氮磷施肥对拟南芥叶片碳氮磷化学计量特征的影响

doi:10.3724/SP.J.1258.2013.00056

摘要/Abstract

摘要：

研究植物碳(C)氮(N)磷(P)化学计量特征, 有助于了解C、N、P元素的分配规律和确定限制植物生长的元素类型, 理解生长速率调控的内在机制。该研究基于盆栽施肥试验, 测定不同N、P供应水平下拟南芥(Arabidopsis thaliana)叶片的生物量和C、N、P含量, 分析拟南芥的限制元素类型、验证生长速率假说、探讨N、P的内稳性差异和C、N、P元素间的异速生长关系。主要结果如下: 盆栽试验基质中限制元素是P, 施N过多可能引起毒害作用; 拟南芥的生长符合生长速率假说, 即随着叶片N:P和C:P的增加, 比生长速率显著减小; 叶片P含量存在显著的调整系数(3.5), 但叶片N含量与基质N含量之间无显著相关; 叶片N和P含量具有显著的异速生长关系, 但不符合N-P^3/4关系, 施P肥导致表征N、P异速生长关系的幂指数(0.209)显著低于施N肥处理(0.466)。该研究首次基于温室培养实验分析了拟南芥C、N、P的化学计量特征及其对N、P添加的响应, 研究结果将为野外研究不同物种、群落或生态系统的化学计量特征提供参考。

关键词: 异速生长关系, 生长速率假说, 内稳性, 叶碳氮磷化学计量特征, 氮磷肥

Abstract:

Aims Arabidopsis thaliana, a widely used model organism in plant biology, is an ideal plant to test the growth rate hypothesis (GRH) and homeostasis theory about plant nutrition. Our objectives are to test i) whether GRH applies to this plant species, ii) how leaf nitrogen (N) and phosphorus (P) of A. thaliana follow the homeostasis theory and iii) whether the allometric relationship between leaf N and P content is consistent with the 3/4 power function (N-P^3/4) for individual plant species.
MethodsBased on a pot experiment in a phytotron with N and P fertilizer additions, we measured the leaf carbon (C), N and P content and leaf biomass of A. thaliana. Specific growth rate (mg·mg^-1·d^-1) was the leaf biomass increment divided by the initial biomass at planting, and by the days after planting. The homeostasis of plant elements is indicated by the exponent (reciprocal of the regulation coefficient) of the power function of leaf nutrient against soil nutrient concentrations.
Important findings P is the limiting nutrient of the culture substrate for A. thaliana, while N fertilization could cause toxic effects in cases of excessive N uptake. The growth of A. thaliana is consistent with GRH—the specific growth rate decreases with increasing leaf N:P or C:P. Leaf P content shows a significant regulation coefficient (3.51) (leaf-P-substrate-P^1/3.51), but leaf N content has no significant relationship with substrate N. There is a significant allometric relationship between leaf N and P content, which is inconsistent with the 3/4 power function (N-P^3/4). The power exponent (0.209) between leaf N content and leaf P content in the P fertilization treatments is significantly lower than the exponent (0.466) in the N fertilization treatments, suggesting that fertilization may affect the allometry between nutrients. Our findings can offer reference for future field studies on plant ecological stoichiometry at scales from species to community to ecosystem.

Key words: allometric relationship, growth rate hypothesis, homeostasis, leaf carbon/nitrogen/phosphorus stoichiometry, nitrogen and phosphorus fertilization

严正兵,金南瑛,韩廷申,方精云,韩文轩. 氮磷施肥对拟南芥叶片碳氮磷化学计量特征的影响. 植物生态学报, 2013, 37(6): 551-557. DOI: 10.3724/SP.J.1258.2013.00056

YAN Zheng-Bing,KIM Nam-Young,HAN Ting-Shen,FANG Jing-Yun,HAN Wen-Xuan. Effects of nitrogen and phosphorus fertilization on leaf carbon, nitrogen and phosphorus stoichiometry of Arabidopsis thaliana. Chinese Journal of Plant Ecology, 2013, 37(6): 551-557. DOI: 10.3724/SP.J.1258.2013.00056

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https://www.plant-ecology.com/CN/Y2013/V37/I6/551

图/表 6

表1 对照和各施肥水平下基质的N、P化学计量特征

Table 1 N and P stoichiometric characteristics in substrate of control (CK) and each fertilization treatment

基质N、P 水平 Substrate N, P level	基质有效氮含量^a Substrate available N^a content (mg·kg^-1)	基质有效磷含量^b Substrate available P^b content (mg·kg^-1)	基质N:P (质量比) Substrate N:P (mass ratio)
CK	180.73	12.12	14.91
N1	205.73	12.12	16.97
N2	230.73	12.12	19.03
N3	280.73	12.12	23.15
N4	380.73	12.12	31.40
N5	580.73	12.12	47.90
N6	780.73	12.12	64.39
P1	180.73	22.12	8.17
P2	180.73	32.12	5.63
P3	180.73	52.12	3.47
P4	180.73	92.12	1.96
P5	180.73	172.12	1.05
P6	180.73	252.12	0.72

图1 不同氮磷肥处理对拟南芥叶生长速率的影响(平均值±标准误差)。A, 氮肥处理。B, 磷肥处理。处理同表1。不同字母表示各基质氮(磷)水平下叶生长速率存在显著差异(p < 0.05)。

Fig. 1 Effect of different N (P) fertilization treatment on leaf growth rate (G) of Arabidopsis thaliana (mean ± SE). A, nitrogen fertilization treatment. B, phosphorus fertilization treatment. Treatment see Table 1. Different letters indicate significant difference (p < 0.05) in leaf growth rate between different substrate N (P) levels.

图2 基质有效P含量与拟南芥叶生长速率的关系。

Fig. 2 Relationship between substrate P content and leaf growth rate (G) of Arabidopsis thaliana.

图3 拟南芥叶N:P、C:P和叶比生长速率(u)的关系。αRMA表示简约主轴回归斜率。A, N:P-u。B, C:P-u。

Fig. 3 Relationship between Leaf N:P, C:P and leaf specific growth rate (u) of Arabidopsis thaliana. αRMA indicates the reduced major axis regression slope. A, N:P-u. B, C:P-u.

图4 拟南芥叶片N、P含量和基质有效N、P含量的关系。H, 调整系数。A, 叶N含量。B, 叶P含量。

Fig. 4 Relationship between leaf N (P) content of Arabidopsis thaliana and substrate available N (P) content. H, regulation coefficient. A , leaf N content. B, leaf P content.

表2 拟南芥叶片C、N和P含量的异速生长关系

Table 2 Allometric relationship among leaf C, N, P content of Arabidopsis thaliana

log Y vs. log X	处理 Treatment	α_RMA	95% CI	n	R²	p
logN vs. logC	施氮肥 N-fertilization	4.15	[2.67, 6.47]	21	0.091	0.183
	施磷肥 P-fertilization	-3.76	[-5.68, -2.49]	21	0.217	0.033
logP vs. logC	施氮肥 N-fertilization	8.92	[5.62, 14.15]	21	0.007	0.725
	施磷肥 P-fertilization	-17.98	[-22.28, -14.51]	21	0.798	<0.000 1
logN vs. logP	施氮肥 N-fertilization	0.466^a	[0.373, 0.581]	21	0.785	<0.000 1
	施磷肥 P-fertilization	0.209^b	[0.141, 0.311]	21	0.282	0.013

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