基于植物功能性状的暖温带5种灌木幼苗生长策略

doi:10.17521/cjpe.2021.0396

摘要/Abstract

摘要：

为了理解中国暖温带常见本地灌木和外来灌木的生长策略和生态位分化机制, 并为植被恢复和重建、区域生态环境可持续发展提供理论依据, 该研究选取3种中国暖温带常见本地灌木——荆条(Vitex negundo var. heterophylla)、酸枣(Ziziphus jujuba var. spinosa)、小花扁担杆(Grewia biloba var. parviflora)和2种外来灌木——紫穗槐(Amorpha fruticosa)和火炬树(Rhus typhina), 通过对植物功能性状的测量, 分析了不同物种幼苗生长策略的差异。该研究在山东大学莱芜房干生态实验站进行, 利用盆栽实验种植5种灌木的当年实生苗, 每个物种20个重复, 测量了叶结构性状、叶营养元素含量、叶绿素含量、叶气体交换参数、叶绿素荧光、生物量产量等性状指标。结果表明: 5种灌木在相同的生长环境下表现出了不同的生理生态特性。外来种火炬树和紫穗槐表现出较高的光合速率, 这对于两者充分利用林下光斑环境非常有利。紫穗槐支撑结构比例高达0.434, 有利于其叶片运动, 进而适应多变的光照环境, 较强的光合适应能力是2个外来种能够定居下来的重要原因。本地种荆条和酸枣对资源的利用比较保守, 两者气体交换值均较低, 充分地利用正午的光照是荆条叶片生理的一大特点。小花扁担杆的叶片有较强的气体交换能力, 但是通过叶绿素荧光分析发现, 其正午过后的实际量子产量恢复比较慢, 另外, 小花扁担杆叶片的比叶质量和叶绿素a/b都最小, 分别为44.539 g·m^-2和2.215, 说明小花扁担杆具有一定的阴生性。良好的种子扩散能力(百粒质量小, 0.947 g)和叶散热能力(周长²/面积(γ′)值最高, 48.389)、保守的光能利用策略、较强的干扰后恢复能力(根冠比高, 5.051)使得荆条成为中国暖温带地区的广布优势种。

关键词: 生长策略, 叶性状, 生理生态学, 植物功能性状, 灌木

Abstract:

Aims The subject of plant functional traits, which reflect plant growth and survival strategies, is an important trending topic in plant ecology. In this study, we selected three native warm temperate shrub species in China: Vitex negundo var. heterophylla, Ziziphus jujuba var. spinosa, Grewia biloba var. parviflora and two alien species: Amorpha fruticosa and Rhus typhina, to compare their growth and survival strategies.
Methods The differences in growth strategies were analyzed by measuring plant functional traits. This experiment was carried out at Fanggan ecological experimental station of Shandong University in Jinan. The seeds of five shrubs were germinated and 20 seedlings of each species were planted in pots containing a mixture of sand and humus (2:1, by volume). The leaf structural traits, leaf nitrogen, phosphorus and potassium, chlorophyll content, leaf gas exchange parameters, chlorophyll fluorescence, biomass, and seedling growth parameters were measured.
Important findings Physiological and ecological characteristics of five shrub seedlings were different under the same condition. 1) The alien species, i.e. R. typhina and A. fruticosa, showed high photosynthetic rates and had a strong ability to adjust to the instantaneous change of light intensity, which may be beneficial for plants to make full use of sunflecks under the canopy. The high supporting structure ratio of A. fruticosa is also more conducive to leaf movements, thus adjusting to the changeable light environment. The supporting structure ratio of A. fruticosa was as high as 0.434, which is conducive to its leaf movement and adjustment to the changeable light environment. The high photosynthetic capacity of R. typhina and A. fruticosa was the vital reason for the establishment of the two alien species. 2) The local species V. negundo var. heterophylla and Z. jujuba var. spinosa demonstrated relatively conservative utilization of resources, and in both of them low gas exchange values were measured. However, V. negundo var. heterophylla could make full use of the midday light. 3) G. biloba var. parviflora showed strong gas exchange capacity, and the actual quantum yield of diurnal variation of fluorescence was slow to recover in the afternoon as demonstrated by the chlorophyll fluorescence analysis. In addition, Grewia biloba var. parviflora had the lowest specific leaf mass (44.539 g·m^-2) and chlorophyll a/b (2.215), which confirmed that it is a shade-tolerant plant. Traits such as good seed dispersal ability, leaf heat dissipation ability, conservative light energy utilization strategy, and strong ability to recover after disturbance, help V. negundo var. heterophylla become the dominant species in the warm temperate zone of China. In summary, our study is helpful to understand the growth strategy and niche differentiation mechanisms in several common shrubs of warm temperate zone of China and provides theoretical basis for vegetation restoration, vegetation reconstruction and sustainable development of regional ecological environment.

