细根对中亚热带森林几种优势树种凋落叶分解的影响
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摘要
为深入理解森林生态系统中进入凋落物层生长的细根对凋落叶分解的影响,通过分解袋控制实验,以多花黑麦草(Lolium multiflorum)根系为研究对象,探讨细根对中亚热带森林中四川山矾、薯豆、香樟和马尾松等4种优势树种单一及其混合凋落叶分解速率的影响.结果显示,进入分解袋中生长的活根生物量因凋落叶性质不同差异极显著(P <0.001),其中薯豆分解袋中的细根在生长高峰期时生物量最大(131.5 mg/袋).凋落物质量和细根存在与否以及二者的交互作用均对分解过程产生显著影响.经过270 d的分解,生长进入分解袋中的细根能一定程度加速凋落叶分解,其中细根对薯豆凋落叶质量损失的相对贡献率最大(57.78%),而对马尾松的贡献率最小(6%),凋落叶初始C/N值显著影响细根对凋落叶质量损失的相对贡献率.同一类型凋落叶在两种根处理条件下,凋落叶表面细菌及真菌群落结构均存在较大差异,有根处理能显著提高细菌群落的多样性及数量,且细根的存在及其吸收作用对3种阔叶树种凋落叶的混合分解产生协同效应.本研究表明进入凋落物层生长的细根生物量与凋落叶初始质量相关,细根通过改变凋落叶表面分解者的群落结构与数量,并且主动调控其生长的养分需求从而加速分解.(图4表7参35)
To understand the influence of fine roots entering the litter layer on the decomposition of litter in the forest ecosystem, a simulation experiment using litterbags was conducted with the root system of Lolium multiflorum as the study object. The effects of living roots on the decomposition rate single and mixed leaf litters of dominant species of mid-subtropical forests were analyzed using the following species: Symplocos setchuensis, Elaeocarpus japonicus, Cinnamomum camphora,and Pinus massoniana. The results indicated that the biomass of the living roots penetrating the litterbags with various leaf litter differed significantly because of the litter quality(P < 0.001), and the fine roots in the litterbags of E. japonicus had the largest biomass at growth peak of the L. multiflorum(131.5 mg/bag). The litter quality, presence or absence of fine roots,and the interaction of the two had significant effects on the decomposition process. After 270 days of decomposition, the fine roots growing in the litter bag accelerated the decomposition of litter to some extent. The fine roots exhibited the largest contribution rate to the loss of the E. japonicus litter(57.78%), whereas the P. massoniana litter(6%) experienced the smallest contribution rate. The initial C/N ratio of litter significantly affected the relative contribution of fine roots to the litter mass loss.Considerable differences in the structure of bacterial and fungal communities on the surface of the leaf litter of the same type under the two root treatments were observed. Treatment with roots remarkably increased the diversity and quantity of bacterial communities, and the presence of fine roots and their absorption had synergistic effects on the mixed decomposition of the litter of three broad-leafed species. In summary, the biomass of fine roots entering the litter layer was related to the initial quality of the litter. The fine roots accelerate decomposition by altering the community structure of the litter surface decomposers and by actively regulating the nutrient requirements for their growth.
To understand the influence of fine roots entering the litter layer on the decomposition of litter in the forest ecosystem, a simulation experiment using litterbags was conducted with the root system of Lolium multiflorum as the study object. The effects of living roots on the decomposition rate single and mixed leaf litters of dominant species of mid-subtropical forests were analyzed using the following species: Symplocos setchuensis, Elaeocarpus japonicus, Cinnamomum camphora,and Pinus massoniana. The results indicated that the biomass of the living roots penetrating the litterbags with various leaf litter differed significantly because of the litter quality(P < 0.001), and the fine roots in the litterbags of E. japonicus had the largest biomass at growth peak of the L. multiflorum(131.5 mg/bag). The litter quality, presence or absence of fine roots,and the interaction of the two had significant effects on the decomposition process. After 270 days of decomposition, the fine roots growing in the litter bag accelerated the decomposition of litter to some extent. The fine roots exhibited the largest contribution rate to the loss of the E. japonicus litter(57.78%), whereas the P. massoniana litter(6%) experienced the smallest contribution rate. The initial C/N ratio of litter significantly affected the relative contribution of fine roots to the litter mass loss.Considerable differences in the structure of bacterial and fungal communities on the surface of the leaf litter of the same type under the two root treatments were observed. Treatment with roots remarkably increased the diversity and quantity of bacterial communities, and the presence of fine roots and their absorption had synergistic effects on the mixed decomposition of the litter of three broad-leafed species. In summary, the biomass of fine roots entering the litter layer was related to the initial quality of the litter. The fine roots accelerate decomposition by altering the community structure of the litter surface decomposers and by actively regulating the nutrient requirements for their growth.
