间伐对杨桦次生林冠层结构及林下光照的影响
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摘要
自然或人为干扰产生的森林冠层结构和林下光照差异是造成林下环境异质性、物种多样性的主要原因,对森林生态系统的结构、过程与格局具有重要作用.以桃山林场2块50 m×50 m的杨桦次生林固定样地为研究对象,分别作为间伐和对照样地,于2012—2016年、2018年获取样地冠层影像,通过方差分析、马尔科夫模型等方法分析其冠层结构、林下光照动态及间伐后林冠的恢复规律.结果表明:间伐措施能显著调整冠层结构和林下光照,并能持续较长的作用时间,但调整效果随时间逐渐降低;间伐后冠层结构及林下光照的变化速度随时间推移而降低,间伐后前3年林冠恢复速度较快,3年后恢复速度降低并趋于平缓;林下光照与林冠开度呈显著正相关,与叶面积指数呈显著负相关,其中林下散射辐射与冠层结构参数的相关性最大,且间伐样地冠层结构与林下光照的相关性大于对照样地;间伐后冠层的恢复速度与林冠开度大小有关,林冠开度越大,林冠恢复越快,向更小林冠开度转移的时间越早.马尔科夫模型能够模拟冠层结构分布的变化,可用来预测冠层结构动态.
The variation of forest canopy structure and understory light caused by natural or human disturbances might account for environmental heterogeneity and species diversity in the understory. These factors play an important role in driving the structure, process and pattern in forest ecosystem. We set up two 0.25 hm~2 permanent plots in secondary Betula platyphylla-Populus davidiana forests on the Taoshan Forest Farm, one of which was thinned in 2012 and the other one remained natural. The canopy images of two plots were collected by hemispherical photography technology from 2012 to 2016 and 2018. Analysis of variance and Markov matrix were applied to examine the dynamics of canopy structure, understory light, and canopy closing process after thinning. The results showed that thinning was effective in adjusting canopy structure and understory light availability. Such process lasted for a long time and the adjusting effect decreased over time. After thinning, the change rate of canopy structure and understory light decreased over time. The tree canopy quickly closed during the first three years and then reached a stable state. Understory light availability was positively correlated with canopy openness and negatively correlated with leaf area index. The correlation between understory scattered radiation and canopy structural parameters was the strongest. The correlation between canopy structure and understory light in the thinning plot was stronger than that of the control plot. After thinning, the recovery rate of canopy structure was related to the canopy openness, with larger canopy openness being accompanied with higher recovery rate. Thus, less time was required for the transfer to smaller canopy openness. The Markov matrix model could simulate changes in distributions of canopy structure and could be used to predict the dynamics of the canopy structure.
The variation of forest canopy structure and understory light caused by natural or human disturbances might account for environmental heterogeneity and species diversity in the understory. These factors play an important role in driving the structure, process and pattern in forest ecosystem. We set up two 0.25 hm~2 permanent plots in secondary Betula platyphylla-Populus davidiana forests on the Taoshan Forest Farm, one of which was thinned in 2012 and the other one remained natural. The canopy images of two plots were collected by hemispherical photography technology from 2012 to 2016 and 2018. Analysis of variance and Markov matrix were applied to examine the dynamics of canopy structure, understory light, and canopy closing process after thinning. The results showed that thinning was effective in adjusting canopy structure and understory light availability. Such process lasted for a long time and the adjusting effect decreased over time. After thinning, the change rate of canopy structure and understory light decreased over time. The tree canopy quickly closed during the first three years and then reached a stable state. Understory light availability was positively correlated with canopy openness and negatively correlated with leaf area index. The correlation between understory scattered radiation and canopy structural parameters was the strongest. The correlation between canopy structure and understory light in the thinning plot was stronger than that of the control plot. After thinning, the recovery rate of canopy structure was related to the canopy openness, with larger canopy openness being accompanied with higher recovery rate. Thus, less time was required for the transfer to smaller canopy openness. The Markov matrix model could simulate changes in distributions of canopy structure and could be used to predict the dynamics of the canopy structure.
