热带次生林两种林下植物叶片生理特性对氮磷添加的响应
|
摘要
为探究热带次生林林下植物叶片对长期氮(N)、磷(P)添加的响应,阐明热带森林植物对土壤低磷环境适应的生理生态机制,基于华南热带次生林长达9年的氮、磷添加实验,测定两种分布较广的林下乡土树种大青(Clerodendrum cyrtophyllum)和紫玉盘(Uvaria microcarpa)的叶片基本属性、叶绿素含量及氮磷元素含量等指标.结果显示:(1)两种植物叶片的基本属性及光合色素存在显著的种间差异(P <0.05);氮和磷添加提高了大青叶片的比叶面积(35%-38%),对两个树种叶片的叶长、叶宽、叶面积等无显著影响;(2)氮添加显著影响两种植物叶片叶绿素a和总叶绿素含量(P=0.036和P=0.043),而磷添加对光合色素含量无显著影响;(3)对照处理下,大青和紫玉盘叶片的N:P分别为28.71与21.75,大于16,表明两个树种仍受到较强的磷限制;磷添加显著提高了两种植物叶片的磷含量,并降低N:P值;(4)相关性分析表明叶片基本属性、光合色素与氮磷元素含量三者之间的相关性关系较弱.综上,热带森林植物叶片氮和磷元素含量对氮磷添加的响应较敏感,叶绿素a和总叶绿素对氮添加的响应较敏感,而其他属性对氮磷添加的响应较小;因此,在热带森林土壤磷长期缺乏的情况下,植物叶片的生理功能对环境变化具有较好的适应性,其蕴含的生物学机制仍需要进一步的探讨.(图3表2参31)
The goal of this study was to investigate the response and adaptation of plant leaves to long-term N and P addition,clarifying the physiological and ecological mechanisms of tropical forest plants to lower soil P availability in southern China.Based on an N and P addition experiment in the secondary tropical forest of southern China, the foliar traits, photosynthetic pigment, and N and P concentrations of two widely distributed tree species, Clerodendrum cyrtophyllum and Uvaria microcarpa were measured. The results obtained were as follows: 1) Inter-species differences were noted in foliar trait,photosynthetic pigment, and N and P concentrations(P < 0.05)—N and P addition increased the specific leaf area(SLA) of C. cyrtophyllum(35%–38%), whereas there was no significant effect on leaf length, width, or area for the two species; 2) N addition significantly influenced the chl a and chl(a + b) concentrations(P = 0.036 and P = 0.043, respectively), whereas P addition had no significant effects on photosynthetic pigment levels; 3) In the CK treatment, the N:P ratios of C. cyrtophyllum and U. microcarpa were 28.71 and 21.75, respectively, and both were higher than 16, indicating that the two tree species were strongly limited by P availability—P addition significantly increased foliar P concentrations and decreased N:P ratios of the two species; 4) Relatively weaker correlations were noted among foliar traits, photosynthetic pigment, and N and P concentrations.Thus, the foliar N and P concentrations were sensitive to N and P addition, and chl a and chl(a + b) were also sensitive to N addition, whereas other foliar traits were slowly responsive to N and P addition in this tropical forest. With the long-term lower soil availability, tropical forest plant leaves are well adaptive to environmental change, although the underlying biological mechanism is yet to be elucidated.