引文
1潘复静,张伟,王克林,何寻阳,梁士楚,韦国富.典型喀斯特峰丛洼地植被群落凋落物C:N:P生态化学计量特征[J].生态学报,2011,31(2):335-343[Pan FJ,Zhang W,Wang KL,He XY,Liang SC,Wei GF.Litter C:N:P ecological stoichiometry character of plant communities in typical Karst Peak-Cluster Depression[J].Acta Ecol Sin,2011,31(2):335-343]
2 Wright SJ,Yavitt JB,Wurzburger N,Turner BL,Tanner EVJ,Sayer EJ,Santiago LS,Kaspari M,Hedin LO,Harms KE,Garcia MN,Corre MD.Potassium,phosphorus,or nitrogen limit root allocation,tree growth,or litter production in a lowland tropical forest[J].Ecology,2011,92(8):1616-1625
3 Wang FC,Fang XM,Ding ZQ,Wan SZ,Chen FS.Effects of understory plant root growth into the litter layer on the leaf litter decomposition of two woody species in a subtropical forest[J].For Ecol Manage,2016,364:39-45
4 Liu R,Huang Z,Luke McCormack M,Zhou X,Wan X,Yu Z,Wang M,Zheng L.Plasticity of fine-root functional traits in the litter layer in response to nitrogen addition in a subtropical forest plantation[J].Plant Soil,2017,415(1-2):317-330
5 Ball BA,Carrillo Y,Molina M.The inf luence of litter composition across the litter-soil interface on mass loss,nitrogen dynamics and the decomposer community[J].Soil Biol Biochem,2014,69(1):71-82
6 Smyth CE,Macey D,Trofymow JA.Long-term litter decay in Canadian forests and the influence of soil microbial community and soil chemistry[J].Soil Biol Biochem,2015,80:251-259
7 Luo D,Cheng R,Shi Z,Wang W.Decomposition of leaves and fine roots in three subtropical plantations in China affected by litter substrate quality and soil microbial community[J].Forests,2017,8(11):412
8 Kong X,Jia Y,Song F,Tian K,Lin H,Bei Z,Jia X,Yao B,Guo P,Tian X.Insight into litter decomposition driven by nutrient demands of symbiosis system through the hypha bridge of arbuscular mycorrhizal fungi[J].Environ Sci Pollut Res,2018,25(6):5369-5378
9 Persson H?,Stadenberg I.Fine root dynamics in a Norway spruce forest(Picea abies(L.)Karst)in eastern Sweden[J].Plant Soil,2010,330(1-2):329-344
10 Brons JK,van Elsas JD.Analysis of bacterial communities in soil by use of denaturing gradient gel electrophoresis and clone libraries,as influenced by different reverse primers[J].Appl Environ Microb,2008,74(9):2717-2727
11 May LA,Smiley B,Schmidt MG.Comparative denaturing gradient gel electrophoresis analysis of fungal communities associated with whole plant corn silage[J].Can J Microbiol,2001,47(9):829-841
12 Lee SH,Lee HJ,Kim SJ,Lee HM,Kang H,Kim YP.Identification of airborne bacterial and fungal community structures in an urban area by T-RFLP analysis and quantitative real-time PCR[J].Sci Total Environ,2010,408(6):1349-1357
13 Wang ZH,Xu WY.Decomposition-rate estimation of leaf litter in karst forests in China based on a mathematical model[J].Plant Soil,2013,367(1-2):563-577
14 González G,Seastedt TR.Soil fauna and plant litter decomposition in tropical and subalpine forests[J].Ecology,2001,82(4):955-964
15陈法霖,张凯,郑华,林学强,欧阳志云,屠乃美.PCR-DGGE技术解析针叶和阔叶凋落物混合分解对土壤微生物群落结构的影响[J].应用与环境生物学报,2011,17(2):145-151[Chen FL,Zhang K,Zheng H,Lin XQ,Quyang ZY,Tu NM.Analyzing the effect of mixed decomposition of conifer and broadleaf litters on soil microbial communities by using PCR-DGGE[J].Chin J Appl Environ Biol,2011,17(2):145-151
16 Bonanomi G,Incerti G,Barile E,Capodilupo M,Antignani V,Mingo A,Lanzotti V,Scala F,Mazzoleni S.Phytotoxicity,not nitrogen immobilization,explains plant litter inhibitory effects:evidence from solid-state 13C NMR spectroscopy[J].New Phytol,2011,191(4):1018-1030
