磷元素和土壤酸化交互作用对核桃幼苗光合特性的影响

doi:10.3724/SP.J.1258.2014.00129

摘要/Abstract

摘要：

核桃(Juglans regia)向南推广种植不可避免地会遇到土壤酸化和缺磷的环境, 这种环境如何影响核桃的生长是生产中需要知晓的基础问题。该文研究了土壤不同pH值对核桃的磷素营养影响以及缺磷对核桃幼苗水分平衡、光合特性和生长的影响。在温室内采用砂培盆栽试验, 研究一年生核桃嫁接幼苗在不同pH值、磷水平基质中的水分关系、光合特性和生长的应对机制。研究设4种处理, 即: 对照(正常供应磷素+ pH 6.0); 正常供应磷素+ pH 3.0; 不添加磷素+ pH 6.0; 不添加磷素+ pH 3.0。结果显示: pH值与磷素对核桃幼苗的影响是两个相互独立的过程, 酸性(pH值3.0)条件下, 核桃幼苗根系生物量降低、根冠比减小, 根系导水率降低, 对磷素的吸收利用减少, 尽管其供磷正常, 但各生长指标及生理指标与磷胁迫条件下反应相似; 两因素具有一定的叠加性, 在磷胁迫条件下, 酸化(pH值3.0)对核桃幼苗的损害进一步加剧。各指标具体变化如下: 酸化及磷胁迫条件下核桃根系水分导度降低, 叶柄木质部结构改变, 导管密度降低, 木质部导管栓塞程度增加, 叶柄导水率下降, 植株水分运输效率降低, 叶片水势降低, 诱导气孔关闭; 气孔导度降低, 光合作用能力下降; 胁迫条件下, 叶绿素荧光参数最大光化学效率低于0.8, 实际光化学效率、光化学淬灭下降, 非光化学淬灭增加, 核桃幼苗受胁迫环境损害, 叶片光系统II光合电子传递活性受到抑制, 光合能力下降。总之, 土壤酸化抑制了核桃幼苗对磷元素的吸收利用, 造成体内缺磷; 磷胁迫及酸化抑制了叶柄木质部的发育, 降低了根系水分导度和叶柄导水率, 干扰了核桃幼苗水分平衡, 通过气孔与非气孔共同调节, 限制了核桃幼苗光合作用, 抑制了核桃幼苗高生长、直径生长及叶面积增加; 但并没有发现土壤酸化和缺磷之间有明显的交互作用。

关键词: 生长, 水力学特性, 核桃, pH值, 磷素营养, 光合特性

Abstract:

AimsOur main purposes were to explore effects of phosphorus deficiency and soil acidification on growth, water balance and phosphorus nutrition in Juglans regia seedlings and to investigate the hydraulic mechanism involved in reduced photosynthesis and growth.

Methods We measured growth, hydraulic characteristics, and gas exchanges in J. regia seedlings. The effects of phosphorus deficiency and soil acidification on xylem anatomic structure, water balance and photosynthetic characteristics were also analyzed.

Important findings Soil acidification inhibited the phosphorous uptake and utilization efficiency in J. regia seedlings. A combination of soil acidification and phosphorous deficiency interrupted water balance, and reduced photosynthesis and growth of J. regia seedlings. Under soil acidification and phosphorus deficiency, non-stomatal factors also played a role in inhibiting photosynthesis.

