灰杨克隆繁殖、自身养分与土壤养分季节变化及其耦合关系的林缘——林内差异
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摘要
阐明灰杨(Populus pruinosa Schrenk)克隆特征变化及受外界环境影响,是增强灰杨林生态服务功能的基础。本研究在中国新疆塔里木河上游灰杨河岸林,沿垂直河道方向在林缘(距河道200 m)、林内(距河道400 m)生境设置样方,于2014年4～10月,每隔20天调查两生境不定芽、未出土和出土克隆分株数量,测定横走侧根及土壤养分含量。结果表明:(1)横走侧根全氮、全磷、全钾、有机碳含量的变化趋势与不定芽和未出土分株数在4～6月逐渐增加、7～10月逐渐减少变化趋势基本一致,与土壤有机质、碱解氮、速效磷及速效钾含量在4～6月逐渐升高或保持最高水平,在7～10月逐渐降低的变化趋势同步;横走侧根碳氮比在4～6月初逐渐降低,随后呈现增大又减小的趋势;氮磷比在4～6月逐渐减小,7月显著减小后逐渐增大(林缘)或基本保持不变(林内)。(2)不论是林缘还是林内,不定芽数量受到横走侧根全氮和氮磷比的影响,未出土分株数受到横走侧根全氮、有机碳含量、碳氮比的影响;林缘不定芽数量与土壤有机质、碱解氮及速效钾含量相关紧密,未出土分株数与土壤碱解氮和速效磷显著相关,土壤养分也与横走侧根全氮、碳氮比、氮磷比紧密相关,而在林内受到土壤养分影响的主要是横走侧根全氮和氮磷比。(3)林内的不定芽、未出土及出土克隆分株数量显著低于林缘,土壤养分含量通过影响横走侧根养分含量而影响不定芽、未出土分株、出土分株数量。灰杨是杨属为数不多的国家保护植物,具有重要的生态服务功能,我们的结果对基于克隆繁殖的灰杨林保护策略提供基础数据支持。
Propagation features elucidation is important for improving ecological service of poplar forests in arid regions. We studied the relationship between the occurrence of clonal organs of Populus pruinosa Schrenk and soil nutrient content during the clonal propagation,and their variations affected by the forest edge. The sample plot was set up vertical to channel of Tarim River upper reaches in Xinjiang,China,in the riparian forest (400 m away from the river channel) and forest edge (200 m away from the river channel) separately. The number of adventitious buds,underground ramets and unearthed ramets in two habitats was investigated every 20 days from April to October in 2014,and the nutrient contents of lateral roots and soil were also measured. The results show that: (1) The content of total nitrogen,total phosphorus,total potassium and organic carbon of lateral roots increased gradually from April to June,and decreased gradually from July to October,which is consistent with the trend of number variation of the adventitious buds and underground ramets during this period,and also show a spontaneously trend with gradually increased soil organic matter content,available nitrogen content,available phosphorus content and available potassium content or maintain at a high level from April to June,and decreased gradually thereafter from July to October. Whereas,the C/N ratio of lateral roots decreased gradually from April to June,after a slightly increasing and decreasing again,the N/P ratio also decreased gradually from April to June,but show an increasing trend in the forest edge or remain unchanging in the forest after a significant decrease in July. (2) The number of adventitious buds was affected by total nitrogen and nitrogenphosphorus ratio of lateral roots,the number of underground ramets was affected by the content of total nitrogen,organic carbon and carbon-nitrogen ratio of lateral roots both in the forest and forest edge. In the forest edge,the number of adventitious buds was close related to the content of soil organic matter,alkali-hydrolyzable nitrogen and available potassium,and there was also a significant correlation between the underground ramets and the content of soil alkali-hydrolyzable nitrogen and available phosphorus,as well as a significant correlation between the soil nutrients and the total nitrogen,C/N ratio and N/P ratio of lateral roots,while,the main factors affected by soil nutrients in the forest were total nitrogen and N/P ratio of lateral roots. (3) The number of adventitious buds,underground ramets and unearthed ramets in the forest was significantly lower than that in the forest edge. Soil nutrient content affected the number of adventitious buds,underground ramets and unearthed ramets by influencing the nutrient content of lateral roots. P. pruinosa is one of the few national protected plants in P. pruinosa forest play an important role on ecological services in local area,our results provide a basic data support for the conservation strategy formulation based on the clonal propagation of P.pruinosa forest.
