清凉峰不同海拔木本植物小枝内叶大小-数量权衡关系

doi:10.3724/SP.J.1258.2012.00281

摘要/Abstract

摘要：

权衡关系是生活史对策理论的基础, 叶大小-数量的权衡关系对理解叶大小进化具有重要的意义。该研究以单叶面积和单叶片干重表示叶大小, 用小枝干重和小枝茎干重表示小枝大小, 采用标准化主轴估计(standardized major axis estimation, SMA)和系统独立比较分析(phylogenetically independent contrast analysis, PIC)的方法, 对浙江省清凉峰自然保护区3个不同海拔落叶阔叶木本植物当年生小枝内的叶大小与数量间的关系进行研究。结果显示, 无论叶大小和小枝大小是用面积或干重表示, 在每个海拔, 叶大小与出叶强度均存在显著的等速负相关关系, 表明在落叶阔叶木本植物中发现的叶大小与出叶强度之间的权衡关系在不同生境物种中是普遍存在的, 植物在叶大小方面的种间变化, 可能不是自然选择的直接产物, 而是叶片数量变化权衡关系的一个副产物。不同海拔间的比较显示, 高海拔物种的叶面积或干重与出叶强度相关关系的y轴截距比中、低海拔物种小, 表明在出叶强度一定的情况下, 高海拔物种比低海拔物种具有更小的叶大小。与高海拔物种相比, 中海拔物种的共同斜率沿共同主轴有一个向上的位移, 表明中海拔物种比高海拔物种具有更大的叶大小, 但出叶强度更小。这些结果表明生境对叶大小-数量的权衡关系具有显著的影响, 中海拔生境具有更适宜植物生长的气候及养分条件, 而高海拔的低温等不利影响使得叶片变小。

关键词: 生境, 叶大小, 出叶强度, 权衡关系

Abstract:

Aims Trade-offs are fundamental to life-history strategies theory, and the leaf size and number trade-off is an important determinant of leaf-size evolution. It also has been proposed that this trade-off is dependent on habitat, but this is not well tested. Our objectives were to test whether the negative, isometric relationship between leaf size and number is conserved in different habitats and to explore the effects of altitude change on the relationship between the leaf size and number.
Methods Leaf area, mass and number and twig mass and stem mass of current-year twigs were measured for 61 deciduous broad-leaved woody species within three altitude-based habitats on Qingliang Mountain, southeastern China. The standardized major axis estimation method and the phylogenetically independent contrast method were used to examine the scaling relationship between leaf size (leaf mass and leaf area) and leafing intensity (twig mass and stem mass) within current-year twigs.
Important findings Significantly negative and isometric scaling relationships between leaf size and leaﬁng intensity were found to be consistent in all three altitude-based habitats, regardless of whether leaf/twig size was expressed as area or mass. However, the intercepts of these relationships significantly decreased with increasing altitude, suggesting that habitats constrain the leaf size that can be supported by a given leafing intensity. The middle-attitude species usually had significant upper shifts along the common slopes relative to the high-altitude species. This suggested that the middle altitude is a more suitable habitat with high nutrients and moderate climate conditions for plants, compared to the high altitude with low temperatures and nutrients.

Key words: habitat, leaf size, lea?ng intensity, trade-off

杨冬梅,占峰,张宏伟. 清凉峰不同海拔木本植物小枝内叶大小-数量权衡关系. 植物生态学报, 2012, 36(4): 281-291. DOI: 10.3724/SP.J.1258.2012.00281

YANG Dong-Mei,ZHAN Feng,ZHANG Hong-Wei. Trade-off between leaf size and number in current-year twigs of deciduous broad-leaved woody species at different altitudes on Qingliang Mountain, southeastern China. Chinese Journal of Plant Ecology, 2012, 36(4): 281-291. DOI: 10.3724/SP.J.1258.2012.00281

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https://www.plant-ecology.com/CN/Y2012/V36/I4/281