17 Pérez-Corona ME,De Aldana BRV.Allelopathic potential of invasive Ulmus pumila on understory plant species[J].Allelopathy J,2013,32(1):101-112
18 Loydi A,Donath T W,Eckstein R L,Ot te A.Non-native species litter reduces germination and growth of resident forbs and grasses:allelopathic,osmotic or mechanical effects?[J].Biol Invasions,2015,17(2):581-595
19 Xu Z,Zhu J,Wu F,Liu Y,Tan B,Yang W.Effects of litter quality and climate change along an elevational gradient on litter decomposition of subalpine forests,Eastern Tibetan Plateau,China[J].J For Res,2016,27(3):505-511
20査同刚,张志强,孙阁,王高敏,贠小琴,王伊琨,刘艳.凋落物分解主场效应及其土壤生物驱动[J].生态学报,2012,32(24):7991-8000[Zha TG,Zhang ZQ,Sun G,Wang GM,Yun XQ,Wang YK,Liu Y.Homefield advantage of litter decomposition and its soil biological driving mechanism:a review[J].Acta Ecol Sin,2012,32(24):7991-8000]
21 Lopez-Iglesias B,Olmo M,Gallardo A,Villar R.Short-term effects of litter from 21 woody species on plant growth and root development[J].Plant Soil,2014,381(1-2):177-191
22 Berg B.Litter decomposition and organic matter turnover in northern forest soils[J].For Ecol Manage,2000,133(1-2):13-22
23 Zhang T,Luo Y,Chen HYH,Ruan H.Responses of litter decomposition and nutrient release to N addition:a meta-analysis of ter restrial ecosystems[J].Appl Soil Ecol,2018,128:35-42
24 Carrillo Y,Bell C,Koyama A,Canarini A,Boot CM,Wallenstein M,Pendall E.Plant traits,stoichiometry and microbes as drivers of decomposition in the rhizosphere in a temperate grassland[J].J Ecol,2017,105(6):1750-1765
25 Bardgett RD,Bowman WD,Kaufmann R,Schmidt SK.A temporal approach to linking aboveground and belowground ecology[J].Trends Ecol Evol,2005,20(11):634-641
26 de Graaff MA,Classen AT,Castro HF,Schadt CW.Labile soil carbon inputs mediate the soil microbial community composition and plant residue decomposition rates[J].New Phytol,2010,188(4):1055-1064
27 Santonja M,Rancon A,Fromin N,Baldy V,H?ttenschwiler S,Fernandez C,Montès N,Mirleau P.Plant litter diversity increases microbial abundance,fungal diversity,and carbon and nitrogen cycling in a Mediterranean shrubland[J].Soil Biol Biochem,2017,111:124-134
28 Schimel JP,H?ttenschwiler S.Nitrogen transfer between decomposing leaves of different N status[J].Soil Biol Biochem,2007,39(7):1428-1436
29 Chapman SK,Langley JA,Hart SC,Koch GW.Plants actively control nitrogen cycling:uncorking the microbial bottleneck[J].New Phytol,2006,169(1):27-34
30 Kuzyakov Y.Priming effects:Interactions between living and dead organic matter[J].Soil Biol Biochem,2010,42(9):1363-1371
31 Fu S,Cheng W.Rhizosphere priming effects on the decomposition of soil organic matter in C4 and C3 grassland soils[J].Plant Soil,2002,238(2):289-294
32 Craine JM,Morrow C,Fierer N.Microbial nitrogen limitation increases decomposition[J].Ecology,2007,88(8):2105-2113
33 Pragadheesh VS,Saroj A,Yadav A,Chanotiya CS,Alam M,Samad A.Chemical characterization and antifungal activity of Cinnamomum camphora essential oil[J].Ind Crops Prod,2013,49(4):628-633
34陈洪,胡庭兴,王茜,蒋雪,周光良,胡红玲,景建飞.香樟凋落叶分解物对辣椒生长发育的影响[J].西北植物学报,2014,34(12):2525-2534[Chen H,Hu TX,Wang Q,Jiang X,Zhou GL,Hu HL,Jing JF.Effect of decomposing leaf litter of Cinnamomum camphora on growth and development of Capsicum annuum[J]Acta Bot Bor Occid Sin,2014,34(12):2525-2534]
35李仲彬,胡庭兴,李霜,陈洪,胡红玲,卫娇娇,丁伟.香樟凋落叶在土壤中分解初期对凤仙花生长和生理特性的影响[J].应用与环境生物学报,2015,21(3):571-579[Li ZB,Hu TX,Li S,Chen H,Hu HL,Wei JJ,Ding W.Effects of initial decomposing leaf litter of Cinnamomum camphora on the growth and physiology of Impatiens balsamina[J].Chin J Appl Environ Biol,2015,21(3):571-579]