Propagation features elucidation is important for improving ecological service of poplar forests in arid regions. We studied the relationship between the occurrence of clonal organs of Populus pruinosa Schrenk and soil nutrient content during the clonal propagation,and their variations affected by the forest edge. The sample plot was set up vertical to channel of Tarim River upper reaches in Xinjiang,China,in the riparian forest (400 m away from the river channel) and forest edge (200 m away from the river channel) separately. The number of adventitious buds,underground ramets and unearthed ramets in two habitats was investigated every 20 days from April to October in 2014,and the nutrient contents of lateral roots and soil were also measured. The results show that: (1) The content of total nitrogen,total phosphorus,total potassium and organic carbon of lateral roots increased gradually from April to June,and decreased gradually from July to October,which is consistent with the trend of number variation of the adventitious buds and underground ramets during this period,and also show a spontaneously trend with gradually increased soil organic matter content,available nitrogen content,available phosphorus content and available potassium content or maintain at a high level from April to June,and decreased gradually thereafter from July to October. Whereas,the C/N ratio of lateral roots decreased gradually from April to June,after a slightly increasing and decreasing again,the N/P ratio also decreased gradually from April to June,but show an increasing trend in the forest edge or remain unchanging in the forest after a significant decrease in July. (2) The number of adventitious buds was affected by total nitrogen and nitrogenphosphorus ratio of lateral roots,the number of underground ramets was affected by the content of total nitrogen,organic carbon and carbon-nitrogen ratio of lateral roots both in the forest and forest edge. In the forest edge,the number of adventitious buds was close related to the content of soil organic matter,alkali-hydrolyzable nitrogen and available potassium,and there was also a significant correlation between the underground ramets and the content of soil alkali-hydrolyzable nitrogen and available phosphorus,as well as a significant correlation between the soil nutrients and the total nitrogen,C/N ratio and N/P ratio of lateral roots,while,the main factors affected by soil nutrients in the forest were total nitrogen and N/P ratio of lateral roots. (3) The number of adventitious buds,underground ramets and unearthed ramets in the forest was significantly lower than that in the forest edge. Soil nutrient content affected the number of adventitious buds,underground ramets and unearthed ramets by influencing the nutrient content of lateral roots. P. pruinosa is one of the few national protected plants in P. pruinosa forest play an important role on ecological services in local area,our results provide a basic data support for the conservation strategy formulation based on the clonal propagation of P.pruinosa forest.
引文
1.李志军,刘建平,于军,等.胡杨、灰叶胡杨生物生态学特性调查[J].西北植物学报,2003,23(7):1292-1296.Li Z J,Liu J P,Yu J,et al.Investigation on the characteristics of biology and ecology of Populus euphratica and Populus pruinosa[J].Acta Botanica Boreali-occidentalia Sinica,2003,23(7):1292-1296.
2.郑亚琼,周正立,李志军.灰叶胡杨横走侧根空间分布与克隆繁殖的关系[J].生态学杂志,2013,32(10):2641-2646.Zheng Y Q,Zhou Z L,Li Z J.Spatial distribution of horizontal lateral roots of Populus pruinosa Schrenk in relation to clonal reproduction[J].Chinese Journal of Ecology,2013,32(10):2641-2646.
3.郑亚琼,张肖,梁继业,等.濒危物种胡杨和灰叶胡杨的克隆生长特征[J].生态学报,2016,36(5):1331-1341.Zheng Y Q,Zhang X,Liang J Y,et al.Clonal growth characteristics of the endangered species Populus euphratica Oliv and Populus pruinosa Schrenk[J].Acta Ecologica Sinica,2016,36(5):1331-1341.
4.李志军,焦培培,周正立,等.灰叶胡杨根蘖繁殖的形态解剖学特征[J].植物学报,2012,47(2):133-140.Li Z J,Jiao P P,Zhou Z L,et al.Morphological and Anatomical features of root sucker propagation of Populus pruinosa[J].Chinese Bulletin of Botany,2012,47(2):133-140.
5.Zheng Y Q,Jiao P P,Zhao Z S,et al.Clonal growth of Populus pruinosa Schrenk and its role in the regeneration of riparian forests[J].Ecological Engineering,2016,94,380-392.
6.赵正帅,郑亚琼,梁继业,等.塔里木河流域胡杨和灰叶胡杨克隆分株空间分布格局[J].应用生态学报,2016,27(2):403-411.Zhao Z S,Zheng Y Q,Liang J Y,et al.Spatial distribution pattern of Populus euphratica and P.pruinosa clonal ramets in Tarim River Basin,China[J].Chinese Journal of Applied Ecology,2016,27(2):403-411.
7.王文杰,祖元刚,杨逢建,等.边缘效应带促进红松生长的光合生理生态学研究[J].生态学报,2003,23(11):2318-2326.Wang W J,Zu Y G,Yang F J,et al.Photosynthetic ecophysiological study on the growth of korean pine(Pinus koraiensis)afforested by the edge-effect belt method[J].Acta Ecologica Sinica,2003,23(11):2318-2326.