The goal of this study was to investigate the response and adaptation of plant leaves to long-term N and P addition,clarifying the physiological and ecological mechanisms of tropical forest plants to lower soil P availability in southern China.Based on an N and P addition experiment in the secondary tropical forest of southern China, the foliar traits, photosynthetic pigment, and N and P concentrations of two widely distributed tree species, Clerodendrum cyrtophyllum and Uvaria microcarpa were measured. The results obtained were as follows: 1) Inter-species differences were noted in foliar trait,photosynthetic pigment, and N and P concentrations(P < 0.05)—N and P addition increased the specific leaf area(SLA) of C. cyrtophyllum(35%–38%), whereas there was no significant effect on leaf length, width, or area for the two species; 2) N addition significantly influenced the chl a and chl(a + b) concentrations(P = 0.036 and P = 0.043, respectively), whereas P addition had no significant effects on photosynthetic pigment levels; 3) In the CK treatment, the N:P ratios of C. cyrtophyllum and U. microcarpa were 28.71 and 21.75, respectively, and both were higher than 16, indicating that the two tree species were strongly limited by P availability—P addition significantly increased foliar P concentrations and decreased N:P ratios of the two species; 4) Relatively weaker correlations were noted among foliar traits, photosynthetic pigment, and N and P concentrations.Thus, the foliar N and P concentrations were sensitive to N and P addition, and chl a and chl(a + b) were also sensitive to N addition, whereas other foliar traits were slowly responsive to N and P addition in this tropical forest. With the long-term lower soil availability, tropical forest plant leaves are well adaptive to environmental change, although the underlying biological mechanism is yet to be elucidated.
引文
1 Selene B,Jurgen H.Functional traits determine tree growth and ecosystem productivity of a tropical montane forest:insights from a longterm nutrient manipulation experiment[J].Glob Change Biol,2018,24:399-409
2 Alvarez-Clare S,Mack MC,Brooks M.A direct test of nitrogen and phosphorus limitation to net primary productivity in a lowland tropical wet forest[J].Ecology,2013,94:1540-1551
3 Cernusak,Winter K,Turner BL.Leaf nitrogen to phosphorus ratios of tropical trees:experimental assessment of physiological and environmental controls[J].New Phyt,2010,185:770-779
4 Ostertag R.Foliar nitrogen and phosphorus accumulation responses after fertilization:an example from nutrient-limited Hawaiian forests[J].Plant Soil,2010,334:85-98
5 Tanner EVJ,Kapos V,Freskos S,Healey JR,Theobald AM.Nitrogen and phosphorus fertilization of Jamaican montane forest trees[J].J Trop Ecol,1990,6:231-238
6 Turner BL,Brenes-Arguedas T,Condit R.Pervasive phosphorus limitation of tree species but not communities in tropical forests[J].Nature,2018,555:367
7 Tanner EVJ,Vitousek PM,Cuevas E.Experimental investigation of nutrient limitation of forest growth on wet tropical mountains[J].Ecology,1998,79:10-22
8 Li Y,Niu SL,Yu GR.Aggravated phosphorus limitation on biomass production under increasing nitrogen loading:a meta-analysis[J].Glob Change Biol,2016,22:934-943
9 Santiago LS,Wright SJ,Harms KE,Yavitt JB,Korine C,Garcia MN,Turner BL.Tropical tree seedling growth responses to nitrogen,phosphorus and potassium addition[J].J Ecol,2012,100:309-316
