栓皮栎实生幼苗生长发育动态特征
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摘要
为了解栓皮栎幼苗各器官在不同季节的生长动态,以便进一步研究和探讨幼苗生长受阻和死亡的原因,本研究详细跟踪0～3年生栓皮栎幼苗的生命进程,测定幼苗各器官生长量、干物质量以及糖分浓度。结果表明:1) 1年生幼苗夏季茎和叶片有明显的生长受阻现象; 2) 1～2年生幼苗根生长除12月至翌年3月的冬季停止伸长外,一直伸长生长,3年生幼苗根则伸长生长缓慢; 3)由种仁干物质消耗和糖分浓度分析可知,1年生幼苗夏季(6月上旬开始)的生长受阻,与种仁剩余量和糖分浓度最低相匹配; 4) 6月上旬种仁总糖浓度开始低于叶片总糖浓度,此时起种仁不再是植株主要的营养供体; 5) 2、3年生幼苗从3月中旬开始生长,至7月份生长变缓,没有生长停滞现象;因此,0～3年生栓皮栎幼苗生态策略为保证主根生长,1年生幼苗存在夏季生长受阻现象,这与幼苗抗性最弱(异养自养转换)的时期密切相关,2、3年生幼苗无生长停滞现象。
[Background] Plant growth dynamics reveals the impacts of the environment on plants.Quercus variabilis is one of the afforestation species widely used in rocky mountainous areas. It is prone to growth retardation or death under environmental stress. In order to further reveal the reasons for the stunted seedling growth and death,this paper traced the life course of each organ. [Methods] In this study,the seedlings of Quercus variabilis were sampled by hand-excavation in the nursery field. The sampling interval was 10 d in the first year,one month in the second year,and one season in the third year,and the growth,dry biomass and sugar concentration of every organ in the seedlings were measured by drying and Anthrone colorimetry. [Results]1) During the first growing season,there was obvious growth-stasis-regrowth of stem and leaf,showing the phenomenon of blocked growth in summer. 2)Except for the winter season( December-March of the next year),the roots of 1/2-year-old seedlings grew continuously. At the age of 3,the rate of root elongation was slower than at 1/2 years,with root thickening and lateral root growth being the main growth. 3) Analysis of seed kernel dry matter consumption and total sugar concentration showed that the growth of seedlings in their first summer( from early June) was blocked,which was in line with the lowest seed kernel residue and total sugar. 4) In early June,the total sugar concentration of the seed kernel began to fall beneath the total sugar concentration of the leaf,indicating that the seed kernel was no longer the main nutrient donor of the plant. 5) Seedlings aged 2 to 3 years began to grow from the middle of March,and growth slowed down in July,and they did not show the phenomenon of blocked growth. [Conclusions] The ecological strategy of 0-3 years old seedlings of Q. variabilis was to ensure the growth of the main roots,the 1-year-old seedlings have stunted growth in summer,which is closely related to the period of weakest seedling resistance transform( heterotrophic autotrophic transformation),and there was no stagnation in the growth of 2 or 3 year old seedlings.
[Background] Plant growth dynamics reveals the impacts of the environment on plants.Quercus variabilis is one of the afforestation species widely used in rocky mountainous areas. It is prone to growth retardation or death under environmental stress. In order to further reveal the reasons for the stunted seedling growth and death,this paper traced the life course of each organ. [Methods] In this study,the seedlings of Quercus variabilis were sampled by hand-excavation in the nursery field. The sampling interval was 10 d in the first year,one month in the second year,and one season in the third year,and the growth,dry biomass and sugar concentration of every organ in the seedlings were measured by drying and Anthrone colorimetry. [Results]1) During the first growing season,there was obvious growth-stasis-regrowth of stem and leaf,showing the phenomenon of blocked growth in summer. 2)Except for the winter season( December-March of the next year),the roots of 1/2-year-old seedlings grew continuously. At the age of 3,the rate of root elongation was slower than at 1/2 years,with root thickening and lateral root growth being the main growth. 3) Analysis of seed kernel dry matter consumption and total sugar concentration showed that the growth of seedlings in their first summer( from early June) was blocked,which was in line with the lowest seed kernel residue and total sugar. 4) In early June,the total sugar concentration of the seed kernel began to fall beneath the total sugar concentration of the leaf,indicating that the seed kernel was no longer the main nutrient donor of the plant. 5) Seedlings aged 2 to 3 years began to grow from the middle of March,and growth slowed down in July,and they did not show the phenomenon of blocked growth. [Conclusions] The ecological strategy of 0-3 years old seedlings of Q. variabilis was to ensure the growth of the main roots,the 1-year-old seedlings have stunted growth in summer,which is closely related to the period of weakest seedling resistance transform( heterotrophic autotrophic transformation),and there was no stagnation in the growth of 2 or 3 year old seedlings.