8.祖元刚,王文杰,王慧梅.边缘效应带和保留带内红松幼林水分生态的差异[J].植物生态学报,2002,26(5):613-620.Zu Y G,Wang W J,Wang H M.Differences in water ecology of a young Pinus koraiensis stand grown in edge-effect belts and reserved belts[J].Chinese Journal of Plant Ecology,2002,26(5):613-620.
9.Wang W J,Yan X F,Shi F C,et al.A trial to accelerate afforestation of Korean pine forests using a strip-cutting method for deciduous broad-leaved secondary forests in Northeastern China-an ecophysiological approach[J].Eurasian Journal of Forest Research,2001,3:27-48.
10.魏晨辉,沈光,裴忠雪,等.不同植物种植对松嫩平原盐碱地土壤理化性质与细根生长的影响[J].植物研究,2015,35(5):759-764.Wei C H,Shen G,Pei Z X,et al.Effects of different plants cultivation on soil physical-chemical properties and fine root growth in saline-alkaline soil in songnen plain,Northeastern China[J].Bulletin of Botanical Research,2015,35(5):759-764.
11.路嘉丽,沈光,王琼,等.树种差异对哈尔滨市土壤理化性质影响及造林启示[J].植物研究,2016,36(4):549-555.Lu J L,Shen G,Wang Q,et al.Effect of urban tree species on soil physicochemical properties in Harbin,Northeastern China,and afforestation implications[J].Bulletin of Botanical Research,2016,36(4):549-555.
12.任蔓莉,魏晨辉,裴忠雪,等.松嫩平原沙土区不同植被类型对土壤相关指标的影响[J].植物研究,2015,35(5):765-771.Ren M L,Wei C H,Pei Z X,et al.Influences of different vegetation types on soil parameters in degraded sandy lands of Songnen Plain[J].Bulletin of Botanical Research,2015,35(5):765-771.
13.Tomlinson K W,O’connor T G.Control of tiller recruitment in bunchgrasses:uniting Physiology and ecology[J].Functional Ecology,2004,18(4):489-496.
14.Koerselman W,Meuleman A F M.The vegetation N∶P ratio:a new tool to detect the nature of nutrient limitation[J].Journal of Applied Ecology,1996,33(6):1441-1450.
15.Güsewell S,Koerselman W,Verhoeven J T A.Biomass N∶P ratios as indicators of nutrient limitation for Plant Populations in wetlands[J].Ecological Applications,2003,13(2):372-384.
16.Güsewell S.N∶P ratios in terrestrial plants:variation and functional significance[J].New Phytologist,2004,164(2):243-266.
17.Güsewell S.High nitrogen:Phosphorus ratios reduce nutrient retention and second-year growth of wetland sedges[J].New Phytologist,2005,166(2):537-550.
18.Verhoeven J T A,Koerselman W,Meuleman A F M.Nitrogen-or Phosphorus-limited growth in herbaceous,wet vegetation:relations with atmospheric inputs and management regimes[J].Trends in Ecology&Evolution,1996,11(12):494-497.
19.Ikegami M,Whigham D F,Werger M J A.Responses of rhizome length and ramet production to resource availability in the clonal sedge Scirpus olneyi A.Gray[J].Plant E-cology,2007,189(2):247-259.
20.Roiloa S R,Retuerto R.Small-scale heterogeneity in soil quality influences photosynthetic efficiency and habitat selection in a clonal plant[J].Annals of Botany,2006,98(5):1043-1052.
21.Hodge A.The plastic plant:root responses to heterogeneous supplies of nutrients[J].New Phytol,2004,62:9-24.
2.郑亚琼,周正立,李志军.灰叶胡杨横走侧根空间分布与克隆繁殖的关系[J].生态学杂志,2013,32(10):2641-2646.Zheng Y Q,Zhou Z L,Li Z J.Spatial distribution of horizontal lateral roots of Populus pruinosa Schrenk in relation to clonal reproduction[J].Chinese Journal of Ecology,2013,32(10):2641-2646.
3.郑亚琼,张肖,梁继业,等.濒危物种胡杨和灰叶胡杨的克隆生长特征[J].生态学报,2016,36(5):1331-1341.Zheng Y Q,Zhang X,Liang J Y,et al.Clonal growth characteristics of the endangered species Populus euphratica Oliv and Populus pruinosa Schrenk[J].Acta Ecologica Sinica,2016,36(5):1331-1341.
4.李志军,焦培培,周正立,等.灰叶胡杨根蘖繁殖的形态解剖学特征[J].植物学报,2012,47(2):133-140.Li Z J,Jiao P P,Zhou Z L,et al.Morphological and Anatomical features of root sucker propagation of Populus pruinosa[J].Chinese Bulletin of Botany,2012,47(2):133-140.