Key words: growth strategy, leaf trait, physiological ecology, plant functional trait, shrub

祁鲁玉, 陈浩楠, 库丽洪·赛热别力, 籍天宇, 孟高德, 秦慧颖, 王宁, 宋逸欣, 刘春雨, 杜宁, 郭卫华. 基于植物功能性状的暖温带5种灌木幼苗生长策略. 植物生态学报, 2022, 46(11): 1388-1399. DOI: 10.17521/cjpe.2021.0396

QI Lu-Yu, CHEN Hao-Nan, Kulihong SAIREBIELI, JI Tian-Yu, MENG Gao-De, QIN Hui-Ying, WANG Ning, SONG Yi-Xin, LIU Chun-Yu, DU Ning, GUO Wei-Hua. Growth strategies of five shrub seedlings in warm temperate zone based on plant functional traits. Chinese Journal of Plant Ecology, 2022, 46(11): 1388-1399. DOI: 10.17521/cjpe.2021.0396

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https://www.plant-ecology.com/CN/Y2022/V46/I11/1388

图/表 6

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性状 Trait	荆条 Vitex negundo var. heterophylla	酸枣 Ziziphus jujuba var. spinosa	小花扁担杆 Grewia biloba var. parviflora	紫穗槐 Amorpha fruticosa	火炬树 Rhus typhina
Area (cm²)	9.261 ± 1.045^c	4.652 ± 0.410^c	54.529 ± 3.560^a	4.509 ± 0.314^c	22.574 ± 0.817^b
L/W	2.380 ± 0.126^b	2.142 ± 0.052^b	1.476 ± 0.053^c	3.157 ± 0.165^a	3.296 ± 0.117^a
L/PL	6.088 ± 0.555^c	27.319 ± 2.846^a	8.940 ± 0.215^c	19.041 ± 0.811^b	-
γ′	48.389 ± 5.577^a	21.526 ± 0.981^c	20.755 ± 0.430^c	20.438 ± 0.708^c	36.152 ± 1.332^b
LMA (g·m^-2)	48.094 ± 0.611^bc	49.303 ± 1.698^ab	44.539 ± 1.513^c	53.040 ± 1.342^a	46.389 ± 1.397^bc
x(c) (cm·cm^-1)	0.417 ± 0.006^c	0.447 ± 0.006^b	0.402 ± 0.004^d	0.483 ± 0.004^a	0.392 ± 0.003^d
N (g·kg^-1)	32.825 ± 0.581^c	42.105 ± 0.768^a	37.878 ± 1.221^b	36.800 ± 1.158^b	27.073 ± 1.103^d
P (g·kg^-1)	3.540 ± 0.186^b	5.968 ± 0.111^a	5.593 ± 0.462^a	5.208 ± 0.548^a	5.793 ± 0.284^a
K (g·kg^-1)	12.015 ± 0.889^b	15.040 ± 0.142^a	12.478 ± 0.514^b	9.895 ± 0.368^c	10.073 ± 0.540^c
N:P	9.371 ± 0.632^a	7.068 ± 0.236^b	6.908 ± 0.586^b	7.274 ± 0.689^b	4.710 ± 0.309^c
Chl a (mg·g^-1)	1.565 ± 0.060^c	2.316 ± 0.112^a	2.055 ± 0.085^b	2.013 ± 0.040^b	1.771 ± 0.064^c
Chl b (mg·g^-1)	0.667 ± 0.043^c	0.878 ± 0.057^ab	0.938 ± 0.064^a	0.812 ± 0.027^abc	0.726 ± 0.037^bc
Chl a/b	2.384 ± 0.156	2.658 ± 0.076	2.215 ± 0.081	2.494 ± 0.096	2.446 ± 0.035
LBP	0.878 ± 0.004^b	-	0.965 ± 0.001^a	0.790 ± 0.006^c	0.764 ± 0.007^d