Key words: growth, hydraulic conductance, Juglans regia, pH value, phosphorus nutrition, photosynthetic characteristics

张翠萍, 孟平, 李建中, 万贤崇. 磷元素和土壤酸化交互作用对核桃幼苗光合特性的影响. 植物生态学报, 2014, 38(12): 1345-1355. DOI: 10.3724/SP.J.1258.2014.00129

ZHANG Cui-Ping, MENG Ping, LI Jian-Zhong, WAN Xian-Chong. Interactive effects of soil acidification and phosphorus deficiency on photosynthetic characteristics and growth in Juglans regia seedlings. Chinese Journal of Plant Ecology, 2014, 38(12): 1345-1355. DOI: 10.3724/SP.J.1258.2014.00129

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2014.00129

https://www.plant-ecology.com/CN/Y2014/V38/I12/1345

图/表 16

表1 磷处理营养液成分

Table 1 Composition of nutrient solution under different phosphorus treatments (mmol·L-1)

营养元素 Nutrition	缺磷 Phosphorus deficiency	正常磷 Normal phosphorus
KNO₃	4	2
K₂HPO₄	-	1
NH₄NO₃	1	2
(NH₄)₂SO₄	3	3
K₂SO₄	-	-
MgSO₄	2	2
KH₂PO₄	-	-
Ca(NO₃)₂	2	2

图1 土壤磷含量(平均值±标准误差)。 CK, 正常供磷素+ pH 6.0; T1, 正常供磷素+ pH 3.0; T2, 不添加磷素+ pH 6.0; T3, 不添加磷素+ pH 3.0。不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 1 Phosphorus content in soil (mean ± SE). CK, normal phosphorus and pH 6.0; T1, normal phosphorus and pH 3.0; T2, phosphorus deficiency and pH 6.0; T3, phosphorus deficiency and pH 3.0. Different small letters indicate significant differences among treatments (p < 0.05).

表2 磷元素与pH值对土壤及核桃幼苗各器官磷含量影响的双因素分析

Table 2 Two-way ANOVA for testing the effects of phosphorus and pH value on phosphorus content in soil and seedlings

	土壤 Soil		根 Root		叶片 Leaf		茎 Shoot		全株 Total plant
	F	Sig.	F	Sig.	F	Sig.	F	Sig.	F	Sig.
Phosphorus (P)	324.527	0.000	121.359	0.000	188.985	0.000	57.784	0.000	431.250	0.000
pH	0.127	0.731	33.458	0.000	129.631	0.000	28.593	0.001	202.247	0.000
P × pH	5.428	0.055	2.314	0.167	56.035	0.000	6.718	0.032	24.811	0.001

图2 核桃各器官中磷含量(平均值±标准误差)。CK, 正常供磷素+ pH 6.0; T1, 正常供磷素+ pH 3.0; T2, 不添加磷素+ pH 6.0; T3, 不添加磷素+ pH 3.0。不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 2 Phosphorus content in different organs of Juglans regia (mean ± SE). CK, normal phosphorus and pH 6.0; T1, normal phosphorus and pH 3.0; T2, phosphorus deficiency and pH 6.0; T3, phosphorus deficiency and pH 3.0. Different small letters indicate significant differences among treatments (p < 0.05).

表3 不同处理条件下核桃幼苗的生长与生物量分配

Table 3 Growth and biomass allocation in Juglans regia seedlings under different treatments

生长指标 Growth variable	处理 Treatment
生长指标 Growth variable	CK	T1	T2	T3
根生物量 Root biomass	109.59^b	102.16^c	121.61^a	89.52^d
茎生物量 Shoot biomass	33.36^a	29.27^b	26.33^b	22.12^c
叶生物量 Leaf biomass	16.76^a	15.10^a	14.31^a	9.77^a
地下生物量/地上生物量 Root biomass / Shoot biomass	2.19^b	2.31^b	3.01^a	2.83^a
高生长 Height increment (cm)	10.23^a	9.18^b	8.68^b	7.68^c
茎生长 Diameter growth (mm)	4.76^a	2.61^b	2.09^bc	1.69^c
叶面积 Leaf area (cm²)	116.96^a	96.43^b	87.43^bc	82.82^c

表4 磷元素与pH值对核桃幼苗生长与生物量分配影响的双因素分析

Table 4 Two-way ANOVA for testing the effects of phosphorus and pH value on growth and biomass allocation in Juglans regia seedlings

