高寒矮生嵩草草甸主要植物物候特征对养分和水分添加的响应

doi:10.3724/SP.J.1258.2014.00013

摘要/Abstract

摘要：

植物物候特征对环境条件的季节和年际变化具有较强的指示作用, 因此研究植物物候特征对环境条件变化的响应, 对理解植物和环境之间的相互作用关系、植物的适应机制和生存策略, 以及应对全球变化都具有重要的意义。该研究基于2009-2011年高寒矮生嵩草(Kobresia humilis)草甸养分水分控制实验的植物物候观测数据资料, 采用巢式方差分析、物候指数和聚类分析方法, 开展了高寒矮生嵩草草甸主要植物物候特征对养分和水分添加的响应研究。结果表明: (1)养分添加处理之间植物返青期和枯黄期均无显著差异, 但养分添加中氮磷处理对主要物种作用较明显, 使莎草科、禾本科、杂类草主要代表植物的返青期和枯黄期推迟。(2)增雪处理效应明显, 主要优势物种无论是何种养分添加, 在增雪处理后均表现出花期物候提前的趋势(p < 0.01), 同时增雪处理使杂类草植物返青期显著提前(p < 0.05)。增水处理对植物的作用效果并不一致, 其中垂穗披碱草(Elymus nutans)和双柱头藨草(Scirpus distigmaticus)的枯黄期显著推迟(p < 0.05), 而杂类草枯黄期提前。(3)养分添加后, 不同物种的物候特征表现出显著差异(p < 0.01), 例如雪白委陵菜(Potentilla nivea)枯黄期显著推迟(p < 0.05), 而双柱头藨草的枯黄期显著提前(p < 0.05), 但物种对养分添加响应的差异以植物类群为单位, 禾本科植物表现为返青期推迟, 而莎草科植物表现为返青期提前。(4)矮生嵩草草甸主要植物营养生长期与果后营养期持续天数之间呈负相关关系, 主要植物物候特征经聚类分析可以分为3个类群, 3个类群经氮磷钾、钾和氮钾三个养分添加处理后植物物候特征变化较大。研究表明, 高寒矮生嵩草草甸植物物候特征在物种水平响应和水分添加后的响应表现出较大差异, 而对养分添加的响应不显著。

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关键词: 返青期, 水分, 养分, 物候, 物候指数, 枯黄期

Abstract:

Aims Phenology is a sensitive, integrated indicator of environmental changes. It is important to research the responses of phenological characteristics to environmental changes for understanding interactions between plants and environment, adaptive mechanisms and survival strategies. Our objective is to examine how phenological characteristics of plants respond to nutrient and water conditions in the eastern Qinghai-Tibet Plateau.
Methods Plant phenological characteristics in alpine Kobresia humilis meadow following nutrient and water additions from 2009 to 2011 were quantitatively analyzed by the methods of nested analysis of variance, phonological index and cluster analysis.
Important findings There were no significant differences in either green-up date or senescence date of plants with nutrient addition, but these dates in several dominant species of grasses, sedges and forbs were postponed following N and P addition. The treatments of nutrient and winter moisture addition moved up flowering dates of dominant species (p < 0.01) and green-up dates of forbs (p < 0.05). Effects of nutrient and summer moisture addition were inconsistent. The senescence dates of Elymus nutans and Scirpus distigmaticus were significantly postponed (p < 0.05) and forbs were postponed. Phenological characteristics of different species had significant differences with nutrient addition (p < 0.01): the senescence dates of Potentilla nivea were significantly postponed (p < 0.05), the senescence dates of Scirpus distigmaticus were significantly moved up (p < 0.05), but the different phenology responses of plants to the nutrient addition were based mainly on plant groups, with the green-up dates of grasses mostly postponed but the green-up dates of sedges moved up. Continuance of vegetative growth and vegetative period after fruiting were negative correlated. Phenological characteristics of plants were divided into three groups by cluster analysis. Phenological characteristics of the treatment of nitrogen-phosphorus-potassium mixed fertilizer, potassium and nitrogen-potassium mixed fertilizer addition varied considerably among plant groups. In summary, phenological characteristics of species in K. humilis meadow displayed large differences after moisture addition, but displayed smaller differences after nutrient addition.