10 Mayor JR,Wright SJ,Turner BL,Austin A.Species-specific responses of foliar nutrients to long-term nitrogen and phosphorus additions in a lowland tropical forest[J].J Ecol,2014,102:36-44
11 Mo QF,Zou B,Li YW,Chen Y,Zhang WX,Mao R,Ding YZ,Wang J,Lu XK,Li XB,Tang JW,Li ZA,Wang FM.Response of plant nutrient stoichiometry to fertilization varied with plant tissues in a tropical forest[J].Sci Reports,2015,5:14605
12莫其锋,陈瑶,王法明,邹碧,李应文,余世钦,李晓波,李志安.华南热带森林两种林下植物氮磷比对氮磷添加的响应[J].应用与环境生物学报,2015,21(5):919-925[Mo QF,Chen Y,Wang FM,Zou B,Li YW,Yu SQ,LI XB,Li ZA.Nitrogen to phosphorus ratios of two understory plant species in response to nitrogen and phosphorus addition in tropical forest of southern China[J].Chin J Appl Environ Biol,2015,21(5):919-925]
13 Zhu FF,Lu XK,Mo JM.Phosphorus limitation on photosynthesis of two dominant understory species in a lowland tropical forest[J].J Plant Ecol,2014,7:526-534
14 Mao QG,Lu XK,Wang C,Zhou KJ,Mo JM.Responses of understory plant physiological traits to a decade of nitrogen addition in a tropical reforested ecosystem[J].For Ecol Ma nage,2017,401:65-74
15 Mao QG,Lu XK,Mo H,Gundersen P,Mo JM.Effects of simulated N deposition on foliar nutrient status,N metabolism and photosynthetic capacity of three dominant understory plant species in a mature tropical forest[J].Sci Total Environ,2018,610-611:555-562
16 Wang FM,Ding YZ,Sayer EJ,Li QL,Zou B,Mo QF,Li YW,Lu XL,Tang JW,Zhu WX,Li ZA.Tropical forest restoration:fast resilience of plant biomass contrasts with slow recovery of stable soil C stocks[J].Func Ecol,2017,31:2344-2355
17 Li J,Li ZA,Wang FM,Zou B,Chen Y,Zhao J,Mo QF,Li YW,Li XB,Xia HP.Effects of nitrogen and phosphorus addition on soil microbial community in a secondary tropical forest of China[J].Biol Fer Soil,2015,51:207-215
18 Chen Y,Sayer EJ,Li ZA,Mo QF,Li YW,Ding YZ,Wang J,Lu XK,Tang JW,Wang FM.Nutrient limitation of woody debris decomposition in a tropical forest:contrasting effects of N and P addition[J].Func Ecol,2016,30:295-304
19 Oberson A.,Friesen DK,Morel C,Tiessen H.Determination of phosphorus released by chloroform fumigation from microbial biomass in high P sorbing tropical soils[J].Soil Biol Biochem,1997,29:1579-1583
20 Lu XK,Mao QG,Gilliam FS,Luo Y,Mo JM.Nitrogen deposition contributes to soil acidification in tropical ecosystems[J].Glob Change Biol,2014,20:3790-3801
21 Bremner JM,Mulvaney CS.Nitrogen-total//Page AL,Miller RH,Keeney DR.Methods of Soil Analysis,Part 2:Chemical and Microbial Properties[M].Madison,Wisconsin:Agronomy Society of American,1982:595-624
22 Liu JX,Huang WJ,Zhou GY,Zhang DQ,Liu SR,Li YY.Nitrogen tophosphorus ratios of tree species in response to elevated carbon dioxide and nitrogen addition in subtropical forests[J].Glob Change Biol,2013,19:208-216
Ali AM,Darvishzadeh R,Skidmore AK,van Duren I.Specific leaf area estimation from leaf and canopy reflectance through optimization and validation of vegetation indices[J].Agri Forest Meteo,2017,236:162-174
Li RH,Zhu SD,Chen HYH,John R,Zhou GY,Zhang DQ,Zhang QM,YL Q.Are functional traits a good predictor of global change impacts on tree species abundance dynamics in a subtropical forest?[J] Ecol Let,2015,18:1181-1189
张云海,何念鹏,张光明,黄建辉,韩兴国.氮沉降强度和频率对羊草叶绿素含量的影响[J].生态学报,2013,33:6786-6794[Zhang YH,He NP,Zhang GM,Huang JH,Han XG.Nitrogen deposition and Leymus chinensis leaf chlorophyll content in Inner Mongolian grassland[J].Acta Ecol Sin,2013,33:6786-6794]
Sperfeld E,Martin-Creuzburg D,Wacker A.Multiple resource limitation theory applied to herbivorous consumers:Liebig's minimum rule vs.interactive co-limitation[J].Ecol Lett,2012,15:142-150