图/表 5

参考文献 70

[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. DOI URL
[2]	Ackerly DD, Reich PB (1999). Convergence and correlations among leaf size and function in seed plants: a comparative test using independent contrasts. American Journal of Botany, 86, 1272-1281. URL PMID
[3]	Barthélémy D, Caraglio Y (2007). Plant architecture: a dynamic, multilevel and comprehensive approach to plant form, structure and ontogeny. Annuals of Botany, 99, 375-407. DOI URL
[4]	Bell AD (1993). Plant Form: an Illustrated Guide to Flowering Plant Morphology. Oxford University Press, New York.
[5]	Bond WJ, Midgley J (1988). Allometry and sexual differences in leaf size. The American Naturalist, 131, 901-910. DOI URL
[6]	Bonsall MB, Jansen VAA, Hassell MP (2004). Life history trade-offs assemble ecological guilds. American Association for the Advancement of Science, 306, 111-114. DOI URL
[7]	Bonser SP, Aarssen LW (1996). Meristem allocation: a new classification theory for adaptive strategies in herbaceous plants. Oikos, 77, 347-352. DOI URL
[8]	Brouat C, Gibernau M, Amsellem L, McKey D (1998). Corner’s rules revisited: ontogenetic and interspecific patterns in leaf-stem allometry. New Phytologist, 139, 459-470. DOI URL
[9]	Cavender-Bares J, Holbrook NM (2001). Hydraulic properties and freezing-induced cavitation in sympatric evergreen and deciduous oaks with contrasting habitats. Plant, Cell & Environment, 24, 1243-1256.
[10]	Chapin FS III (1980). The mineral nutrition of wild plants. Annual Review of Ecology and Systematics, 11, 233-260. DOI URL
[11]	Cornelissen JHC (1999). A triangular relationship between leaf size and seed size among woody species: allometry, ontogeny, ecology and taxonomy. Oecologia, 118, 248-255. DOI URL PMID
[12]	Corner EJH (1949). The durian theory or the origin of the modern tree. Annals of Botany, 13, 367-414. DOI URL
[13]	Dombroskie SL, Aarssen LW (2010). Within-genus size distributions in angiosperms: small is better. Perspectives in Plant Ecology, Evolution and Systematics, 12, 283-293. DOI URL
[14]	Falster DS, Warton DI, Wright IJ (2006). User’s Guide to SMATR: Standardised Major Axis Tests & Routines Version 2.0. http://www.bio.mq.edu.au/ecology/SMATR/.Cited March 11,2012.
[15]	Falster DS, Westoby M (2003). Leaf size and angle vary widely across species: What consequences for light interception? New Phytologist, 158, 509-525. DOI URL
[16]	Field C, Mooney HA (1986). The photosynthesis-nitrogen relationship in wild plants. In: Givnish TJ ed. On the Economy of Plant Form and Function. Cambridge University Press, Cambridge, UK. 25-55.
[17]	Fonseca CR, Overton JMC, Collins B, Westoby M (2000). Shifts in trait-combinations along rainfall and phosphorus gradients. Journal of Ecology, 88, 964-977. DOI URL
[18]	Givnish TJ (1978a). On the adaptive significance of leaf height in forest herbs, with particular reference to tropical trees. In: Tomlinson PB, Zimmermann MH eds. Tropical Trees as Living Systems. Cambridge University Press, Cambridge, UK. 351-380.
[19]	Givnish TJ (1978b). Ecological aspects of plant morphology: leaf form in relation to environment. Acta Biotheoretica, 27, 83-142.
[20]	Givnish TJ (1979 375-407.
[21]	Givnish TJ (1984). Leaf and canopy adaptations in tropical forests. In: Medina E, Mooney HA, Vázquez-Yánez C eds. Physiological Ecology of Plants of the Wet Tropics. Junk Press, The Hague. 51-84.
[22]	Givnish TJ, Vermeij GJ (1976). Sizes and shapes of liane leaves. The American Naturalist, 110, 743-778. DOI URL
[23]	Gleason HA, Cronquist A (1991). Manual of Vascular Plants of Northeastern United States and Adjacent Canada. The New York Botanical Garden, Bronx.
[24]	Heuret P, Meredieu C, Coudurier T, Courdier F, Barthélémy D (2006). Ontogenetic trends in the morphological features of main stem annual shoots of Pinus pinaster( Pinaceae). American Journal of Botany, 93, 1577-1587. DOI URL
[25]	Huang CL (黄成林) (1992). Florogeographical analysis of Mount Qingliangfeng Natural Reserve in Tianmu Mountains. Journal of Zhejiang Forestry College (浙江林学院学报) 9, 277-282. (in Chinese with English abstract)
[26]	King DA (1998). Influence of leaf size on tree architecture: first branch height and crown dimensions in tropical rain forest trees. Trees-Structure and Function, 12, 438-445. DOI URL
[27]	Kleiman D, Aarssen LW (2007). The leaf size/number trade-off in trees. Journal of Ecology, 95, 376-382. DOI URL
[28]	Leishman MR (2001). Does the seed size/number trade-off model determine plant community structure? An assessment of the model mechanisms and their generality. Oikos, 93, 294-302. DOI URL
[29]	Li T, Deng JM, Wang GX, Cheng DL, Yu ZL (2009). Isometric scaling relationship between leaf number and size within current-year shoots of woody species across contrasting habitats. Polish Journal of Ecology, 57, 659-667.
[30]	Mcculloh KA, Sperry JS (2005). Patterns in hydraulic architecture and their implications for transport efficiency. Tree Physiology, 25, 257-267. DOI URL PMID
[31]	McDonald PG, Fonseca CR, Overton JM, Westoby M (2003). Leaf-size divergence along rainfall and soil-nutrient gradients: Is the method of size reduction common among clades? Functional Ecology, 17, 50-57. DOI URL
[32]	Milla R (2009). The leafing intensity premium hypothesis tested across clades, growth forms and altitudes. Journal of Ecology, 97, 972-983. DOI URL
[33]	Moles AT, Westoby M (2000). Do small leaves expand faster than large leaves, and do shorter expansion times reduce herbivore damage? Oikos, 90, 517-524. DOI URL
[34]	Molles MC, Cahill JF (1999). Ecology: Concepts and Applications. McGraw-Hill, New York. 509.
[35]	Niinemets Ü, Portsmuth A, Tobias M (2006). Leaf size modifies support biomass distribution among stems, petioles and mid-ribs in temperate plants. New Phytologist, 171, 91-104. DOI URL PMID
[36]	Niklas KJ (1988). The role of phyllotatic pattern as a “developmental constraint” on the interception of light by leaf surfaces. Evolution, 42, 1-16. DOI URL PMID
[37]	Niklas KJ, Cobb ED, Niinemets Ü, Reich PB, Sellin A, Shipley B, Wright IJ (2007). “Diminishing returns” in the scaling of functional leaf traits across and within species groups. Proceedings of the National Academy of Sciences of the United States of America, 104, 8891-8896. DOI URL PMID
