不同生育期入侵植物大狼把草的生物量分配格局及异速生长分析

doi:10.11934/j.issn.1673-4831.2017.02.008

摘要/Abstract

摘要：

生物量分配格局和异速生长关系与植物生态策略的形成密切相关，对探讨植物对环境的适应性，揭示入侵植物的入侵机制有着重要意义。目前，对于入侵植物不同生育期之间的生物量分配格局和异速生长关系的差异研究较少。以入侵植物大狼把草（Bidens frondosa）为研究对象，采用挖掘法和称重法对大狼把草苗期、营养期和繁殖期的生长特性和生物量分配格局进行研究，并利用标准化主轴估计（standardized major axis estimation，SMA）方法对构件形态特性与生物量间的异速生长关系进行分析。结果表明：株高、植株基径、分枝数和叶片数均随着生育期的推进而增加，根长、总叶面积和平均单叶叶面积均在营养期达到最高。叶生物量分配比在苗期最大（48.88%），茎生物量分配比在生殖期最大（59.89%）；苗期和营养期根生物量分配比显著高于生殖期（P<0.05），茎生物量分配比随生育期的推进而升高，而叶生物量分配比随生育期的推进而下降，且不同生育期间存在显著差异（P<0.05）。大狼把草植株基径与各构件生物量间呈异速生长关系，总叶面积与茎、叶生物量大致呈等速生长关系，但不同生育期间、构件间的异速生长关系存在一定差异。可见，不同生育期大狼把草的生长特性、生物量分配格局及异速生长关系存在差异，显示了其灵活的生长和资源分配策略。

关键词: 生育期, 大狼把草, 入侵, 生物量分配, 异速生长

Abstract:

The relationship between biomass distribution and allometric growth is closely related to formation of plant ecological strategies, it is of great significance to understand adaptability of a plant to environment and to reveal mechanism of invasive plants invading a region. At present, not much has been reported concerning relationship between biomass distribution and allometric growth relative to growth stage of an invasive plant. A study on Bidens frondosa, an invasive species, was carried out using excavation and weight methods to explore growth characteristics and biomass distribution of the plant at its seedling stage, vegetative growing stage and reproductive stage. And then analysis was done of allometric relations between modular morphological characteristics and biomass with the standardized major axis estimation method (SMA). Results show that plant height, base stem diameter, branch number and leaf number increased with the plant growing from one stage to the next stage, and root length, total leaf area and mean single leaf area peaked at the vegetative growth stage. Leaf peaked in biomass distribution ratio at the seedling stage period (48.88%), while stem did at the reproductive stage (59.89%). Root at the seedling and vegetative growth stages was significantly higher in biomass distribution ratio than that at the reproductive stage (P<0.05). Stem increased in biomass distribution ratio with the plant growing from one stage to the next stage, while leaf did reversely and varied sharply with the stage (P<0.05). Moreover, base stem diameter of the plant was in allometric relationships with all the other plant parameters in biomass, whereas total leaf area was in isometric relationships with stem and leaf biomass. However, the allometric relationships between tissues of the plant varied with the growth stage. Obviously, growth characteristics, biomass distribution and allometric relationships of B. frondosa vary with the growth stage, indicating its flexible growth and resource allocation strategy.

Key words: growth stage, Bidens frondosa, invasion, biomass distribution, allometry

中图分类号:

Q945.79

闫小红, 何春兰, 周兵, 王宁, 尹增芳. 不同生育期入侵植物大狼把草的生物量分配格局及异速生长分析[J]. 生态与农村环境学报, 2017, 33(2): 150-158.

YAN Xiao-hong, HE Chun-lan, ZHOU Bing, WANG Ning, YIN Zeng-fang. Biomass Distribution and Allometric Analysis of Bidens frondosa Relative to Growth Stage[J]. Journal of Ecology and Rural Environment, 2017, 33(2): 150-158.