性状 Trait	荆条 Vitex negundo var. heterophylla	酸枣 Ziziphus jujuba var. spinosa	小花扁担杆 Grewia biloba var. parviflora	紫穗槐 Amorpha fruticosa	火炬树 Rhus typhina
Area (cm²)	9.261 ± 1.045^c	4.652 ± 0.410^c	54.529 ± 3.560^a	4.509 ± 0.314^c	22.574 ± 0.817^b
L/W	2.380 ± 0.126^b	2.142 ± 0.052^b	1.476 ± 0.053^c	3.157 ± 0.165^a	3.296 ± 0.117^a
L/PL	6.088 ± 0.555^c	27.319 ± 2.846^a	8.940 ± 0.215^c	19.041 ± 0.811^b	-
γ′	48.389 ± 5.577^a	21.526 ± 0.981^c	20.755 ± 0.430^c	20.438 ± 0.708^c	36.152 ± 1.332^b
LMA (g·m^-2)	48.094 ± 0.611^bc	49.303 ± 1.698^ab	44.539 ± 1.513^c	53.040 ± 1.342^a	46.389 ± 1.397^bc
x(c) (cm·cm^-1)	0.417 ± 0.006^c	0.447 ± 0.006^b	0.402 ± 0.004^d	0.483 ± 0.004^a	0.392 ± 0.003^d
N (g·kg^-1)	32.825 ± 0.581^c	42.105 ± 0.768^a	37.878 ± 1.221^b	36.800 ± 1.158^b	27.073 ± 1.103^d
P (g·kg^-1)	3.540 ± 0.186^b	5.968 ± 0.111^a	5.593 ± 0.462^a	5.208 ± 0.548^a	5.793 ± 0.284^a
K (g·kg^-1)	12.015 ± 0.889^b	15.040 ± 0.142^a	12.478 ± 0.514^b	9.895 ± 0.368^c	10.073 ± 0.540^c
N:P	9.371 ± 0.632^a	7.068 ± 0.236^b	6.908 ± 0.586^b	7.274 ± 0.689^b	4.710 ± 0.309^c
Chl a (mg·g^-1)	1.565 ± 0.060^c	2.316 ± 0.112^a	2.055 ± 0.085^b	2.013 ± 0.040^b	1.771 ± 0.064^c
Chl b (mg·g^-1)	0.667 ± 0.043^c	0.878 ± 0.057^ab	0.938 ± 0.064^a	0.812 ± 0.027^abc	0.726 ± 0.037^bc
Chl a/b	2.384 ± 0.156	2.658 ± 0.076	2.215 ± 0.081	2.494 ± 0.096	2.446 ± 0.035
LBP	0.878 ± 0.004^b	-	0.965 ± 0.001^a	0.790 ± 0.006^c	0.764 ± 0.007^d