[38]	Niklas KJ, Enquist BJ (2002). Canonical rules for plant organ biomass partitioning and annual allocation. American Journal of Botany, 89, 812-819. DOI URL PMID
[39]	Ogawa K (2008). The leaf mass/number trade-off of Kleiman and Aarssen implies constancy of leaf biomass, its density and carbon uptake in forest stands: scaling up from shoot to stand level. Journal of Ecology, 96, 188-191.
[40]	Parkhurst DF, Loucks OL (1972). Optimal leaf size in relation to environment. The Journal of Ecology, 60, 505-537. DOI URL
[41]	Pickup M, Westoby M, Basden A (2005). Dry mass costs of deploying leaf area in relation to leaf size. Functional Ecology, 19, 88-97. DOI URL
[42]	Pitman EJG (1939). A note on normal correlation. Biometrika, 31, 9-12. DOI URL
[43]	Poorter HS, Pepin S, Rijkers T, de Jong Y, Evans JR, Körner C (2006). Construction costs, chemical composition and payback time of high- and low-irradiance leaves. Journal of Experimental Botany, 57, 355-371. DOI URL PMID
[44]	Preston KA, Ackerly DD (2003). Hydraulic architecture and the evolution of shoot allometry in contrasting climates. American Journal of Botany, 90, 1502-1512. DOI URL PMID
[45]	Qian H (钱宏), Wang SL (汪思龙) (1988). Major forest vegetation types and their distributions in Qingliang Mountain Natural Reserve (JQNR) of Jixi County, Anhui Province. Journal of Ecology (生态学杂志) 7, 32-36. (in Chinese with English abstract)
[46]	Reich PB, Ellsworth DS, Walters MB, Vose JM, Gresham C, Volin C, Bowman WD (1999). Generality of leaf trait relationships: a test across six biomes. Ecology, 80, 1955-1969. DOI URL
[47]	Reich PB, Walters MB, Ellsworth DS (1992). Leaf life-span in relation to leaf, plant, and stand characteristics among diverse ecosystems. Ecological Monographs, 62, 365-392. DOI URL
[48]	Shao XP (邵小平) (2011). Vegetation types and vertical distribution zones spectrum analysis of Qingliang Mountain Natural Reserve in Anhui. Anhui Agricultural Science Bulletin (安徽农学通报) 17, 55-56. (in Chinese with English abstract).
[49]	Song CS (宋朝枢) (1997). Scientific Survey of the Qingliang Mountain Natural Reserve in Zhejiang (浙江清凉峰自然保护区科学考察集). China Forestry Publishing House, Beijing. (in Chinese)
[50]	Sun SC, Jin DM, Shi PL (2006). The leaf size-twig size spectrum of temperate woody species along an altitudinal gradient: an invariant allometric scaling relationship. Annals of Botany, 97, 97-107. DOI URL PMID
[51]	Walter A, Schurr U (2005). Dynamics of leaf and root growth: endogenous control versus environmental impact. Annuals of Botany, 95, 891-900. DOI URL
[52]	Warton DI, Weber NC (2002). Common slope tests for bivariate errors-in-variables models. Biometrical Journal, 44, 161-174. DOI URL
[53]	Warton DI, Wright IJ, Falster DS, Westoby M (2006). Bivariate line-fitting methods for allometry. Biological Reviews, 81, 259-291. DOI URL PMID
[54]	Watson MA, Casper BB (1984). Morphogenetic constraints on patterns of carbon distribution in plants. Annual Review of Ecology and Systematics, 15, 233-258. DOI URL
[55]	Westoby M, Falster DS, Moles AT, Vesk PA, Wright IJ (2002). Plant ecological strategies: some leading dimensions of variation between species. Annual Review of Ecology and Systematics, 33, 125-159. DOI URL
[56]	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
[57]	Whitman T, Aarssen LW (2010). The leaf size/number trade-off in herbaceous angiosperms. Journal of Plant Ecology, 3, 49-58. DOI URL
[58]	Woodward FI (1987). Climate and Plant Distribution. Cambridge University Press, Cambridge, UK. 174.
[59]	Woodward FI, Lomas MR, Kelly CK (2004). Global climate and the distribution of plant biomes. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 359, 1465-1476. DOI URL PMID
[60]	Wright IJ, Ackerly DD, Bongers F, Harms KE, Ibarra-Manriquez G, Martinez-Ramos M, Mazer SJ, Muller- Landau HC, Paz H, Pitman NCA, Poorter L, Silman MR, Vriesendorp CF, Webb CO, Westoby M, Wright SJ (2007). Relationships among ecologically important dimensions of plant trait variation in seven Neotropical forests. Annals of Botany, 99, 1003-1015. DOI URL PMID
[61]	Wright IJ, Reich PB, Cornelissen JHC, Falster DS, Groom PK, Hikosaka K, Lee W, Lusk CH., Niinemets Ü, Oleksyn J, Osada N, Poorter H, Warton DI, Westobyet M (2005). Modulation of leaf economic traits and trait relationships by climate. Global Ecology and Biogeography, 14, 411-421. DOI URL
[62]	Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornelissen JHC, Diemer M, Flexas J, Garnier E, Groom PK, Gulias J, Hikosaka K, Lamont BB, Lee T, Lee W, Lusk C, Midgley JJ, Navas ML, Niinemets U, Oleksyn J, Osada N, Poorter H, Poot P, Prior L, Pyankov VI, Roumet C, Thomas SC, Tjoelker MG, Veneklaas EJ, Villar R (2004). The worldwide leaf economics spectrum. Nature, 428, 821-827. DOI URL PMID
[63]	Wright IJ, Westoby M, Reich PB (2002). Convergence towards higher leaf mass per area in dry and nutrient-poor habitats has different consequences for leaf life span. Journal of Ecology, 90, 534-543. DOI URL
[64]	Xiang S, Liu YL, Fang F, Wu N, Sun SC (2009a). Stem architectural effect on leaf size, leaf number, and leaf mass fraction in plant twigs of woody species. International Journal of Plant Sciences, 170, 999-1008. DOI URL
[65]	Xiang S, Wu N, Sun SC (2009b). Within-twig biomass allocation in subtropical evergreen broad-leaved species along an altitudinal gradient: allometric scaling analysis. Trees-Structure and Function, 23, 637-647. DOI URL
[66]	Xiang S, Wu N, Sun SC (2010). Testing the generality of the “leafing intensity premium” hypothesis in temperate broad-leaved forests: a survey of variation in leaf size within and between habitats. Evolutionary Ecology, 24, 685-701. DOI URL
[67]	Yagi T (2004). Within-tree variations in shoot differentiation patterns of 10 tall tree species in a Japanese cool- temperate forest. Canadian Journal of Botany, 82, 228-243. DOI URL
[68]	Yagi T (2006). Relationships between shoot size and branching patterns in 10 broad-leaved tall tree species in a Japanese cool-temperate forest. Canadian Journal of Botany, 84, 1894-1907. DOI URL
[69]	Yagi T, Kikuzawa K (1999). Patterns in size-related variations in current-year shoot structure in eight deciduous tree species. Journal of Plant Research, 112, 343-352. DOI URL
[70]	Yang DM, Li GY, Sun SC (2008). The generality of leaf size versus number trade-off in temperate woody species. Annuals of Botany, 102, 623-629. DOI URL