参考文献

[1] 郝靖,张婕,张沛沛,等.煤矸石场植被自然恢复初期草本植物生物量研究[J].草业学报,2013,22(4):51-60.[HAO Jing,ZHANG Jie,ZHANG Pei-pei,et al.A Study on the Biomass of Herbs at the Initial Natural Reclamation Stage of Plants in Gangue Fields[J].Acta Prataculturae Sinica,2013,22(4):51-60.]
[2] HILLS J M,MURPHY K J,PULFORD I D,et al.A Method for Classifying European Riverine Wetland Ecosystems Using Functional Vegetation Groups[J].Functional Ecology,1994,8(2):242-252.
[3] BAZZAZ F A,GRACE J.Plant Resource Allocation[M].London,UK:Academic Press,1997:1-37.
[4] WEINER J.Allocation,Plasticity and Allometry in Plants[J].Perspectives in Plant Ecology,Evolution and Systematics,2004,6(4):207-215.
[5] 代海军,何怀江,赵秀海,等.阔叶红松林两种主要树种的生物量分配格局及异速生长模型[J].应用与环境生物学报,2013,19(4):718-722.[DAI Hai-jun,HE Huai-jiang,ZHAO Xiu-hai,et al.Biomass Allocation Patterns and Allometric Models of Two Dominant Tress Species in Broad-Leaved and Korean Pine Mixed Forest[J].Chinese Journal of Applied and Environmental Biology,2013,19(4):718-722.]
[6] 陶冶,张元明.准噶尔荒漠6种类短命植物生物量分配与异速生长关系[J].草业学报,2014,23(2):38-48.[TAO Ye,ZHANG Yuan-ming.Biomass Allocation Patterns and Allometric Relationships of Six Ephemeroid Species in Junggar Basin,China[J].Acta Prataculturae Sinica,2014,23(2):38-48.]
[7] SHIBUYA M,YAJIMA T,YOSHIDA M,et al.Different Biomass-Allocation Patterns Among Four Tree Species in Heavily Disturbed Sites on a Volcanic Mountain in Hokkaido,Northern Japan[J].Ecological Research,2007,22(1):177-182.
[8] HOSSAIN M D,HANAFI M M,SALEH G,et al.Growth,Photosynthesis and Biomass Allocation of Different Kenaf (Hibiscus cannabinus L.) Accessions Grown on Sandy Soil[J].Australian Journal of Crop Science,2012,6(3):480-487.
[9] 王振兴,朱锦懋,王健,等.闽楠幼树光合特性及生物量分配对光环境的响应[J].生态学报,2012,32(12):3841-3848.[WANG Zhen-xing,ZHU Jin-mao,WANG Jian,et al.The Response of Photosynthetic Characters and Biomass Allocation of P. bournei Young Trees to Different Light Regimes[J].Acta Ecologica Sinica,2012,32(12):3841-3848.]
[10] PERRIN P M,MITCHELL F J G.Effects of Shade on Growth,Biomass Allocation and Leaf Morphology in European Yew (Taxus baccata L.)[J].European Journal of Forest Research,2013,132(2):211-218.
[11] SELZER L J,LENCINAS M V,MARTÍNEZ-PASTUR G J,et al.Light and Soil Moisture Effects on Biomass and Its Allocation in Osmorhiza depauperatua Philippi (Apiaceae)[J].Ecological Research,2013,28(3):469-480.
[12] 程军回,张元明.水分胁迫下荒漠地区2种草本植物生物量分配策略[J].干旱区研究,2012,29(3):432-439.[CHEN Jun-hui,ZHANG Yuan-ming.Strategies for Biomass Allocation of Two Desert Plant Species Under Water Stress[J].Arid Zone Research,2012,29(3):432-439.]
[13] YANG Y,WANG G,YANG L,et al.Effects of Drought and Warming on Biomass,Nutrient Allocation,and Oxidative Stress in Abies fabri in Eastern Tibetan Plateau[J].Journal of Plant Growth Regulation,2013,32(2):298-306.
[14] WANG J C,DUAN B L,ZHANG Y B.Effects of Experimental Warming on Growth,Biomass Allocation,and Needle Chemistry of Abies faxoniana in Even-Aged Monospecific Stands[J].Plant Ecology,2012,213(1):47-55.