2 Wright SJ,Yavitt JB,Wurzburger N,Turner BL,Tanner EVJ,Sayer EJ,Santiago LS,Kaspari M,Hedin LO,Harms KE,Garcia MN,Corre MD.Potassium,phosphorus,or nitrogen limit root allocation,tree growth,or litter production in a lowland tropical forest[J].Ecology,2011,92(8):1616-1625
3 Wang FC,Fang XM,Ding ZQ,Wan SZ,Chen FS.Effects of understory plant root growth into the litter layer on the leaf litter decomposition of two woody species in a subtropical forest[J].For Ecol Manage,2016,364:39-45
4 Liu R,Huang Z,Luke McCormack M,Zhou X,Wan X,Yu Z,Wang M,Zheng L.Plasticity of fine-root functional traits in the litter layer in response to nitrogen addition in a subtropical forest plantation[J].Plant Soil,2017,415(1-2):317-330
5 Ball BA,Carrillo Y,Molina M.The inf luence of litter composition across the litter-soil interface on mass loss,nitrogen dynamics and the decomposer community[J].Soil Biol Biochem,2014,69(1):71-82
6 Smyth CE,Macey D,Trofymow JA.Long-term litter decay in Canadian forests and the influence of soil microbial community and soil chemistry[J].Soil Biol Biochem,2015,80:251-259
7 Luo D,Cheng R,Shi Z,Wang W.Decomposition of leaves and fine roots in three subtropical plantations in China affected by litter substrate quality and soil microbial community[J].Forests,2017,8(11):412
8 Kong X,Jia Y,Song F,Tian K,Lin H,Bei Z,Jia X,Yao B,Guo P,Tian X.Insight into litter decomposition driven by nutrient demands of symbiosis system through the hypha bridge of arbuscular mycorrhizal fungi[J].Environ Sci Pollut Res,2018,25(6):5369-5378
9 Persson H?,Stadenberg I.Fine root dynamics in a Norway spruce forest(Picea abies(L.)Karst)in eastern Sweden[J].Plant Soil,2010,330(1-2):329-344
10 Brons JK,van Elsas JD.Analysis of bacterial communities in soil by use of denaturing gradient gel electrophoresis and clone libraries,as influenced by different reverse primers[J].Appl Environ Microb,2008,74(9):2717-2727
11 May LA,Smiley B,Schmidt MG.Comparative denaturing gradient gel electrophoresis analysis of fungal communities associated with whole plant corn silage[J].Can J Microbiol,2001,47(9):829-841
12 Lee SH,Lee HJ,Kim SJ,Lee HM,Kang H,Kim YP.Identification of airborne bacterial and fungal community structures in an urban area by T-RFLP analysis and quantitative real-time PCR[J].Sci Total Environ,2010,408(6):1349-1357
13 Wang ZH,Xu WY.Decomposition-rate estimation of leaf litter in karst forests in China based on a mathematical model[J].Plant Soil,2013,367(1-2):563-577
14 González G,Seastedt TR.Soil fauna and plant litter decomposition in tropical and subalpine forests[J].Ecology,2001,82(4):955-964
15陈法霖,张凯,郑华,林学强,欧阳志云,屠乃美.PCR-DGGE技术解析针叶和阔叶凋落物混合分解对土壤微生物群落结构的影响[J].应用与环境生物学报,2011,17(2):145-151[Chen FL,Zhang K,Zheng H,Lin XQ,Quyang ZY,Tu NM.Analyzing the effect of mixed decomposition of conifer and broadleaf litters on soil microbial communities by using PCR-DGGE[J].Chin J Appl Environ Biol,2011,17(2):145-151
16 Bonanomi G,Incerti G,Barile E,Capodilupo M,Antignani V,Mingo A,Lanzotti V,Scala F,Mazzoleni S.Phytotoxicity,not nitrogen immobilization,explains plant litter inhibitory effects:evidence from solid-state 13C NMR spectroscopy[J].New Phytol,2011,191(4):1018-1030
17 Pérez-Corona ME,De Aldana BRV.Allelopathic potential of invasive Ulmus pumila on understory plant species[J].Allelopathy J,2013,32(1):101-112