27 Vitousek PM,Porder S,Houlton BZ,Chadwick OA.Terrestrial phosphorus limitation:mechanisms,implications,and nitrogenphosphorus interactions[J].Ecol Appl,2010,20:5-15
28 Wright JS,Yavitt JB,Wurzburger N,Turner BL,Tanner E VJ,Sayer E J,Santiago L S.,Kaspari M,Hedin LO,Harms KE,Garcia M N,Corre M D.Potassium,phosphorus,or nitrogen limit root allocation tree growth,or litter production in a lowland tropical forest[J].Ecology,2011,92:1616-1625
29 Hidaka A,Kitayama K.Allocation of foliar phosphorus fractions and leaf traits of tropical tree species in response to decreased soil phosphorus availability on Mount Kinabalu,Borneo[J].J Ecol,2011.99:849-857
30 Hidaka A,Kitayama K.Relationship between photosynthetic phosphorus-use efficiency and foliar phosphorus fractions in tropical tree species[J].Ecol Evol,2013,3:4872-4880
31 Zhang GH,Zhang LL,Wen DZ.Photosynthesis of subtropical forest species from different successional status in relation to foliar nutrients and phosphorus fractions[J].Sci Rep,2018,8:10455
2 Alvarez-Clare S,Mack MC,Brooks M.A direct test of nitrogen and phosphorus limitation to net primary productivity in a lowland tropical wet forest[J].Ecology,2013,94:1540-1551
3 Cernusak,Winter K,Turner BL.Leaf nitrogen to phosphorus ratios of tropical trees:experimental assessment of physiological and environmental controls[J].New Phyt,2010,185:770-779
4 Ostertag R.Foliar nitrogen and phosphorus accumulation responses after fertilization:an example from nutrient-limited Hawaiian forests[J].Plant Soil,2010,334:85-98
5 Tanner EVJ,Kapos V,Freskos S,Healey JR,Theobald AM.Nitrogen and phosphorus fertilization of Jamaican montane forest trees[J].J Trop Ecol,1990,6:231-238
6 Turner BL,Brenes-Arguedas T,Condit R.Pervasive phosphorus limitation of tree species but not communities in tropical forests[J].Nature,2018,555:367
7 Tanner EVJ,Vitousek PM,Cuevas E.Experimental investigation of nutrient limitation of forest growth on wet tropical mountains[J].Ecology,1998,79:10-22
8 Li Y,Niu SL,Yu GR.Aggravated phosphorus limitation on biomass production under increasing nitrogen loading:a meta-analysis[J].Glob Change Biol,2016,22:934-943
9 Santiago LS,Wright SJ,Harms KE,Yavitt JB,Korine C,Garcia MN,Turner BL.Tropical tree seedling growth responses to nitrogen,phosphorus and potassium addition[J].J Ecol,2012,100:309-316
10 Mayor JR,Wright SJ,Turner BL,Austin A.Species-specific responses of foliar nutrients to long-term nitrogen and phosphorus additions in a lowland tropical forest[J].J Ecol,2014,102:36-44
11 Mo QF,Zou B,Li YW,Chen Y,Zhang WX,Mao R,Ding YZ,Wang J,Lu XK,Li XB,Tang JW,Li ZA,Wang FM.Response of plant nutrient stoichiometry to fertilization varied with plant tissues in a tropical forest[J].Sci Reports,2015,5:14605
12莫其锋,陈瑶,王法明,邹碧,李应文,余世钦,李晓波,李志安.华南热带森林两种林下植物氮磷比对氮磷添加的响应[J].应用与环境生物学报,2015,21(5):919-925[Mo QF,Chen Y,Wang FM,Zou B,Li YW,Yu SQ,LI XB,Li ZA.Nitrogen to phosphorus ratios of two understory plant species in response to nitrogen and phosphorus addition in tropical forest of southern China[J].Chin J Appl Environ Biol,2015,21(5):919-925]
13 Zhu FF,Lu XK,Mo JM.Phosphorus limitation on photosynthesis of two dominant understory species in a lowland tropical forest[J].J Plant Ecol,2014,7:526-534
14 Mao QG,Lu XK,Wang C,Zhou KJ,Mo JM.Responses of understory plant physiological traits to a decade of nitrogen addition in a tropical reforested ecosystem[J].For Ecol Ma nage,2017,401:65-74
