基于异速生长理论的准噶尔盆地荒漠灌丛形态研究

doi:10.3724/SP.J.1258.2011.00471

摘要/Abstract

摘要：

为了揭示荒漠灌丛形态的发生发展机制并认识其在荒漠生态系统中的功能, 从形态和结构决定功能的原理出发, 对生长在准噶尔荒漠东南部的岛状灌丛进行了形态学调查。依据Malthusian方程微分形式, 根据异速生长理论, 建立了冠幅与株高生长、灌丛表面积与体积生长的数学关系式, 利用植被调查数据进行了验证, 并最终得出不同灌丛在不同株高时的情景示意图。结果表明: 1)将荒漠灌丛形态假设成半三轴椭球体是合理的; 2)虽然灌丛形态发展趋势可以是扁平、近半球和竖直3种类型, 但是形态建成后, 一般维持在扁平和近半球两种类型; 3) 18类荒漠灌丛的体积和表面积的数量关系具有一定的一致性, 可能与同处于相同环境条件下的水分利用效率相近有关。

关键词: 异速生长理论, 冠幅, 冠型, 荒漠灌丛, 相对生长速率

Abstract:

Aims Our objectives were to investigate how shape and structure affect shrub architecture and to understand its function in the desert ecosystem through simulating differences in crown architecture of desert shrubs in China’s Junggar Basin.

Methods We analyzed shrub height and crown form. The difference between observed crown area and height and the estimated crown model was found to be a good measure of degree of crown development. A Malthusian conceptual model is proposed where crown area and height modify the crown architecture, shrub surface area and volume. We assumed that the three-dimensional structure of desert shrubs is the triaxial ellipsoid and chose shrub desert island in Junggar Basin as our object of study. According to plant allometric theory, we started from the differential form of the Malthusian equation and set up models describing (a) crown area and height growth and (b) surface area and volume growth. In addition, simulations were used for calculating the potential crown architecture of the desert shrub.

Important findings The assumption that the vertical distribution of desert shrub crown shape is semi-triaxial ellipsoid is valid. Three crown types were identified: flat, nearly hemispherical, and upright. These crown types corresponded well with the vertical distribution patterns of maximum height. The consistency of quantitative relationship between volume and surface area of desert shrubs is suggested as a useful tool for characterizing similar water-use strategy in the same environmental condition.

Key words: allometric theory, crown area, crown-type, desert shrub, relative growth rate

李嵩, 郑新军, 唐立松, 李彦. 基于异速生长理论的准噶尔盆地荒漠灌丛形态研究. 植物生态学报, 2011, 35(5): 471-479. DOI: 10.3724/SP.J.1258.2011.00471

LI Song, ZHENG Xin-Jun, TANG Li-Song, LI Yan. Morphological investigation of desert shrubs of China’s Junggar Basin based on allometric theory. Chinese Journal of Plant Ecology, 2011, 35(5): 471-479. DOI: 10.3724/SP.J.1258.2011.00471

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2011.00471

https://www.plant-ecology.com/CN/Y2011/V35/I5/471

图/表 5

表1 灌丛类型及其数量特征

Table 1 Shrub types and their quantitative characteristics

物种 Species	代号Code	相对密度Relative density (RD) (%)	相对频度Relative frequency (RF) (%)
白刺 Nitraria sibirica	Ns	1.2	13.3
淡枝沙拐枣 Calligonum leucocladum	Cal	3.3	56.7
白梭梭 Haloxylon persicum	Hp	4.0	56.7
柽柳 Tamarix spp.	Tr	2.9	26.7
黑果枸杞 Lycium ruthenicum	Lr	0.8	6.7
花花柴 Karelinia caspica	Kc	1.5	3.3
铃铛刺 Halimodendrom halodendron	Hh	0.4	6.7
疏叶骆驼刺 Alhagi sparsifolia	Als	1.4	6.7
毛足假木贼 Anabasis eriopoda	Ae	1.3	6.7
琵琶柴 Reaumuria soongorica	Rs	24.2	33.3
梭梭 Haloxylon ammodendron	Ha	38.5	93.3
驼绒藜 Ceratoides lateens	Cel	0.7	3.3
小叶碱蓬 Suaeda microphylla	Sm	0.7	13.3
盐节木 Halocnemum strobilaceum	Hs	0.6	3.3
盐生假木贼 Anabasis salsa	Ans	0.7	3.3
盐穗木 Halostachys caspica	Hc	2.3	10.0
盐爪爪 Kalidium spp.	Kf	13.2	16.7
准噶尔沙蒿 Artemisia songarica	Ars	2.3	13.3