引文
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[11] CARDILLO E,BERNAL C J. Morphological response and growth of cork oak(Quercus suber L.)seedlings at different shade levels[J]. Forest Ecology and Management,2006,222(1/2/3):296.
[12] SEVILLANO I,SHORT I,GRANT J,et al. Effects of light availability on morphology,growth and biomass allocation of Fagus sylvatica and Quercus robur seedlings[J].Forest Ecology and Management,2016,374:11.
[13] WEINER J. Allocation,plasticity and allometry in plants[J]. Perspectives in Plant Ecology Evolution and Systematics,2004,6(4):207.
[14] LLORET F,CASANOVAS C,PENUELAS J. Seedling survival of Mediterranean shrubland species in relation to root:Shoot ratio,seed size and water and nitrogen use[J]. Functional Ecology,1999.,13(2):210.
[15] HARTMANN H,ADAMS H D,HAMMOND W M,et al. Identifying differences in carbohydrate dynamics of seedlings and mature trees to improve carbon allocation in models for trees and forests[J]. Environmental and Experimental Botany,2018,152:7.
[16] DUMSCHOTT K,RICHTER A,LOESCHER W H,et al. Post photosynthetic carbon partitioning to sugar alcohols and consequences for plant growth[J]. Phytochemistry,2017,144(12):243.
[17] CHAVES I,PASSARINHO J A,CAPITAFO C,et al.Temperature stress effects in Quercus suber leaf metabolism[J]. Journal of Plant Physiology,2011,168(15):1729.
[18] WANG Ao,WANG Xue,TOGNETTI R,et al. Elevation alters carbon and nutrient concentrations and stoichiometry in Quercus aquifolioides in southwestern China[J].Science of the Total Environment,2018,622:1463.
[19] THORNLEY J H M. A model of leaf tissue growth,acelimation and senescence[J]. Annals of Botany,1991,67(3):219.
[20] THORNLEY J H M. A transport-resistance model of forest growth and partitioning[J]. Annals of Botany,1991,68(3):211.
[21]张继祥，魏钦平，陆佩玲，等．美国黑核桃实生苗干物质积累与分配过程的数值模拟[J]．生物数学学报，2004,19(1):109.ZHANG Jixiang,WEI Qinping LU Peiling. Mathematically modeling of ecophysiological process response to environmental factors for American black walnut seedlings(Ⅳ):Dynamic models of the accumulation and distribution for black walnut's dry matter[J]. Journal of Biomathematics,2004,19(1):109.
[22] BAQUEDANO F J,CASTILLO F J. Drought tolerance in the Mediterranean species Quercus coccifera,Quercus ilex,Pinus halepensis,and Juniperus phoenicea[J]. Photosynthetica,2007,45:229.
[23] SREE K S,RAJAM M. Genetic Engineering strategies for biotic stress tolerance in plants[M]∥BAHADUR B,RAJAM M,SAHIJRAM L,et al. Plant Biology and Biotechnology. New Delhi:Springer,2015:611.
[24] TAIZ L,ZEIGER E. Plant physiology[M]. 5th ed. Sunderland,Massachusetts. USA:Sinauer Associates Inc.Publishers,2010,591.