Key words: green-up date, moisture, nutrient, phenology, phenology index, senescence date

叶鑫, 周华坤, 刘国华, 姚步青, 赵新全. 高寒矮生嵩草草甸主要植物物候特征对养分和水分添加的响应. 植物生态学报, 2014, 38(2): 147-158. DOI: 10.3724/SP.J.1258.2014.00013

YE Xin, ZHOU Hua-Kun, LIU Guo-Hua, YAO Bu-Qing, ZHAO Xin-Quan. Responses of phenological characteristics of major plants to nutrient and water additions in Kobresia humilis alpine meadow. Chinese Journal of Plant Ecology, 2014, 38(2): 147-158. DOI: 10.3724/SP.J.1258.2014.00013

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2014.00013

https://www.plant-ecology.com/CN/Y2014/V38/I2/147

图/表 12

表1 每年每小区化肥施加量(g)

Table 1 Amount of applied fertilizer per plot per year (g)

养分类别 Nutrient category	养分形态 Nutrient form	添加量 Addition (g·36 m^-2·a^-1)
N	CO(NH₂)₂	771.43
P	Ca(H₂PO₄)₂·H₂O	1 464.39
K	K₂SO₄	802.40
C	C₁₂H₂₂O₁₁	5 148.00
Ca^*	CaCO₃	538.39
Mg^*	MgCO₃	374.67
Fe^*	FeSO₄·7H₂O	3 043.58
Mn^*	MnSO₄	247.54
Zn^*	ZnSO₄·7H₂O	158.26
Cu^*	CuSO₄·5H₂O	141.51
B^*	Na₂B₄O₇·10H₂O	31.77
Mo^*	Na₂MoO₄·2H₂O	9.08

图1 实验样地设计图。I, 养分添加; II, 养分添加并夏季增雨; III, 养分添加并冬季增雪; IV, 预留区; ①观测区; ②采样区; ③其他实验区。

Fig. 1 Design draws of experimental plots. I, nutrient addition; II, nutrient addition and summer water addition; III, nutrient addition and winter snow addition; IV, reserve zone; ①, observation zone; ②, sampling zone; ③ other experimental plot.

图2 养分添加后矮生嵩草的开花比率。C, 碳; CK, 对照; K, 钾; N, 氮; NK, 氮钾混合肥料; NP, 氮磷混合肥料; NPK, 氮磷钾混合肥料; P, 磷; PK, 磷钾混合肥料。159 days, 儒略历第159天; 162 days, 儒略历第162天。

Fig. 2 Florescence ratios of Kobresia humilis after nutrient addition. C, carbon; CK, control; K, potassium; N, nitrogen; NK, nitrogen-potassium mixed fertilizer; NP, nitrogen-phosp- horus mixed fertilizer; NPK, nitrogen-phosphorus-potassium mixed fertilizer; P, phosphorus; PK, phosphorus-potassium mixed fertilizer. 159 days, the 159th day of Julian calendar; 162 days, the 162th day of Julian calendar.

图3 养分添加后第三年麻花艽返青期和枯黄期物候比率。图注同图2。

Fig. 3 Phenological ratios of green-up date and senescence date of Gentiana straminea in the 3rd year after nutrient addition. Notes see Fig. 2.

图4 养分添加后麻花艽的返青比率。图注同图2。

Fig. 4 Green-up ratios of Gentiana straminea after nutrient addition. Notes see Fig. 2.

图5 养分添加第二年垂穗披碱草和双柱头藨草的枯黄比率(平均值±标准误差)。Nutrient, 养分添加; Nutrient + Snow, 养分添加并冬季增雪。不同大、小写字母表示差异显著(p < 0.05)。

Fig. 5 Senescence ratios of Elymus nutans and Scirpus distigmaticus in the 2nd year after nutrient addition (mean ± SE). Nutrient, nutrient addition; Nutrient + Snow, nutrient addition and winter snow addition. Different upper-case and lower-case letters indicate significant difference (p < 0.05).

图6 养分添加第三年矮生嵩草6月初的开花比率。Nutrient, 养分添加; Nutrient + Snow, 养分添加并冬季增雪; Nutrient + Water, 养分添加并夏季增水; C + CK, 碳+对照; Other nutrients, 除碳、对照外的养分添加。

Fig. 6 Florescence ratios of Kobresia humilis in early June of the 3rd year after nutrient addition. Nutrient, nutrient addition; Nutrient + Snow, nutrient addition and winter snow addition; Nutrient + Water, nutrient addition and summer water addition; C + CK, carbon + control; Other nutrients, nutrient addition expect carbon and control.

