生长环境与株龄对凤丹当年生果枝的生物量分配的影响

doi:10.7525/j.issn.1673-5102.2018.02.006

摘要/Abstract

摘要： 生物量分配研究是了解作物产量形成机制的基础。凤丹是以杨山牡丹（Paeonia ostii）为原种形成的栽培类群，也是新型木本油料作物油用牡丹的主栽品种。该文通过比较凤丹主要产区6龄种群、以及安徽铜陵和上海地区的4、6、8龄种群的当年生果枝大小与生物量分配，探讨环境和株龄对小枝生长与繁殖的影响。结果表明，凤丹植株的果枝数量随株龄增大而上升，不同株龄植株的单个果枝大小和果枝内生物量分配没有明显差异，果枝水平上没有株龄效应，但整株水平上株龄效应明显；果枝大小及生物量各指标之间存在明显相关性，但相关性在种群间变化较大；不同种群的果枝大小及生物量分配差异明显，果枝生物量分配明显随纬度、降水和光辐射强度变化而变化；异速生长方程模拟显示，大部分种群的果枝繁殖与营养生物量或总生物量之间为等速生长关系（斜率=1），但种群间截距变化较大。这些结果表明栽培环境对凤丹果枝的生长与繁殖有显著效应。

关键词: 果枝, 生物量分配, 株龄, 异速生长, 凤丹

Abstract: Biomass allocation is critical for understanding the yield formation of crops. Fengdan is the cultivated type originated from Paeonia ostii and the predominant variety of oil tree Paeonia that is newly cultivated as one of the oil tree crops in China. In order to examine the effects of growth environment and tree age on twig vegetative and reproductive growth, we compared the biomass characteristics of fruit twigs among eight 6-year-old Fengdan populations from different sites and that between 4, 6 and 8-year populations cultivated at Shanghai, and Tongling of Anhui Province. The number of fruit twigs increased with age increase, while there was no significant difference of twig growth and reproduction between different aged populations, indicating age effects existed at individual level but not twig level. There were significant correlations between the trait measurements, whereas, such correlations varied among populations. The body sizes and biomass allocations of fruit twigs were different among different populations, and the variations of biomass allocation associated with environmental variables. By allometric relationship modeling analysis, the fruit twigs from most of populations displayed constant biomass allocation between growth and reproduction functions(slope=1) with varied intercepts among populations. Therefore, the cultivation environment has significant effects on the growth and reproduction of Fengdan.

Key words: fruit twig, biomass allocation, tree age, allometric relationship, Fengdan (Paeonia ostia)

中图分类号:

Q949.746.5

马菡泽, 汪成忠, 李鼎, 范国祯, 胡永红, 宋志平. 生长环境与株龄对凤丹当年生果枝的生物量分配的影响[J]. 植物研究, 2018, 38(2): 201-211.

MA Han-Ze, WANG Cheng-Zhong, LI Ding, FAN Guo-Zhen, HU Yong-Hong, SONG Zhi-Ping. Effects of Growth Environment and Tree Age on Biomass Allocation within Fruit Twigs of Fengdan(Paeonia ostii)[J]. Bulletin of Botanical Research, 2018, 38(2): 201-211.

