y × C × W C × F × W Y × C × F × W	1, 54 1, 54 2, 54 2, 54 2, 54 2, 54 1, 54 4, 54 2, 54 4, 54 2, 54 4, 54	112.857 11.280 70.733 2.503 11.719 3.512 1.467 6.424 0.027 1.256 0.601 0.716	0.000^ 0.001^ 0.000^ 0.091 0.000^ 0.037^* 0.231 0.000^** 0.973 0.299 0.552 0.585	10.594 2.056 0.406 0.447 4.550 4.752 2.068 0.585 0.184 0.601 1.641 0.150	0.002^** 0.157 0.668 0.642 0.015^* 0.013^* 0.156 0.675 0.833 0.664 0.203 0.962	109.522 3.098 53.104 0.316 17.097 1.786 4.249 8.301 0.697 1.626 4.779 0.375	0.000^ 0.084 0.000^ 0.730 0.000^** 0.177 0.044^* 0.000^** 0.502 0.181 0.012^* 0.825	0.273 2.305 0.522 0.490 0.854 0.251 0.001 2.521 2.340 0.925 0.268 0.410	0.603 0.135 0.596 0.615 0.432 0.779 0.982 0.052 0.106 0.457 0.766 0.800
总变异 Total variance		107

图1 不同年份刈割、施肥和浇水处理对矮嵩草补偿高度、补偿密度和补偿地上生物量的影响(平均值±标准误差)。F, 施肥; H1, 重度刈割; H3, 中度刈割; NF, 不施肥; NH, 不刈割; NW, 不浇水; W, 浇水。相同字母及ns分别表示处理间或其互作效应无显著差异; 不同字母及*分别表示处理间或其互作效应差异显著(p < 0.05)。

Fig. 1 Effects of clipping, fertilizing and watering on compensatory height, compensatory density and compensatory aboveground biomass of Kobresia humilis in different years (mean ± SE). F, fertilized; H1, stubbled 1 cm; H3, stubbled 3 cm; NF, unfertilized; NH, unclipped; NW, unwatered; W, watered. The same letters and “ns” indicate no significant difference between treatments or their interaction, respectively; and different letters and “*” indicate significant differences between treatments or their interaction, respectively (p < 0.05).

表2 刈割、施肥和浇水对矮嵩草补偿地上生物量(GB)、生物量相对增长率(RGRB)、比叶面积(SLA)和净光合速率(Pn)影响的方差分析

Table 2 ANOVA for the effects of clipping, fertilizing and watering on compensatory aboveground biomass (GB), relative growth rate of biomass (RGRB), specific leaf area (SLA) and net photosynthetic rate (Pn) of Kobresia humilis

	变异来源 Source of variance	自由度 df (i, j)	SLA		P_n		G_B		RGR_B
	变异来源 Source of variance	自由度 df (i, j)	F-test	p	F-test	p	F-test	p	F-test	p
主区 Whole plot	C B	2, 4 2, 4	149.715 2.304	0.000^** 0.216	4.232 1.874	0.103 0.267	2.327 2.308	0.214 0.216	23.222 0.857	0.006^** 0.490
副区 Subplot	F W C × F C × W F × W C × F × W	1, 18 1, 18 2, 18 2, 18 1, 18 2, 18	11.644 0.709 0.834 0.292 0.177 0.499	0.003^** 0.411 0.450 0.750 0.679 0.616	20.358 0.583 0.358 0.436 1.418 0.643	0.000^** 0.455 0.704 0.653 0.249 0.537	20.546 0.827 0.781 1.570 0.009 3.450	0.000^** 0.375 0.473 0.235 0.924 0.054	31.969 0.734 3.101 2.697 1.747 7.958	0.000^ 0.403 0.070 0.095 0.203 0.003^
总变异 Total variance		35

表3 不同处理下矮嵩草分株高度、密度及地上生物量的补偿指数

Table 3 Compensation index of height, density and aboveground biomass of Kobresia humilis ramet under different treatments

补偿指数G/C Compensation index G/C	刈割处理 Clipping treatment				交互作用 Interactive effect
补偿指数G/C Compensation index G/C	NH	H3	H1		H3×NF× NW	H3×F× NW	H3×NF× W	H3×F× W	H1×NF× NW	H1×F× NW	H1×NF× W	H1×F× W
G_H/C G_D/C G_B/C	1 1 1	0.613 1.309 0.624	0.492 0.824 1.119		0.717 1.566 0.657	0.576 2.203 0.633	0.594 1.053 0.733	0.595 1.035 0.523	0.519 1.281 1.414	0.429 1.138 0.671	0.526 0.705 1.217	0.523 0.557 1.532

图2 不同年份刈割、施肥和浇水处理对矮嵩草相对增长率的影响(平均值±标准误差)。F, 施肥; H1, 重度刈割; H3, 中度刈割; NF, 不施肥; NH, 不刈割; NW, 不浇水; W, 浇水。相同字母及ns分别表示处理间或其互作效应无显著差异, 不同字母及*分别表示处理间或其互作效应差异显著(p < 0.05)。

Fig. 2 Effects of clipping, fertilizing and watering on relative growth rate of Kobresia humilis in different years (mean ± SE). F, fertilized; H1, stubbled 1 cm; H3, stubbled 3 cm; NF, unfertilized; NH, unclipped; NW, unwatered; W, watered. The same letters and “ns” indicate no significant difference between treatments and their interaction, respectively; and different letters and “*” indicate significant differences between treatments and their interaction, respectively (p < 0.05).

图3 刈割和施肥处理对矮嵩草比叶面积和净光合速率的影响(平均值±标准误差)。F, 施肥; H1, 重度刈割; H3, 中度刈割; NF, 不施肥; NH, 不刈割。相同字母及ns分别表示处理间或其互作效应无显著差异; 不同字母及*分别表示处理间或其互作效应差异显著(p < 0.05)。

Fig. 3 Effects of clipping and fertilizing on specific leaf area and net photosynthetic rate of Kobresia humilis (mean ± SE). F, fertilized; H1, stubbled 1 cm; H3, stubbled 3 cm; NF, unfertilized; NH, unclipped. The same letters and “ns” indicate no significant difference between treatments or their interaction respectively; and different letters and “*” indicate significant differences between treatments or their interaction, respectively (p < 0.05).

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