图1 18种灌木的株高(A)、冠幅面积(B)、灌丛表面积(C)和体积(D)的分布。箱型图左、右的竖线为样本的25%和75%分位数, 箱型顶端和底部的差值为四分位极值; 箱型中间的竖线为样本中值, 若线不在箱型中央, 则表明存在偏度; 箱型向左或向右延伸的直线称为“触须”, 若没有统计异常值, 则样本最大值为右触须顶部, 样本最小值为左触须的底部。若样本中存在距离箱型左侧或右侧大于1.5倍内四分位极值的数值, 则认为样本中有统计异常值(如图用“+”表示)。若触须外无数据, 则底部触须有一点; 箱型上下两侧的V型槽口对应样本中值95%置信区间。物种代号见表1。

Fig. 1 Distribution of 18 shrubs’ height (A), crown area (B), shrub surface area (C) and volume (D). On each box, the reft and right edges of the box are the 25th and 75th percentiles, the difference of the top and bottom is quartile extreme; the central mark is the median, if the line doesn’t lie the central of the box, it shows skewness; box to the left or right extension of the line were known as “whisker”, if there were no statistical outliers, the maximum value of the sample should be the top of the right whiskers, and the sample minimum should be the bottom of the left whisker. It will be considered as statistical outliers in the samples if there are values more than 1.5 times of the quartile extreme from the left or right side of the box (see Figure with “+” indicates). A dot will be at the bottom of the tentacles if no data out of tentacles; the notch of both sides of the box corresponding to 95% confidence interval of the samples. Species code see Table 1.

图2 18类灌丛的冠幅与株高的相互关系。物种代号见表1。

Fig. 2 Relationship between height (H) and crown area (CA) of 18 shrubs. Species code see Table 1.

图3 18类灌丛的表面积与体积的相互关系。物种代号见表1。

Fig. 3 Relationship between surface area and volume of 18 shrubs. Species code see Table 1.

图4 18种灌木的冠型模拟图。物种代号见表1。

Fig. 4 Crown-type simulative diagrams of 18 desert shrubs. Species code see Table 1.