参考文献

1. Poorter H,Niklas K J,Reich P B,et al.Biomass allocation to leaves,stems and roots:meta-analyses of interspecific variation and environmental control[J].New Phytologist,2011,193(1):30-50.
2. Weiner J.Allocation,plasticity and allometry in plants[J].Perspectives in Plant Ecology,Evolution and Systematics,2004,6(4):207-215.
3. 高杰,郭子健,刘艳红.北京松山不同龄级天然油松林生物量分配格局及其影响因子[J].生态学杂志,2016,35(6):1475-1480. Gao J,Guo Z J,Liu Y H.Biomass allocation pattern and its influencing factors across natural Chinese pine forests of different ages in Songshan,Beijing[J].Chinese Journal of Ecology,2016,35(6):1475-1480.
4. 姚红,谭敦炎.胡卢巴属4种短命植物的生长特点及繁殖分配动态[J].新疆农业大学学报,2005,28(3):26-29. Yao H,Tan D Y.The growth characteristics and the reproductive allocation dynamics in four ephemeral species of Trigonella[J].Journal of Xinjiang Agricultural University,2005,28(3):26-29.
5. 刘贵周,蔡传涛,罗媛,等.两种萝芙木生物量分配与估测模型研究[J].云南植物研究,2009,31(1):63-66. Liu G Z,Cai C T,Zhou Y,et al.Biomass allocation and biomass estimation models of Rauvolfia vomitoria and Rauvolfia verticillata(Apocynaceae)[J].Acta Botanica Yunnanica,2009,31(1):63-66.
6. 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.
7. Weiner J,Lovett D J,Lovett D L.The influence of competition on plant reproduction[M].//Lovett Doust J,Lovett Doust L.Plant Reproductive Ecology:Patterns and Strategies.New York:Oxford University Press,1988:228-245.
8. 祝介东,孟婷婷,倪健,等.不同气候带间成熟林植物叶性状间异速生长关系随功能型的变异[J].植物生态学报2011,35(7):687-698. Zhu J D,Meng T T,NI J,et al.Within-leaf allometric relationships of mature forests in different bioclimatic zones vary with plant functional types[J].Chinese Journal of Plant Ecology,2011,35(7):687-698.
9. 杨冬梅,毛林灿,彭国全.常绿和落叶阔叶木本植物小枝内生物量分配关系研究:异速生长分析[J].植物研究,2011,31(4):472-477. Yang D M,Mao L C,Peng G Q.Within-twig biomass allocation in evergreen and deciduous broad-leaved species:allometric scaling analyses[J].Bulletin of Botanical Research,2011,31(4):472-477.
10. 李亚男,杨冬梅,孙书存,等.杜鹃花属植物小枝大小对小枝生物量分配及叶面积支持效率的影响:异速生长分析[J].植物生态学报,2008,32(5):1175-1183. Li Y N,Yang D M,Sun S C,et al.Effects of twig size on biomass allocation within twigs and on lamina area supporting efficiency in Rholdlden:allometric scaling analyses[J].Journal of Plant Ecology,2008,32(5):1175-1183.
11. Gu D,Yang D R,Compton S G,et al.Age at pollination modifies relative male and female reproductive success in a monoecious fig tree[J].Symbiosis,2012,57(2):73-81.
12. López F,Fungairiño S,De Las Heras P,et al.Age changes in the vegetative vs.reproductive allocation by module demographic strategies in a perennial plant[J].Plant Ecology,2001,157(1):13-21.
13. Carbone M S,Czimczik C I,Keenan T F,et al.Age,allocation and availability of nonstructural carbon in mature red maple trees[J].New Phytologist,2013,200(4):1145-1155.
14. Enright N J.Age,reproduction and biomass allocation in Rhopalostylis sapida(Nikau Palm)[J].Austral Journal of Ecology,2010,10(4):461-467.
15. Xiang S A,Wu N,Sun S C.Within-twig biomass allocation in subtropical evergreen broad-leaved species along an altitudinal gradient:allometric scaling analysis[J].Trees,2009,23(3):637-647.