[15] 朱铁霞,乌日娜,于永奇.不同氮肥施用量下菊芋株高及各器官生物量分配动态研究[J].草地学报,2014,22(1):199-202.[ZHU Tie-xia,WU Ri-na,YU Yong-qi.Effect of Nitrogen on the Plant Height and Dynamic Biomass Allocation of Helianthus tuberosus[J].Acta Agrestia Sinica,2014,22(1):199-202.]
[16] 赵光影,刘景双,王洋,等.大气CO₂浓度升高和氮肥施用对三江平原湿地小叶章生物量及根冠比的影响[J].生态与农村环境学报,2009,25(1):38-41.[ZHAO Guang-ying,LIU Jing-shuang,WANG Yang,et al.Effects of Elevated CO₂ Concentration and Application of N Fertilizer on Biomass and Root/Shoot Ratio of Calamagrostis angustifolia in Typical Wetland of Sanjiang Plain[J].Journal of Ecology and Rural Envrionment,2009,25(1):38-41.]
[17] PRENTIS P J,WILSON J R U,DORMONTT E E,et al.Adaptive Evolution in Invasive Species[J].Trends Plant Science,2008,13(6):288-294.
[18] PYŠEK P,RICHARDSON D M.Traits Associated With Invasiveness in Alien Plants:Where Do We Stand?[M]//NENTWIG W.Biological Invasions,Section Ⅱ.Berlin,Germany:Verlag-Springer,2007.
[19] SULTAN S E.Phenotypic Plasticity and Plant Adaptation[J].Plant Biology,1995,44(4):363-383.
[20] 耿宇鹏,张文驹,李博,等.表型可塑性与外来植物的入侵能力[J].生物多样性,2004,12(4):447-455.[GENG Yu-peng,ZHANG Wen-ju,LI Bo,et al.Phenotypic Plasticity and Invasiveness of Alien Plants[J].Biodiversity Science,2004,12(4):447-455.]
[21] 潘玉梅,唐赛春,岑艳喜,等.钻形紫菀开花期种群构件的生物量分配[J].热带亚热带植物学报,2010,18(2):176-181.[PAN Yu-mei,TANG Sai-chun,CEN Yan-xi,et al.Biomass Allocation on the Modules of Aster subulatus Michx. Population at Flowering Stage[J].Journal of Tropical and Subtropical Botany,2010,18(2):176-181.]
[22] 潘玉梅,唐赛春,蒲高忠,等.外来入侵植物小飞蓬种群构件生物量结构特征[J].广西植物,2009,29(4):476-480.[PAN Yu-mei,TANG Sai-chun,PU Gao-zhong,et al.Module Biomass Structure of the Alien Invasive Plant Conyza canadensis[J].Guihaia,2009,29(4):476-480.]
[23] FENG Y L,WANG J F,SANG W G.Biomass Allocation,Morphology and Photosynthesis of Invasive and Noninvasive Exotic Species Grown at Four Irradiance Levels[J].Acta Oecologica,2007,31(1):40-47.
[24] GENG Y P,PAN X Y,XU C Y,et al.Plasticity and Ontogenetic Drift of Biomass Allocation in Response to Above-and Below-Ground Resource Availabilities in Perennial Herbs:A Case Study of Alternanthera philoxeroides[J].Ecological Research,2007,22(2):255-260.
[25] 任丽娟,王国祥,仇乐,等.江苏潮滩湿地不同生境互花米草形态与生物量分配特征[J].生态与农村环境学报,2010,26(3):220-226.[REN Li-juan,WANG Guo-xiang,QIU Le,et al.Morphology and Biomass Distribution of Spartina alterniflora Growing in Different Tidal Flat Habitats in Jiangsu[J].Journal of Ecology and Rural Envrionment,2010,26(3):220-226.]
[26] ZHENG Y L,FENG Y L,LIU W X,et al.Growth,Biomass Allocation,Morphology,and Photosynthesis of Invasive Eupatorium Adenophorum and Its Native Congeners Grown at Four Irradiances[J].Plant Ecology,2009,203(2):263-271.
[27] 李天林,申时才,徐高峰,等.薇甘菊不同时期的营养繁殖及其生物量分配特征[J].西北植物学报,2012,32(7):1377-1383.[LI Tian-lin,SHEN Shi-cai,XU Gao-feng,et al.Vegetative Reproduction and Biomass Allocation Characteristics of Mikania micrantha H. B. K Under Different Seedling Periods[J].Acta Botanica Boreali-Occidentalia Sinica,2012,32(7):1377-1383.]
[28] MCALPINE K G,JESSON L K.Biomass Allocation,Shade Tolerance and Seedling Survival of the Invasive Species Berberis darwinii (Darwin's Barberry)[J].New Zealand Journal of Ecology,2007,31(1):1-12.