	根生物量 Root biomass		茎生物量 Shoot biomass		叶生物量 Leaf biomass		地下生物量/地上生物量 Root biomass / Shoot biomass		高生长 Height increment (cm)		茎生长 Diameter growth (mm)		叶面积 Leaf area (cm²)
	F	Sig.	F	Sig.	F	Sig.	F	Sig.	F	Sig.	F	Sig.	F	Sig.
Phosphorus (P)	0.056	0.817	38.236	0.000	0.106	0.749	46.398	0.000	13.89	0.001	87.615	0.000	30.435	0.000
pH	224.433	0.000	13.113	0.002	0.108	0.747	0.085	0.775	9.360	0.004	45.942	0.000	8.775	0.004
P × pH	87.391	0.000	0.003	0.958	0.858	0.368	2.238	0.154	0.028	0.868	23.349	0.000	3.307	0.073

图3 各处理条件下核桃根系水分流速与压力差的关系。CK, 正常供磷素+ pH 6.0; T1, 正常供磷素+ pH 3.0; T2, 不添加磷素+ pH 6.0; T3, 不添加磷素+ pH 3.0。

Fig. 3 The relationships between pressure drop and water velocity in Juglans regia roots under different treatments. CK, normal phosphorus and pH 6.0; T1, normal phosphorus and pH 3.0; T2, phosphorus deficiency and pH 6.0; T3, phosphorus deficiency and pH 3.0.

图4 不同处理条件下核桃幼苗的根系导水率(平均值±标准误差)。CK, 正常供磷素+ pH 6.0; T1, 正常供磷素+ pH 3.0; T2, 不添加磷素+ pH 6.0; T3, 不添加磷素+ pH 3.0。不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 4 Root hydraulic conductivity in Juglans regia seedlings under different treatments (mean ± SE). CK, normal phosphorus and pH 6.0; T1, normal phosphorus and pH 3.0; T2, phosphorus deficiency and pH 6.0; T3, phosphorus deficiency and pH 3.0. Different small letters indicate significant differences among treatments (p < 0.05).

图5 不同处理条件下核桃叶片水势(平均值±标准误差)。CK, 正常供磷素+ pH 6.0; T1, 正常供磷素+ pH 3.0; T2, 不添加磷素+ pH 6.0; T3, 不添加磷素+ pH 3.0。不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 5 Leaf water potential (WP) in Juglans regia under different treatments (mean ± SE). CK, normal phosphorus and pH 6.0; T1, normal phosphorus and pH 3.0; T2, phosphorus deficiency and pH 6.0; T3, phosphorus deficiency and pH 3.0. Different small letters indicate significant differences among treatments (p < 0.05).

图6 不同处理条件下核桃叶柄导水率和导水损失率(平均值±标准误差)。CK, 正常供磷素+ pH 6.0; T1, 正常供磷素+ pH 3.0; T2, 不添加磷素+ pH 6.0; T3, 不添加磷素+ pH 3.0。不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 6 Hydraulic conductivity and percentage loss of hydraulic conductivity (PLC) in Juglans regia petioles under different treatments (mean ± SE). CK, normal phosphorus and pH 6.0; T1, normal phosphorus and pH 3.0; T2, phosphorus deficiency and pH 6.0; T3, phosphorus deficiency and pH 3.0. Different small letters indicate significant differences among treatments (p < 0.05).

表5 磷元素与pH值对核桃幼苗根系导水率及叶柄水分运输影响双因素分析

Table 5 Two-way ANOVA for testing the effects of phosphorus and pH value on root hydraulic conductivity and petiole water transport in Juglans regia seedlings

	根系导水率 Root hydraulic conductance		导水损失率 Percentage loss of hydraulic conductance		导水率 Hydraulic conductivity		导管密度 Vessel density (No.·mm^-2)		导管直径 Vessel diameter (μm)
	F	Sig.	F	Sig.	F	Sig.	F	Sig.	F	Sig.
Phosphorus (P)	5.132	0.043	59.903	0.000	33.211	0.000	76.056	0.000	6.285	0.017
pH	5.578	0.036	23.256	0.000	19.911	0.000	61.606	0.000	2.637	0.114
P × pH	0.367	0.556	4.646	0.051	1.069	0.317	2.714	0.108	0.525	0.474