16. Bonser S P,Aarssen L W.Meristem allocation:A new classification theory for adaptive strategies in herbaceous plants[J].Oikos,1996,77(2):347-352.
17. Pallas B,Mialet-Serra I,Rouan L,et al.Effect of source/sink ratios on yield components,growth dynamics and structural characteristics of oil palm(Elaeis guineensis) bunches[J].Tree Physiology,2013,33(4):409-424.
18. Cherbiy-Hoffmanna S U,Hallb A J,Rousseauxa M C.Fruit,yield,and vegetative growth responses to photosynthetically active radiation during oil synthesis in olive trees[J].Scientia Horticulturae,2013,150:110-116.
19. 赵宝军,宫永红.早实核桃优良单株产量主要影响因子的相关分析及通径分析[J].中国果树,2013,(3):27-29. Zhao B J,Gong Y H.The correlative analysis and pathway analysis of high productive early-fruiting walnut[J].China Fruit,2013,(3):27-29.
20. 陈春芳,吴云华,余崇彪,等.早实核桃产量影响因子研究初报[J].湖北林业科技,2008(4):21-23. Chen C F,Wu Y H,Yu C B,et al.Initial report on study of impact factor of early-fruit bearing production for Juglans regia L.[J].Hubei Forestry Science and Technology,2008(4):21-23.
21. Martens S N,Ustin S L,Norman J M.Measurement of tree canopy architecture[J].International Journal of Remote Sensing,1991,12(7):1525-1545.
22. 陈虹,董玉芝,朱小虎,等.新疆早实核桃品种测土配方施肥肥效试验初报[J].新疆农业科学,2010,47(8):1584-1589. Chen H,Dong Y Z,Zhu X H,et al.Preliminary discussion on the effects of fertilizer experiment with "3414" design in precocious walnut[J].Xinjiang Agricultural Sciences,2010,47(8):1584-1589.
23. 唐健,宋贤冲,曹继钊,等.不同产量油茶林树体结构及生物量分配差异[J].湖北农业科学,2013,52(12):2848-2850. Tang J,Song X C,Cao J Z,et al.Different yield oil Camellia tree structure and differences of biomass allocation[J].Hubei Agricultural Sciences,2013,52(12):2848-2850.
24. 康志雄,邹达明,王芷虔.用灰色系统理论分析油茶产量形成[J].浙江林业科技,1993,13(3):62-65. Kang Z X,Zou D M,Wang Z J.Analysis on formation of teaoil Camellia output by grey system theory[J].Zhejiang Forestry Science and Technology,1993,13(3):62-65.
25. 李晓青,韩继刚,刘炤,等.不同地区凤丹经济性状及其籽油脂肪酸成分分析[J].粮食与油脂,2014,27(4):43-46. Liu X Q,Han J G,Liu Z,et al.Economic characteristics investigation and seed oil fatty acid composition analysis of Paeonia ostii plants in different areas[J].Cereals & Oils,2014,27(4):43-46.
26. 陈慧玲,李洪喜,张建华,等.‘保康紫斑’牡丹生长适应性及结籽性状[J].林业工程学报,2014,28(4):43-46. Chen H L,Li H X,Zhang J H,et al.The growth adaptation and seed-bearing characteristics of Paeonia rockii ‘Baokang Ziban’ in Hubei Province[J].Journal of Forestry Engineering,2014,28(4):43-46.
27. 赵玉如,于晓金,谷欣,等.油用牡丹产量构成初步观测及分析[J].山东林业科技,2014,(5):79-81. Zhao Y R,Yu X J,Gu X,et al.The analysis and observation of yield components of oil peony[J].Journal of Shandong Forestry Science and Technology,2014(5):79-81.
28. 王佳.杨山牡丹遗传多样性与江南牡丹品种资源研究[D].北京:北京林业大学,2009:72-129. Wang J.Genetic diversity of Paeonia ostii and germplasm resources of tree peony cultivars form Chinese Jiangnan area[D].Beijing:Beijing Forestry University,2009:72-129.
29. 李嘉珏.中国牡丹与芍药[M].北京:中国林业出版社,1999:120-157. Li J Y.Chinese peony and Paeonia lactiflora[M].Beijing:China Forestry Publishing House,1999:120-157.
30. Harvey P H,Pagel M D.The comparative method in evolutionary biology[M].Oxford:Oxford University Press,1991:169-172.