[25]束怀瑞．果树栽培生理学[M]．北京:中国农业出版社，1993,53.SHU Huairui. Physiology of fruit cultivation[M]. Beijing:China Agricultural Press,1993,53.
[26] HOLCH A E. Development of roots and shoots of certain deciduous tree seedlings in different forest sites[J]. Ecology,1931,12(2):259.
[27]刘国军，张希明，吕朝燕，等．不同供水条件下梭梭幼苗生长动态的研究[J]．中国沙漠，2012,32(2):388.LIU Guojun,ZHANG Ximing,LChaoyan,et al. Seedling growth dynamic of Haloxylon ammodendron under different water supply[J]. Journal of Desert Research,2012,32(2):388.
[28] DANNER B T,KNAPP A K. Growth dynamics of oak seedlings(Quercus macrocarpa Michx. and Quercus muhlenbergii Engelm.)from gallery forests:Implications for forest expansion into grasslands[J]. Trees,2001,15(5):271.
[29] VILLAR-SALVADOR P,HEREDIA N,MILLARD P.Remobilization of acorn nitrogen for seedling growth in holm oak(Quercus ilex),cultivated with contrasting nutrient availability[J]. Tree Physiology,2010,30(2):257.
[2] COSTA A,VILLA-HERNANDEZ S,ROJO A P,et al.Can native shrubs facilitate the early establishment of contrasted co-occurring oaks in Mediterranean grazed areas?[J]. Journal of Vegetation Science,2017,28:1047.
[3] ZAVALETA E S,HULVEY K B,FULFROST B. Regional patterns of recruitment success and failure in two endemic California oaks[J]. Diversity and Distributions,2007,13(6):735.
[4] COSTA A,PEREIRA H,MADEIRA M. Analysis of spatial patterns of oak decline in cork oak woodlands in Mediterranean conditions[J]. Annals of Forest Science,2010,67(2):204.
[5] GUGGER P F,PENALOZA-RAMIREZ J M,WRIGHT J W,et al. Whole-transcriptome response to water stress in a California endemic oak,Quercus lobata[J]. Tree Physiology,2017,37(5):632.
[6] PERKINS D,UHL E,BIBER P,et al. Impact of climate trends and drought events on the growth of oaks(Quercus robur L. and Quercus petraea(Matt.)Liebl.)within and beyond their natural range[J]. Forests,2018,9(108):1.
[7]杨舒婷，曲博，李谦盛，等．弗吉尼亚栎幼苗对高温胁迫的生理响应[J]．江西农业大学学报，2015,37(1):90.YANG Shuting,QU Bo,LI Qiansheng,et al. Physiological response of Quercus virginiana seedlings to high temperature stress[J]. Acta Agriculturae Universitatis Jiangxiensis,2015,37(1):90.
[8]程龙霞，施曼，祝遵凌．欧洲鹅耳枥一年生播种苗年生长动态探究[J]．中国野生植物资源，2015,34(5):46.CHENG Longxia,SHI Man,ZHU Zunling. Research on the dynamic growth of one-year-old Carpinus betulus[J].Chinese Wild Plant Resources,2015,34(5):46.
[9]蔡年辉，许玉兰，白青松，等．不同种群高山松1年生播种苗木生长节律及其变异[J]．东北林业大学学报，2013,41(5):11.CAI Nianhui,XU Yulan,BAI Qinsong,et al. Annual seedling growth dynamic rhythm and variation of one-year Pinus densata among populations[J]. Journal of Northeast Forestry University,2013,41(5):11.
[10] CANELLAS I,San MIGUEL A. Biomass of root and shoot systems of Quercus coccifera shrublands in eastern Spain[J]. Annals of Forest Science,2000,57(8):803.
[11] CARDILLO E,BERNAL C J. Morphological response and growth of cork oak(Quercus suber L.)seedlings at different shade levels[J]. Forest Ecology and Management,2006,222(1/2/3):296.
[12] SEVILLANO I,SHORT I,GRANT J,et al. Effects of light availability on morphology,growth and biomass allocation of Fagus sylvatica and Quercus robur seedlings[J].Forest Ecology and Management,2016,374:11.