表2 2010年5月1日(儒略历第121天)不同养分添加处理间的植物返青比率

Table 2 Green-up ratios of plant with different nutrient additions on May 1st, 2010 (the 121th day of Julian calendar)

物种 Species	养分添加 Nutrient addition
物种 Species	NPK	K	NP	N	P	PK	NK	C	Control
矮生嵩草 Kobresia humilis	0.83	0.83	1.00⁺	0.77	0.75	1.00⁺	1.00⁺	0.92	0.75
垂穗披碱草 Elymus nutans	0.58	0.55	0.31^-	0.39^-	0.67	0.62	0.50^-	0.62	0.75
双柱头藨草 Scirpus distigmaticcus	0.46^-	1.00	0.50^-	0.75	0.63^-	0.82	0.89	0.67	1.00
麻花艽 Gentiana straminea	0.46⁺	0.42⁺	0.17	0.08	0.50⁺	0.33	0.15	0.25	0.08
美丽凤毛菊 Saussurea superba	0.42^-	0.58	0.33^-	0.73⁺	0.55	0.83⁺	0.69⁺	0.58	0.55
黑褐穗薹草 Carex atrofusca	0.60	0.71⁺	0.75⁺	0.67	1.00⁺	0.20^-	0.70	0.62	0.50
雪白委陵菜 Potentilla nivea	0.67^-	0.91	0.54^-	0.92⁺	1.00⁺	0.91	0.92⁺	0.67	0.83
冷地早熟禾 Poa crymophila	0.70	0.67^-	0.70	0.70	0.70	0.83	0.42^-	0.62	0.83
异针茅 Stipa aliena	0.08^-	0.36	0.30^-	0.33^-	0.42	0.46	0.50	0.58	0.88

图7 养分添加第二年物种的返青比率。不同小写字母表示差异显著(p < 0.05)。

Fig. 7 Green-up ratios of species in the 2nd year after nutrient addition. Different lower-case letters indicate significant difference (p < 0.05).

表3 2010年10月22日(儒略历第295天)不同养分添加处理间植物种的枯黄比率

Table 3 Senescence ratios of plant species with different nutrient additions on Oct. 22th, 2010 (the 295th day of Julian calendar)

物种 Species	养分添加 Nutrient addition								Control
物种 Species	NPK	K	NP	N	P	PK	NK	C	Control
矮生嵩草 Kobresia humilis	0.93⁺	0.71	0.95⁺	0.83	0.83	0.82	0.86⁺	0.78^-	0.80
垂穗披碱草 Elymus nutans	0.79	0.96⁺	0.87	0.88	0.85	0.91	0.89	0.83^-	0.91
双柱头藨草 Scirpus distigmaticcus	0.89⁺	0.52	0.72⁺	0.62	0.93⁺	0.58	0.57	0.66	0.54
麻花艽 Gentiana straminea	0.75	0.90⁺	0.94⁺	0.77	0.88⁺	0.84	0.87	0.73^-	0.84
美丽凤毛菊 Saussurea superba	0.92	0.88	0.97⁺	0.95	0.98⁺	0.94	0.99⁺	0.83^-	0.89
黑褐穗薹草 Carex atrofusca	0.72	0.89	0.82	0.90	0.92⁺	0.78^-	0.88	0.85	0.91
雪白委陵菜 Potentilla nivea	0.83	0.77	0.93	0.83	0.81	0.88	0.94	0.74^-	0.96
异叶米口袋 Gueldenstaedtia diversifolia	0.96	1.00⁺	0.89	0.97	0.96	0.95	1.00⁺	0.99⁺	0.95
甘青剪股颖 Agrostis hugoniana	0.85	0.93	0.86	0.95⁺	0.89	0.95⁺	0.90	0.96⁺	0.86
异针茅 Stipa aliena	0.80	0.78	0.73	0.81	0.71	0.85	0.68^-	0.86⁺	0.85
冷地早熟禾 Poa crymophila	0.63	0.87⁺	0.78	0.80⁺	0.61	0.67^-	0.69	0.88⁺	0.70

图8 养分添加第二年物种枯黄比率。不同小写字母表示差异显著(p < 0.05)。

Fig. 8 Senescence ratios of species in the 2nd year after nutrient addition. Different lower-case letters indicate significant difference (p < 0.05).

图9 矮生嵩草草甸植被物候持续天数的聚类分析。

Fig. 9 Cluster analysis of vegetation phenological lasting days of Kobresia humilis meadow.

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