图7 核桃幼苗在不同处理条件下气体交换的变异(平均值±标准误差)。CK, 正常供磷素+ pH 6.0; T1, 正常供磷素+ pH 3.0; T2, 不添加磷素+ pH 6.0; T3, 不添加磷素+ pH 3.0。不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 7 Variations of gas exchanges in Juglans regia seedlings under different treatments (mean ± SE). CK, normal phosphorus and pH 6.0; T1, normal phosphorus and pH 3.0; T2, phosphorous deficiency and pH 6.0; T3, phosphorus deficiency and pH 3.0. Different small letters indicate significant differences among treatments (p < 0.05).

表6 核桃幼苗叶柄木质部解剖结构(平均值±标准误差)

Table 6 Anatomical structure of petiole xylem in Juglans regia seedlings under different treatments (mean ± SE)

处理 Treatment	CK	T1	T2	T3
导管密度 Vessel density (No.·mm^-2)	318 ± 41.35^a	236 ± 23.30^b	229 ± 20.88^b	175 ± 16.41^c
导管直径 Vessel diameter (μm)	28.43 ± 3.32^a	27.74 ± 2.04^a	27.0 ± 2.31^ab	24.8 ± 1.84^b

表7 磷元素与pH值对核桃幼苗光合参数影响的双因素分析

Table 7 Two-way ANOVA for testing the effects of phosphorus and pH value on phosphorus characteristics in Juglans regia seedlings

	P_n		G_s		T_r		F_v/F_m		Φ_PSII		q_P		NPQ
	F	Sig.	F	Sig.	F	Sig.	F	Sig.	F	Sig.	F	Sig.	F	Sig.
Phosphorus (P)	48.676	0.000	19.368	0.000	20.750	0.000	15.241	0.001	20.073	0.000	26.746	0.000	22.131	0.000
pH	17.963	0.000	7.118	0.016	29.513	0.000	7.402	0.013	11.444	0.003	7.512	0.013	17.880	0.001
P × pH	1.447	0.245	0.247	0.626	4.459	0.052	0.013	0.910	0.066	0.800	0.748	0.398	0.184	0.674

图8 不同处理条件下核桃幼苗叶绿素荧光参数(平均值±标准误差)。CK, 正常供磷素+ pH 6.0; T1, 正常供磷素+ pH 3.0; T2, 不添加磷素+ pH 6.0; T3, 不添加磷素+ pH 3.0。ΦPSII, PSII有效光化学量子效率; Fv/Fm, PSII最大光化学量子效率; NPQ, 非光化学淬灭; qP, 光化学淬灭。不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 8 Parameters of chlorophyll fluorescence in Juglans regia seedlings under different treatments (mean ± SE). CK, normal phosphorus and pH 6.0; T1, normal phosphorus and pH 3.0; T2, phosphorus deficiency and pH 6.0; T3, phosphorus deficiency and pH 3.0. ΦPSII, quantum yield of PSII; Fv/Fm, maximum efficiency of PSII; NPQ, non-photochemical quenching; qP, photochemical quenching. Different small letters indicate significant differences among treatments (p < 0.05).

图9 不同处理条件下叶柄木质部导管的显微结构。 A, 正常供磷素+ pH 6.0。B, 正常供磷素+ pH 3.0。C, 不添加磷素 + pH 6.0。D, 不添加磷素+ pH 3.0。

Fig. 9 Microstructure of petiole xylem vessel under different treatments. A, Normal phosphorus and pH 6.0. B, Normal phosphorus and pH 3.0. C, Phosphorus deficiency and pH 6.0. D, Phosphorus deficiency and pH 3.0.

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