[29] 王艳婷,孙瑜琳,王美玲,等.节节麦不同生育期叶片蜡质组成和超微形态的变化[J].麦类作物学报,2014,34(11):1516-1521.[WANG Yan-ting,SUN Yu-lin,WANG Mei-ling,et al.Composition and Ultrastructure Variation of Leaf Culticular Wax at Different Developing Stages of Aegilops tauschii[J].Journal of Triticeae Crops,2014,34(11):1516-1521.]
[30] KIM M S,SHIM K B,PARK S H,et al.Changes in Cuticular Waxes of Developing Leaves in Sesame (Sesamum indicum L.)[J].Journal of Crop Science and Biotechnology,2009,12(3):161-167.
[31] 陆霞梅,周长芳,安树青,等.植物的表型可塑性、异速生长及其入侵能力[J].生态学杂志,2007,26(9):1438-1444.[LU Xia-mei,ZHOU Chang-fang,AN Shu-qing,et al.Phenotypic Plasticity,Allometry and Invasiveness of Plants[J].Chinese Journal of Ecology,2007,26(9):1438-1444.]
[32] WEST G B,BROWN J H,ENQUIST B J.A General Model for the Origin of Allometric Scaling Laws in Biology[J].Science,1997,276(5309):122-126.
[33] NIKLAS K J.Modelling Below-and Above-Ground Biomass for Non-Woody and Woody Plants[J].Annals of Botany,2005,95(2):315-321.
[34] MCCONNAUGHAY K D M,COLEMAN J S.Biomass Allocation in Plants:Ontogeny or Optimality? A Test Along Three Resource Gradients[J].Ecology,1999,80(8):2581-2593.
[35] 董点,林天喜,唐景毅,等.紫椴生物量分配格局及异速生长方程[J].北京林业大学学报,2014,36(4):54-63.[DONG Dian,LIN Tian-xi,TANG Jing-yi,et al.Biomass Allocation Patterns and Allometric Models of Tilia amurensis[J].Journal of Beijing Forestry University,2014,36(4):54-63.]
[36] 汪珍川,杜虎,宋同清,等.广西主要树种(组)异速生长模型及森林生物量特征[J].生态学报,2015,35(13):4462-4472.[WANG Zhen-chuan,DU Hu,SONG Tong-qing,et al.Allometric Models of Major Tree Species and Forest Biomass in Guangxi[J].Acta Ecologica Sinica,2015,35(13):4462-4472.]
[37] XIAO C W,CEULEMANS R.Allometric Relationships for Below-and Above-Ground Biomass of Young Scots Pines[J].Forest Ecology and Management,2004,203(1/2/3):177-186.
[38] LIM H,LEE K H,LEE K H,et al.Biomass Expansion Factors and Allometric Equations in an Age Sequence for Japanese Cedar (Cryptomeria japonica) in Southern Korea[J].Journal of Forest Research,2013,18(4):316-322.
[39] 李钰,赵成章,侯兆疆,等.高寒退化草地狼毒种群个体大小与茎、叶的异速生长[J].生态学杂志,2013,32(2):241-246.[LI Yu,ZHAO Cheng-zhang,HOU Zhao-jiang,et al.Body Size and Stem and Leaf Allometry of Stellera chamaejasme in Degraded Alpine Grassland[J].Chinese Journal of Ecology,2013,32(2):241-246.]
[40] SCHIMID B,WINER J.Plastic Relationship Between Reproductive and Vegetative Mass in Solidago altissima[J].Evolution,1993,47(1):61-74.
[41] 万方浩,刘全儒,谢明.生物入侵:中国外来入侵植物图鉴[M].北京:科学出版社,2012.[WAN Fang-hao,LIU Quan-ru,XIE Ming.Biological Invasions:Color Illustrations of Invasive Alien Plants in China[M].Beijing:Science Press,2012.]
[42] ŠUMBEROVÁK,TZONEV R,VLADIMIROV V.Bidens frondosa (Asteraceae):A New Alien Species for the Bulgarian Flora[J].Phytologia Balcanica,2004,10(2/3):179-181.
[43] COSKUNELCEBI K,TERZIOgLU S,VLADIMIROV V.A New Alien Species for the Flora of Turkey:Bidens frondosa L.(Asteraceae)[J].Turkish Journal of Botany,2007,31(5):477-479.
[44] BRÄNDEL M.Dormancy and Germination of Heteromorphic Achenes of Bidens frondosa[J].Flora:Morphology,Distribution,Functional Ecology of Plants,2004,199(3):228-233.