指标 Index (y-x)	海拔 Altitude	样本量 Number of samples	决定系数 R²	斜率(95%置信区间) Slope (95% confidence interval)
单叶面积-基于枝干重的出叶强度 Individual leaf area-leafing intensity based on twig mass	低海拔 L	32	0.924	-1.004 (-1.113, -0.906)
	中海拔 M	29	0.887	-0.971 (-1.108, -0.851)
	高海拔 H	19	0.676	-0.854 (-1.139, -0.641)
单叶面积-基于茎干重的出叶强度 Individual leaf area-leafing intensity based on stem mass	低海拔 L	32	0.431	-0.968 (-1.278, -0.734)
	中海拔 M	29	0.306	-0.784 (-1.084, -0.568)
	高海拔 H	19	0.385	-0.752 (-1.116, -0.508)
单叶片干重-基于枝干重的出叶强度 Individual lamina mass-leafing intensity based on twig mass	低海拔 L	32	0.988	-1.012 (-1.054, -0.972)
	中海拔 M	29	0.986	-1.025 (-1.075, -0.978)
	高海拔 H	19	0.958	-1.046 (-1.161, -0.943)
单叶片干重-基于茎干重的出叶强度 Individual leaf area-leafing intensity based on stem mass	低海拔 L	32	0.494	-0.976 (-1.269, -0.751)
	中海拔 M	29	0.289	-0.829 (-1.149, -0.597)
	高海拔 H	19	0.309	-0.921 (-1.392, -0.609)