18 Loydi A,Donath T W,Eckstein R L,Ot te A.Non-native species litter reduces germination and growth of resident forbs and grasses:allelopathic,osmotic or mechanical effects?[J].Biol Invasions,2015,17(2):581-595
19 Xu Z,Zhu J,Wu F,Liu Y,Tan B,Yang W.Effects of litter quality and climate change along an elevational gradient on litter decomposition of subalpine forests,Eastern Tibetan Plateau,China[J].J For Res,2016,27(3):505-511
20査同刚,张志强,孙阁,王高敏,贠小琴,王伊琨,刘艳.凋落物分解主场效应及其土壤生物驱动[J].生态学报,2012,32(24):7991-8000[Zha TG,Zhang ZQ,Sun G,Wang GM,Yun XQ,Wang YK,Liu Y.Homefield advantage of litter decomposition and its soil biological driving mechanism:a review[J].Acta Ecol Sin,2012,32(24):7991-8000]
21 Lopez-Iglesias B,Olmo M,Gallardo A,Villar R.Short-term effects of litter from 21 woody species on plant growth and root development[J].Plant Soil,2014,381(1-2):177-191
22 Berg B.Litter decomposition and organic matter turnover in northern forest soils[J].For Ecol Manage,2000,133(1-2):13-22
23 Zhang T,Luo Y,Chen HYH,Ruan H.Responses of litter decomposition and nutrient release to N addition:a meta-analysis of ter restrial ecosystems[J].Appl Soil Ecol,2018,128:35-42
24 Carrillo Y,Bell C,Koyama A,Canarini A,Boot CM,Wallenstein M,Pendall E.Plant traits,stoichiometry and microbes as drivers of decomposition in the rhizosphere in a temperate grassland[J].J Ecol,2017,105(6):1750-1765
25 Bardgett RD,Bowman WD,Kaufmann R,Schmidt SK.A temporal approach to linking aboveground and belowground ecology[J].Trends Ecol Evol,2005,20(11):634-641
26 de Graaff MA,Classen AT,Castro HF,Schadt CW.Labile soil carbon inputs mediate the soil microbial community composition and plant residue decomposition rates[J].New Phytol,2010,188(4):1055-1064
27 Santonja M,Rancon A,Fromin N,Baldy V,H?ttenschwiler S,Fernandez C,Montès N,Mirleau P.Plant litter diversity increases microbial abundance,fungal diversity,and carbon and nitrogen cycling in a Mediterranean shrubland[J].Soil Biol Biochem,2017,111:124-134
28 Schimel JP,H?ttenschwiler S.Nitrogen transfer between decomposing leaves of different N status[J].Soil Biol Biochem,2007,39(7):1428-1436
29 Chapman SK,Langley JA,Hart SC,Koch GW.Plants actively control nitrogen cycling:uncorking the microbial bottleneck[J].New Phytol,2006,169(1):27-34
30 Kuzyakov Y.Priming effects:Interactions between living and dead organic matter[J].Soil Biol Biochem,2010,42(9):1363-1371
31 Fu S,Cheng W.Rhizosphere priming effects on the decomposition of soil organic matter in C4 and C3 grassland soils[J].Plant Soil,2002,238(2):289-294
32 Craine JM,Morrow C,Fierer N.Microbial nitrogen limitation increases decomposition[J].Ecology,2007,88(8):2105-2113
33 Pragadheesh VS,Saroj A,Yadav A,Chanotiya CS,Alam M,Samad A.Chemical characterization and antifungal activity of Cinnamomum camphora essential oil[J].Ind Crops Prod,2013,49(4):628-633
34陈洪,胡庭兴,王茜,蒋雪,周光良,胡红玲,景建飞.香樟凋落叶分解物对辣椒生长发育的影响[J].西北植物学报,2014,34(12):2525-2534[Chen H,Hu TX,Wang Q,Jiang X,Zhou GL,Hu HL,Jing JF.Effect of decomposing leaf litter of Cinnamomum camphora on growth and development of Capsicum annuum[J]Acta Bot Bor Occid Sin,2014,34(12):2525-2534]
35李仲彬,胡庭兴,李霜,陈洪,胡红玲,卫娇娇,丁伟.香樟凋落叶在土壤中分解初期对凤仙花生长和生理特性的影响[J].应用与环境生物学报,2015,21(3):571-579[Li ZB,Hu TX,Li S,Chen H,Hu HL,Wei JJ,Ding W.Effects of initial decomposing leaf litter of Cinnamomum camphora on the growth and physiology of Impatiens balsamina[J].Chin J Appl Environ Biol,2015,21(3):571-579]