[45] HAN Y G,CHO Y H,KIM Y J,et al.Insect Herbivores Associated With the Introduced Weed Bidens frondosa L.(Asteraceae) in Korea,and Their Potential Role as Augmentative Biological Control Agents[J].Entomological Research,2009,39(6):394-400.
[46] 闫小红,曾建军,周兵,等.外来入侵植物大狼把草提取物的化感潜力[J].扬州大学学报(农业与生命科学版),2012,33(2):88-94.[YAN Xiao-hong,ZENG Jian-jun,ZHOU Bing,et al.Allelopathic Potential of the Extracts From Alien Invasive Plant Bidens frondosa[J].Journal of Yangzhou University (Agricultural and Life Science Edition),2012,33(2):88-94.]
[47] 周兵,闫小红,肖宜安,等.外来入侵植物大狼把草种群构件生物量结构研究[J].广西植物,2012,32(5):650-655.[ZHOU Bing,YAN Xiao-hong,XIAO Yi-an,et al.Module Biomass Structure Traits of the Alien Invasive Bidens frondosa Population[J].Guihaia,2012,32(5):650-655.]
[48] FALSTER D S,WARTON D I,WRIGHT I J.User's Guide to SMATR:Standardized Major Axis Tests & Routines Version 2.0.[Z/OL].(2006-03-11)[2016-01-15].http://www.bio.mq.edu.au/ecology/SMATR/.
[49] SCHLICHTING C D,SMITH H.Phenotypic Plasticity:Linking Molecular Mechanisms With Evolutionary Outcomes[J].Evolutionary Ecology,2002,16(3):189-211.
[50] SLOT M,KLOOSTER S H J,STERCK F J,et al.A Lifetime Perspective of Biomass Allocation in Quercus pubescens Trees in a Dry,Alpine Valley[J].Trees,2012,26(5):1661-1668.
[51] 梁艳,张小翠,陈学林.多年生龙胆属植物个体大小与花期资源分配研究[J].西北植物学报,2008,28(12):2400-2407.[LIANG Yan,ZHANG Xiao-cui,CHEN Xue-lin.Individual Size and Resource Allocation in Perennial Gentiana[J].Acta Botanica Boreali-Occidentalia Sinica,2008,28(12):2400-2407.]
[52] 梁坤伦,张洪荣,张丽静,等.紫穗槐表型可塑性及植株资源分配对高寒生境的响应[J].草业科学,2012,29(3):440-446.[LIANG Kun-lun,ZHANG Hong-rong,ZHANG Li-jing,et al.Response of Phenotypic Plasticity and Plant Resource Allocation of Amorpha fruiticosa to Alpine Habitat[J].Pratacultural Science,2012,29(3):440-446.]
[53] BROUWER R.Distribution of Dry Matter in the Plant[J].Netherlands Journal of Agricultural Science,1962,10(3):361-376.
[54] 刘龙昌,范伟杰,董雷鸣,等.入侵植物小花山桃草种群构件生物量结构及种子萌发特征[J].广西植物,2012,32(1):69-76.[LIU Long-chang,FAN Wei-jie,DONG Lei-ming,et al.Module Biomass Structure and Seed Germination Traits of the Exotic Invasive Plant Gaura parviflora[J].Guihaia,2012,32(1):69-76.]
[55] 齐淑艳,徐文铎,文言.外来入侵植物牛膝菊种群构件生物量结构[J].应用生态学报,2006,17(12):2283-2286.[QI Shu-yan,XU Wen-duo,WEN Yan.Biomass Structure of Exotic Invasive Plant Galinsona parviflora[J].Chinese Journal of Applied Ecology,2006,17(12):2283-2286.]
[56] TILMAN D.Plant Strategies and the Dynamics and Structure of Plant Communities[M].Princeton,USA:Princeton University Press,1988:37-52.
[57] STOHLGREN T J,CROSIER C,CHONG G W,et al.Life-History Habitat Matching in Invading Non-Native Plant Species[J].Plant and Soil,2005,277(1):7-18.
[58] COCK A G.Genetical Aspects of Metrical Growth and Form in Animals[J].Quarterly Review of Biology,1966,41(2):131-190.
[59] 汪金松,张春雨,范秀华,等.臭冷杉生物量分配格局及异速生长模型[J].生态学报,2011,31(14):3918-3927.[WANG Jin-song,ZHANG Chun-yu,FAN Xiu-hua,et al.Biomass Allocation Patterns and Allometric Models of Abies nephrolepis Maxim[J].Acta Ecologica Sinica,2011,31(14):3918-3927.]
[60] NIKLAS K J.Plant Biomechanics:An Engineering Approach to Plant Form and Function[M].Chicago,USA:University of Chicago Press,1992:261-312.