5.Zheng Y Q,Jiao P P,Zhao Z S,et al.Clonal growth of Populus pruinosa Schrenk and its role in the regeneration of riparian forests[J].Ecological Engineering,2016,94,380-392.
6.赵正帅,郑亚琼,梁继业,等.塔里木河流域胡杨和灰叶胡杨克隆分株空间分布格局[J].应用生态学报,2016,27(2):403-411.Zhao Z S,Zheng Y Q,Liang J Y,et al.Spatial distribution pattern of Populus euphratica and P.pruinosa clonal ramets in Tarim River Basin,China[J].Chinese Journal of Applied Ecology,2016,27(2):403-411.
7.王文杰,祖元刚,杨逢建,等.边缘效应带促进红松生长的光合生理生态学研究[J].生态学报,2003,23(11):2318-2326.Wang W J,Zu Y G,Yang F J,et al.Photosynthetic ecophysiological study on the growth of korean pine(Pinus koraiensis)afforested by the edge-effect belt method[J].Acta Ecologica Sinica,2003,23(11):2318-2326.
8.祖元刚,王文杰,王慧梅.边缘效应带和保留带内红松幼林水分生态的差异[J].植物生态学报,2002,26(5):613-620.Zu Y G,Wang W J,Wang H M.Differences in water ecology of a young Pinus koraiensis stand grown in edge-effect belts and reserved belts[J].Chinese Journal of Plant Ecology,2002,26(5):613-620.
9.Wang W J,Yan X F,Shi F C,et al.A trial to accelerate afforestation of Korean pine forests using a strip-cutting method for deciduous broad-leaved secondary forests in Northeastern China-an ecophysiological approach[J].Eurasian Journal of Forest Research,2001,3:27-48.
10.魏晨辉,沈光,裴忠雪,等.不同植物种植对松嫩平原盐碱地土壤理化性质与细根生长的影响[J].植物研究,2015,35(5):759-764.Wei C H,Shen G,Pei Z X,et al.Effects of different plants cultivation on soil physical-chemical properties and fine root growth in saline-alkaline soil in songnen plain,Northeastern China[J].Bulletin of Botanical Research,2015,35(5):759-764.
11.路嘉丽,沈光,王琼,等.树种差异对哈尔滨市土壤理化性质影响及造林启示[J].植物研究,2016,36(4):549-555.Lu J L,Shen G,Wang Q,et al.Effect of urban tree species on soil physicochemical properties in Harbin,Northeastern China,and afforestation implications[J].Bulletin of Botanical Research,2016,36(4):549-555.
12.任蔓莉,魏晨辉,裴忠雪,等.松嫩平原沙土区不同植被类型对土壤相关指标的影响[J].植物研究,2015,35(5):765-771.Ren M L,Wei C H,Pei Z X,et al.Influences of different vegetation types on soil parameters in degraded sandy lands of Songnen Plain[J].Bulletin of Botanical Research,2015,35(5):765-771.
13.Tomlinson K W,O’connor T G.Control of tiller recruitment in bunchgrasses:uniting Physiology and ecology[J].Functional Ecology,2004,18(4):489-496.
14.Koerselman W,Meuleman A F M.The vegetation N∶P ratio:a new tool to detect the nature of nutrient limitation[J].Journal of Applied Ecology,1996,33(6):1441-1450.
15.Güsewell S,Koerselman W,Verhoeven J T A.Biomass N∶P ratios as indicators of nutrient limitation for Plant Populations in wetlands[J].Ecological Applications,2003,13(2):372-384.
16.Güsewell S.N∶P ratios in terrestrial plants:variation and functional significance[J].New Phytologist,2004,164(2):243-266.
17.Güsewell S.High nitrogen:Phosphorus ratios reduce nutrient retention and second-year growth of wetland sedges[J].New Phytologist,2005,166(2):537-550.
18.Verhoeven J T A,Koerselman W,Meuleman A F M.Nitrogen-or Phosphorus-limited growth in herbaceous,wet vegetation:relations with atmospheric inputs and management regimes[J].Trends in Ecology&Evolution,1996,11(12):494-497.
19.Ikegami M,Whigham D F,Werger M J A.Responses of rhizome length and ramet production to resource availability in the clonal sedge Scirpus olneyi A.Gray[J].Plant E-cology,2007,189(2):247-259.
20.Roiloa S R,Retuerto R.Small-scale heterogeneity in soil quality influences photosynthetic efficiency and habitat selection in a clonal plant[J].Annals of Botany,2006,98(5):1043-1052.
21.Hodge A.The plastic plant:root responses to heterogeneous supplies of nutrients[J].New Phytol,2004,62:9-24.
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