	LMA	γ′	L/W	L/PL	LBP	N	P	K	N:P	Chl a	Chl b	Chl a/b	Chl t
Area	-0.048	0.168	0.525^**	-0.027	-0.700^**	-0.600^**	0.096	-0.814^**	-0.573^**	-0.172	-0.089	-0.134	-0.146
LMA		0.071	0.425^**	0.281	-0.243	0.269	0.116	0.098	0.170	-0.083	-0.210	0.232	-0.078
γ′			0.386^**	-0.486^**	-0.199	-0.635^**	-0.400	-0.140	0.081	-0.650^**	-0.575^**	-0.035	-0.671^**
L/W				0.293	-0.787^**	-0.367	-0.050	-0.490^*	-0.155	-0.263	-0.414^*	0.340	-0.305
L/PL					-0.446^*	0.579^*	0.424	0.135	-0.262	0.557^**	0.425^*	0.302	0.564^**
LBP						0.582^*	-0.131	0.715^**	0.491	0.142	0.361	-0.473^*	0.199
N							0.387	0.638^**	0.280	0.722^**	0.418	0.325	0.689^**
P								0.118	-0.704^**	0.480^*	0.316	0.097	0.423
K									0.281	0.242	0.189	0.203	0.236
N:P										-0.078	-0.156	0.156	-0.051
Chl a											0.796^**	0.166	0.970^**
Chl b												-0.378^*	0.895^**
Chl a/b													-0.013

	LMA	γ′	L/W	L/PL	LBP	N	P	K	N:P	Chl a	Chl b	Chl a/b	Chl t
Area	-0.048	0.168	0.525^**	-0.027	-0.700^**	-0.600^**	0.096	-0.814^**	-0.573^**	-0.172	-0.089	-0.134	-0.146
LMA		0.071	0.425^**	0.281	-0.243	0.269	0.116	0.098	0.170	-0.083	-0.210	0.232	-0.078
γ′			0.386^**	-0.486^**	-0.199	-0.635^**	-0.400	-0.140	0.081	-0.650^**	-0.575^**	-0.035	-0.671^**
L/W				0.293	-0.787^**	-0.367	-0.050	-0.490^*	-0.155	-0.263	-0.414^*	0.340	-0.305
L/PL					-0.446^*	0.579^*	0.424	0.135	-0.262	0.557^**	0.425^*	0.302	0.564^**
LBP						0.582^*	-0.131	0.715^**	0.491	0.142	0.361	-0.473^*	0.199
N							0.387	0.638^**	0.280	0.722^**	0.418	0.325	0.689^**
P								0.118	-0.704^**	0.480^*	0.316	0.097	0.423
K									0.281	0.242	0.189	0.203	0.236
N:P										-0.078	-0.156	0.156	-0.051
Chl a											0.796^**	0.166	0.970^**
Chl b												-0.378^*	0.895^**
Chl a/b													-0.013

性状 Trait	荆条 Vitex negundo var. heterophylla	酸枣 Ziziphus jujuba var. spinosa	小花扁担杆 Grewia biloba var. parviflora	紫穗槐 Amorpha fruticosa	火炬树 Rhus typhina
HSM (g)	0.947 ± 0.013^c	17.095 ± 0.119^a	4.605 ± 0.025^b	0.833 ± 0.007^c	0.919 ± 0.010^c
RMR	0.295 ± 0.015^a	0.187 ± 0.014^bc	0.162 ± 0.020^c	0.218 ± 0.017^b	0.181 ± 0.018^bc
SMR	0.284 ± 0.014^b	0.328 ± 0.020^a	0.328 ± 0.012^a	0.340 ± 0.005^a	0.179 ± 0.005^c
LMR	0.420 ± 0.008^c	0.485 ± 0.012^b	0.510 ± 0.009^b	0.442 ± 0.021^c	0.639 ± 0.019^a
R/S	0.422 ± 0.031^a	0.231 ± 0.020^b	0.196 ± 0.030^b	0.281 ± 0.027^b	0.224 ± 0.027^b
BR/AR	5.051 ± 0.209^a	1.184 ± 0.121^b	1.629 ± 0.329^b	1.842 ± 0.248^b	5.485 ± 0.596^a
B/MS	0.266 ± 0.053^c	0.772 ± 0.080^b	1.109 ± 0.103^a	0^d	0^d
P/L	0.094 ± 0.006^c	-	0.038 ± 0.003^d	0.271 ± 0.013^b	0.308 ± 0.016^a
STMR	0.320 ± 0.016^b	0.328 ± 0.020^b	0.346 ± 0.012^b	0.434 ± 0.004^a	0.329 ± 0.010^b