[13] WEINER J. Allocation,plasticity and allometry in plants[J]. Perspectives in Plant Ecology Evolution and Systematics,2004,6(4):207.
[14] LLORET F,CASANOVAS C,PENUELAS J. Seedling survival of Mediterranean shrubland species in relation to root:Shoot ratio,seed size and water and nitrogen use[J]. Functional Ecology,1999.,13(2):210.
[15] HARTMANN H,ADAMS H D,HAMMOND W M,et al. Identifying differences in carbohydrate dynamics of seedlings and mature trees to improve carbon allocation in models for trees and forests[J]. Environmental and Experimental Botany,2018,152:7.
[16] DUMSCHOTT K,RICHTER A,LOESCHER W H,et al. Post photosynthetic carbon partitioning to sugar alcohols and consequences for plant growth[J]. Phytochemistry,2017,144(12):243.
[17] CHAVES I,PASSARINHO J A,CAPITAFO C,et al.Temperature stress effects in Quercus suber leaf metabolism[J]. Journal of Plant Physiology,2011,168(15):1729.
[18] WANG Ao,WANG Xue,TOGNETTI R,et al. Elevation alters carbon and nutrient concentrations and stoichiometry in Quercus aquifolioides in southwestern China[J].Science of the Total Environment,2018,622:1463.
[19] THORNLEY J H M. A model of leaf tissue growth,acelimation and senescence[J]. Annals of Botany,1991,67(3):219.
[20] THORNLEY J H M. A transport-resistance model of forest growth and partitioning[J]. Annals of Botany,1991,68(3):211.
[21]张继祥，魏钦平，陆佩玲，等．美国黑核桃实生苗干物质积累与分配过程的数值模拟[J]．生物数学学报，2004,19(1):109.ZHANG Jixiang,WEI Qinping LU Peiling. Mathematically modeling of ecophysiological process response to environmental factors for American black walnut seedlings(Ⅳ):Dynamic models of the accumulation and distribution for black walnut's dry matter[J]. Journal of Biomathematics,2004,19(1):109.
[22] BAQUEDANO F J,CASTILLO F J. Drought tolerance in the Mediterranean species Quercus coccifera,Quercus ilex,Pinus halepensis,and Juniperus phoenicea[J]. Photosynthetica,2007,45:229.
[23] SREE K S,RAJAM M. Genetic Engineering strategies for biotic stress tolerance in plants[M]∥BAHADUR B,RAJAM M,SAHIJRAM L,et al. Plant Biology and Biotechnology. New Delhi:Springer,2015:611.
[24] TAIZ L,ZEIGER E. Plant physiology[M]. 5th ed. Sunderland,Massachusetts. USA:Sinauer Associates Inc.Publishers,2010,591.
[25]束怀瑞．果树栽培生理学[M]．北京:中国农业出版社，1993,53.SHU Huairui. Physiology of fruit cultivation[M]. Beijing:China Agricultural Press,1993,53.
[26] HOLCH A E. Development of roots and shoots of certain deciduous tree seedlings in different forest sites[J]. Ecology,1931,12(2):259.
[27]刘国军，张希明，吕朝燕，等．不同供水条件下梭梭幼苗生长动态的研究[J]．中国沙漠，2012,32(2):388.LIU Guojun,ZHANG Ximing,LChaoyan,et al. Seedling growth dynamic of Haloxylon ammodendron under different water supply[J]. Journal of Desert Research,2012,32(2):388.
[28] DANNER B T,KNAPP A K. Growth dynamics of oak seedlings(Quercus macrocarpa Michx. and Quercus muhlenbergii Engelm.)from gallery forests:Implications for forest expansion into grasslands[J]. Trees,2001,15(5):271.
[29] VILLAR-SALVADOR P,HEREDIA N,MILLARD P.Remobilization of acorn nitrogen for seedling growth in holm oak(Quercus ilex),cultivated with contrasting nutrient availability[J]. Tree Physiology,2010,30(2):257.
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