参考文献 26

[1]	Ackerly DD, Donoghue MJ (1998). Leaf size, sapling allometry, and Corner’s rules: phylogeny and correlated evolution in maples (Acer). The American Naturalist, 152, 767-791. URL PMID
[2]	Allen AP, Pockman WT, Restrepo C, Milne BT (2008). Allometry, growth and population regulation of the desert shrub Larrea tridentata. Functional Ecology, 22, 197-204. DOI URL
[3]	Archibald S, Bond WJ (2003). Growing tall vs growing wide: tree architecture and allometry of Acacia karroo in forest, savanna, and arid environments. Oikos, 102, 3-14. DOI URL
[4]	Brown JH, Gillooly JF, Allen AP, van Savage M, West GB (2004). Toward a metabolic theory of ecology. Ecology, 85, 1771-1789. DOI URL
[5]	Chesson P, Gebauer RLE, Schwinning S, Huntly N, Wiegand K, Ernest MSK, Sher A, Novoplansky A, Weltzin JF (2004). Resource pulses, species interactions, and diversity maintenance in arid and semi-arid environments. Oecologia, 141, 236-253. DOI URL PMID
[6]	Coomes DA, Grubb PJ (1998). A comparison of 12 tree species of Amazonian caatinga using growth rates in gaps and understorey, and allometric relationships. Functional Ecology, 12, 426-435. DOI URL
[7]	Fowler N (1986). The role of competition in plant communities in arid and semiarid regions. Annual Review of Ecology and Systematics, 17, 89-110. DOI URL
[8]	Han WX (韩文轩), Fang JY (方精云) (2008). Review on the mechanism models of allometric scaling laws: 3/4 vs. 2/3 power. Journal of Plant Ecology (Chinese Version) (植物生态学报), 32, 951-960. (in Chinese with English abstract)
[9]	Hein S, Spiecker H (2008). Crown and tree allometry of open-grown ash (Fraxinus excelsior L.) and sycamore (Acer pseudoplatanus L.). Agroforestry Systems, 73, 205-218. DOI URL
[10]	Ishii H, Clement JP, Shaw DC (2000). Branch growth and crown form in old coastal Douglas-fir. Forest Ecology and Management, 131, 81-91. DOI URL
[11]	Kidron GJ (2009). The effect of shrub canopy upon surface temperatures and evaporation in the Negev Desert. Earth Surface Processes and Landforms, 34, 123-132. DOI URL
[12]	Kohyama T, Hotta M (1990). Significance of allometry in tropical saplings. Functional Ecology, 4, 515-521. DOI URL
[13]	Liu JS (刘峻杉), Xu X (徐霞), Zhang Y (张勇), Tian YQ (田玉强), Gao Q (高琼) (2010). Effect of rainfall interannual variability on the biomass and soil water distribution in a semiarid shrub community. Science China (Life Sciences) (中国科学(生命科学)), 53, 729-737. (in Chinese)
[14]	Maguire DA, Hann DW (1987). Equations for predicting sapwood area at crown base in southwestern Oregon Douglas-fir. Canadian Journal of Forest Research, 17, 236-241. DOI URL
[15]	Reynolds JF, Kemp PR, Ogle K, Fernández RJ (2004). Modifying the “pulse-reserve” paradigm for deserts of North America: precipitation pulses, soil water, and plant responses. Oecologia, 141, 194-210. URL PMID
[16]	Schwinning S, Belnap J, Bowling DR, Ehleringer JR (2008). Sensitivity of the Colorado plateau to change: climate, ecosystems, and society. Ecology and Society, 13, 28.
[17]	Schwinning S, Sala OE, Loik ME, Ehleringer JR (2004). Thresholds, memory, and seasonality: understanding pulse dynamics in arid/semi-arid ecosystems. Oecologia, 141, 191-193. URL PMID
[18]	West GB, Woodruff WH, Brown JH (2002). Allometric scaling of metabolic rate from molecules and mitochondria to cells and mammals. Proceedings of the National Academy of Sciences of the United States of America, 99, 2473-2478. URL PMID
[19]	Westoby M, Wright IJ (2003). The leaf size-twig size spectrum and its relationship to other important spectra of variation among species. Oecologia, 135, 621-628. DOI URL PMID
[20]	Xiao S, Chen SY, Zhao LQ, Wang G (2006). Density effects on plant height growth and inequality in sunflower populations. Journal of Integrative Plant Biology, 48, 513-519. DOI URL
[21]	Xu H (许皓), Li Y (李彦), Xie JX (谢静霞), Cheng L (程磊), Zhao Y (赵彦), Liu R (刘冉) (2010). Influence of solar radiation and groundwater table on carbon balance of phreatophytic desert shrub Tamarix. Chinese Journal of Plant Ecology (植物生态学报), 34, 375-386. (in Chinese with English abstract)
[22]	Xu KX (徐克学) (2004). Biomathematics (生物数学). Science Press, Beijing. (in Chinese)
[23]	Zhang JT (张金屯) (2004). Quantitative Ecology (数量生态学). Science Press, Beijing. (in Chinese)
[24]	Zhao HL (赵哈林), Su YZ (苏永中), Zhang H (张华), Zhao LY (赵丽娅), Zhou RL (周瑞莲) (2007). Multiple effects of shrub on soil properties and understory vegetation in Horqin Sand Land, Inner Mongolia. Journal of Desert Research (中国沙漠), 27, 385-390. (in Chinese with English abstract)
[25]	Zheng XJ (郑新军), Wang QX (王勤学), Liu R (刘冉), Li Y (李彦) (2009). The condensation water input to the saline- alkaline desert ecosystem in the southeastern edge of the Junggar Basin. Progress in Natural Science (自然科学进展), 19, 1175-1186. (in Chinese)
[26]	Zhu YJ (朱雅娟), Jia ZQ (贾志清), Lu Q (卢琦), Hao YG (郝玉光), Zhang JB (张景波), Li L (李磊), Qi YL (綦艳林) (2010). Water use strategy of five shrubs in Ulanbuh Desert. Scientia Silvae Sinicae (林业科学), 46(4), 1-7. (in Chinese with English abstract)