15 Mao QG,Lu XK,Mo H,Gundersen P,Mo JM.Effects of simulated N deposition on foliar nutrient status,N metabolism and photosynthetic capacity of three dominant understory plant species in a mature tropical forest[J].Sci Total Environ,2018,610-611:555-562
16 Wang FM,Ding YZ,Sayer EJ,Li QL,Zou B,Mo QF,Li YW,Lu XL,Tang JW,Zhu WX,Li ZA.Tropical forest restoration:fast resilience of plant biomass contrasts with slow recovery of stable soil C stocks[J].Func Ecol,2017,31:2344-2355
17 Li J,Li ZA,Wang FM,Zou B,Chen Y,Zhao J,Mo QF,Li YW,Li XB,Xia HP.Effects of nitrogen and phosphorus addition on soil microbial community in a secondary tropical forest of China[J].Biol Fer Soil,2015,51:207-215
18 Chen Y,Sayer EJ,Li ZA,Mo QF,Li YW,Ding YZ,Wang J,Lu XK,Tang JW,Wang FM.Nutrient limitation of woody debris decomposition in a tropical forest:contrasting effects of N and P addition[J].Func Ecol,2016,30:295-304
19 Oberson A.,Friesen DK,Morel C,Tiessen H.Determination of phosphorus released by chloroform fumigation from microbial biomass in high P sorbing tropical soils[J].Soil Biol Biochem,1997,29:1579-1583
20 Lu XK,Mao QG,Gilliam FS,Luo Y,Mo JM.Nitrogen deposition contributes to soil acidification in tropical ecosystems[J].Glob Change Biol,2014,20:3790-3801
21 Bremner JM,Mulvaney CS.Nitrogen-total//Page AL,Miller RH,Keeney DR.Methods of Soil Analysis,Part 2:Chemical and Microbial Properties[M].Madison,Wisconsin:Agronomy Society of American,1982:595-624
22 Liu JX,Huang WJ,Zhou GY,Zhang DQ,Liu SR,Li YY.Nitrogen tophosphorus ratios of tree species in response to elevated carbon dioxide and nitrogen addition in subtropical forests[J].Glob Change Biol,2013,19:208-216
Ali AM,Darvishzadeh R,Skidmore AK,van Duren I.Specific leaf area estimation from leaf and canopy reflectance through optimization and validation of vegetation indices[J].Agri Forest Meteo,2017,236:162-174
Li RH,Zhu SD,Chen HYH,John R,Zhou GY,Zhang DQ,Zhang QM,YL Q.Are functional traits a good predictor of global change impacts on tree species abundance dynamics in a subtropical forest?[J] Ecol Let,2015,18:1181-1189
张云海,何念鹏,张光明,黄建辉,韩兴国.氮沉降强度和频率对羊草叶绿素含量的影响[J].生态学报,2013,33:6786-6794[Zhang YH,He NP,Zhang GM,Huang JH,Han XG.Nitrogen deposition and Leymus chinensis leaf chlorophyll content in Inner Mongolian grassland[J].Acta Ecol Sin,2013,33:6786-6794]
Sperfeld E,Martin-Creuzburg D,Wacker A.Multiple resource limitation theory applied to herbivorous consumers:Liebig's minimum rule vs.interactive co-limitation[J].Ecol Lett,2012,15:142-150
27 Vitousek PM,Porder S,Houlton BZ,Chadwick OA.Terrestrial phosphorus limitation:mechanisms,implications,and nitrogenphosphorus interactions[J].Ecol Appl,2010,20:5-15
28 Wright JS,Yavitt JB,Wurzburger N,Turner BL,Tanner E VJ,Sayer E J,Santiago L S.,Kaspari M,Hedin LO,Harms KE,Garcia M N,Corre M D.Potassium,phosphorus,or nitrogen limit root allocation tree growth,or litter production in a lowland tropical forest[J].Ecology,2011,92:1616-1625
29 Hidaka A,Kitayama K.Allocation of foliar phosphorus fractions and leaf traits of tropical tree species in response to decreased soil phosphorus availability on Mount Kinabalu,Borneo[J].J Ecol,2011.99:849-857
30 Hidaka A,Kitayama K.Relationship between photosynthetic phosphorus-use efficiency and foliar phosphorus fractions in tropical tree species[J].Ecol Evol,2013,3:4872-4880
31 Zhang GH,Zhang LL,Wen DZ.Photosynthesis of subtropical forest species from different successional status in relation to foliar nutrients and phosphorus fractions[J].Sci Rep,2018,8:10455