31. Falster D S,Warton D I,Wright I J.User's guide to SMATR:standardised major axis tests & routines version 2.0[Z].2006.
32. Samuelson L J,Stokes T A,Butnor J R,et al.Ecosystem carbon density and allocation across a chronosequence of longleaf pine forests[J].Ecological Applications,2017,27(1):244-259.
33. Yang X,Yang Z Y,Wang X Y,et al.Tree biomass allocation and it's model additivity for Casuarina equisetifoliain a tropical forest of Hainan Island,China[J].PLoS One,2016,11(3):e0151858.
34. Li H,Li C Y,Zha T S,et al.Patterns of biomass allocation in an age-sequence of secondary Pinus bungeana forests in China[J].The Forestry Chronicle,2014,90(2):169-176.
35. 赵方,杨永平.中华山蓼不同海拔居群的繁殖分配研究[J].植物分类学报,2008,46(6):830-835. Zhao F,Yang Y P.Reproductive allocation in a dioecious perennial Oxyria sinensis(Polygonaceae) along altitudinal gradients[J].Journal of Systematics and Evolution,2008,46(6):830-835.
36. 赵志刚,杜国祯,任青吉.5种毛茛科植物个体大小依赖的繁殖分配和性分配[J].植物生态学报,2004,28(1):9-16. Zhao Z G,Du G Z,Ren Q J.Size-dependent reproduction and sex allocation in five species of Ranunculaceae[J].Chinese Journal of Plant Ecology,2004,28(1):9-16.
37. 王一峰,刘启茜,裴泽宇,等.青藏高原3种风毛菊属植物的繁殖分配与海拔高度的相关性[J].植物生态学报,2012,36(1):39-46. Wang Y F,Liu Q Q,Pei Z Y,et al.Correlation between altitude and reproductive allocation in three Saussurea species on China's Qinghai-Tibetan Plateau[J].Chinese Journal of Plant Ecology,2012,36(1):39-46.
38. 廖万金,张全国,张大勇.不同海拔藜芦种群繁殖特征的初步研究[J].植物生态学报,2003,27(2):240-248. Liao W J,Zhang Q G,Zhang D Y.A preliminary study on the reproductive features of Veratrum nigrum L. along an altitudinal gradient[J].Chinese Journal of Plant Ecology,2003,27(2):240-248.
39. Smith C C,Fretwell S D.The optimal balance between size and number of offspring[J].The American Naturalist,1974,108(962):499-506.
40. Li W,Pan W W,Tan R,et al.Loss of preferred habitat and pollen limitation threatens reproduction in a rare mountain Paeonia delavayi[J].Journal of Mountain Science,2016,13(12):2147-2154.
41. 刘左军,杜国祯,陈家宽.不同生境下黄帚橐吾(Ligularia virgaurea)个体大小依赖的繁殖分配[J].植物生态学报,2002,26(1):44-50. Liu Z J,Du G Z,Chen J K.Size-dependent reproductive allocation of Ligularia virgaurea in different habitats[J].Chinese Journal of Plant Ecology,2002,26(1):44-50.
42. 牛克昌,赵志刚,罗燕江,等.施肥对高寒草甸植物群落组分种繁殖分配的影响[J].植物生态学报,2006,30(5):817-826. Niu K C,Zhao Z G,Luo Y J,et al.Fertilization effects on species reproductive allocation in an alpine meadow plant community[J].Journal of Plant Ecology,2006,30(5):817-826.
43. Niklas K J.Plant allometry:is there a grand unifying theory?[J].Biological Reviews,2004,79(4):871-889.
44. Guo H,Weiner J,Mazer S J,et al.Reproductive allometry in Pedicularis species changes with elevation[J].Journal of Ecology,2012,100(2):452-458.
45. Weiner J,Rosenmeier L,Massoni E S,et al.Is reproductive allocation in Senecio vulgaris plastic?[J].Botany,2009,87(5):475-481.
46. Weiner J,Campbell L G,Pino J,et al.The allometry of reproduction within plant populations[J].Journal of Ecology,2009,97(6):1220-1233.

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