指标 Index (y-x)	海拔 Altitude	样本量 Number of samples	决定系数 R²	斜率(95%置信区间) Slope (95% confidence interval)
单叶面积-基于枝干重的出叶强度 Individual leaf area-leafing intensity based on twig mass	低海拔 L	32	0.924	-1.004 (-1.113, -0.906)
	中海拔 M	29	0.887	-0.971 (-1.108, -0.851)
	高海拔 H	19	0.676	-0.854 (-1.139, -0.641)
单叶面积-基于茎干重的出叶强度 Individual leaf area-leafing intensity based on stem mass	低海拔 L	32	0.431	-0.968 (-1.278, -0.734)
	中海拔 M	29	0.306	-0.784 (-1.084, -0.568)
	高海拔 H	19	0.385	-0.752 (-1.116, -0.508)
单叶片干重-基于枝干重的出叶强度 Individual lamina mass-leafing intensity based on twig mass	低海拔 L	32	0.988	-1.012 (-1.054, -0.972)
	中海拔 M	29	0.986	-1.025 (-1.075, -0.978)
	高海拔 H	19	0.958	-1.046 (-1.161, -0.943)
单叶片干重-基于茎干重的出叶强度 Individual leaf area-leafing intensity based on stem mass	低海拔 L	32	0.494	-0.976 (-1.269, -0.751)
	中海拔 M	29	0.289	-0.829 (-1.149, -0.597)
	高海拔 H	19	0.309	-0.921 (-1.392, -0.609)

指标 Index (y-x)	回归系数 Regression coefficient	回归截距 y-intercept	决定系数 R²
单叶面积-基于枝干重的出叶强度 Individual leaf area-leafing intensity based on twig mass	-0.957	0.016	0.919
单叶面积-基于茎干重的出叶强度 Individual leaf area-leafing intensity based on stem mass	-0.658	0.052	0.543
单叶片干重-基于枝干重的出叶强度 Individual lamina mass-leafing intensity based on twig mass	-1.008	0.005	0.991
单叶片干重-基于茎干重的出叶强度 Individual leaf area-leafing intensity based on stem mass	-0.684	0.043	0.570

指标 Index (y-x)	回归系数 Regression coefficient	回归截距 y-intercept	决定系数 R²
单叶面积-基于枝干重的出叶强度 Individual leaf area-leafing intensity based on twig mass	-0.957	0.016	0.919
单叶面积-基于茎干重的出叶强度 Individual leaf area-leafing intensity based on stem mass	-0.658	0.052	0.543
单叶片干重-基于枝干重的出叶强度 Individual lamina mass-leafing intensity based on twig mass	-1.008	0.005	0.991
单叶片干重-基于茎干重的出叶强度 Individual leaf area-leafing intensity based on stem mass	-0.684	0.043	0.570