引文
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[3] Li X (李祥),Zhu Y-J (朱玉杰),Dong X-B (董希斌).Canopy structural parameters of Larix gmelini after cutting.Journal of Northeast Forestry University (东北林业大学学报),2015,43(2):1-5 (in Chinese)
[4] Law BE,Cescatti A,Baldocchi DD.Leaf area distribution and radiative transfer in open-canopy forests:Implications for mass and energy exchange.Tree Physiology,2001,21:777-787
[5] Rich PM,Clark DB,Clark DA,et al.Long-term study of solar radiation regimes in a tropical wet forest using quantum sensors and hemispherical photography.Agricultural and Forest Meteorology,1993,65:107-127
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[7] Pang Y (庞勇),Li Z-Y (李增元),Chen E-X (陈尔学),et al.Lidar technology and its application in forestry.Scientia Silvae Sinicae (林业科学),2005,41(3):129-136 (in Chinese)
[8] Chianucci F,Cutini A.Digital hemispherical photography for estimating forest canopy properties:Current controversies and opportunities.iForest Biogeosciences and Forestry,2012,5:290-295
[9] Ou Y-D (区余端),Su Z-Y (苏志尧).Dynamics of canopy structure and understory light in montane evergreen broadleaved forest following a natural disturbance in North Guangdong.Acta Ecologica Sinica (生态学报),2012,32(18):5637-5645 (in Chinese)
[10] Zhang T (张甜),Zhu Y-J (朱玉杰),Dong X-B (董希斌).Canopy structure and light characters after tending felling in Daxing’an Mountains.Journal of Northeast Forestry University (东北林业大学学报),2016,44(10):1-7 (in Chinese)
[11] Fan S-H (范少辉),Liu W-Q (刘蔚漪),Su W-H (苏文会),et al.A comparison of light environmental chara-cteristics of different Phyllostachys heterocycla cv.pubescens forest tapes in Northern Fujian.Journal of Bamboo Research (竹子研究汇刊),2012,31(4):7-11 (in Chinese)
[12] Beaudet M,Messier C.Variation in canopy openness and light transmission following selection cutting in northern hardwood stands:An assessment based on hemispherical photographs.Agricultural and Forest Meteo-rology,2002,110:217-228
[13] Zhu J-J (朱教君),Kang H-Z (康宏樟),Hu L-L (胡理乐).Estimation on optical porosity or canopy closure for a forest stand with hemispherical images.Chinese Journal of Ecology (生态学杂志),2005,24(10):1234-1240 (in Chinese)
[14] Hu L-L (胡理乐),Zhu J-J (朱教君),Li J-S (李俊生),et al.Reviews on methods of measuring light intensities with in forest gaps.Acta Ecologica Sinica (生态学报),2009,29(9):5056-5065 (in Chinese)
[15] Liu Z-L (刘志理),Jin G-Z (金光泽).Estimation of leaf area index of three forest types in Xiaoxing’an Mountains of Northeast China.Chinese Journal of Applied Ecology (应用生态学报),2012,23(9):2437-2444 (in Chinese)
[16] Jia X-R (贾小容),Su Z-R (苏志尧),Ou Y-D (区余端),et al.Canopy structural parameters and understory light regimes of 3 artificial forest stands in South China.Guihaia (广西植物),2011,31(4):473-478 (in Chinese)
[17] Zhu JJ,Matsuzaki T,Lee FQ,et al.Effect of gap size created by thinning on seedling emergency,survival and establishment in a coastal pine forest.Forest Ecology and Management,2003,182:339-354
[18] Canham CD.Growth and canopy architecture of shade-tolerant trees:Response to canopy gaps.Ecology,1988,69:786-795
[19] Lieffers VJ,Messier C,Stadt KJ,et al.Predicting and managing light in the understory of boreal forests.Canadian Journal of Forest Research,1999,29:796-811
[20] Ameztegui A,Coll L,Benavides R,et al.Understory light predictions in mixed conifer mountain forests:Role of aspect-induced variation in crown geometry and openness.Forest Ecology and Management,2012,276:52-61
[21] Zhang SY,Chauret G,Swift DE,et al.Effects of precommercial thinning on tree growth and lumber quality in a jack pine stand in New Brunswick,Canada.Canadian Journal of Forest Research,2006,36:945-952
[22] Lei X-D (雷向东),Lu Y-C (陆元昌),Zhang H-R (张会儒),et al.Effects of thinning on mixed stands of Larix olgensis,Abies nephrolepis and Picea jazoensis.Scientia Silvae Sinicae (林业科学),2005,41(4):78-85 (in Chinese)
[23] Rich PM,Clark DB,Clark DA,et al.Long-term study of solar radiation regimes in a tropical wet forest using quantum sensors and hemispherical photography.Agricultural and Forest Meteorology,1993,65:107-127
[24] Davi H,Baret F,Huc R,et al.Effect of thinning on LAI variance in heterogeneous forests.Forest Ecology and Management,2008,256:890-899
[25] Pelt RV,Franklin JF.Influence of canopy structure on the understory environment in tall,old-growth,conifer forests.Canadian Journal of Forest Research,2000,30:1231-1245
[26] Beaudet M,Messier C,Leduc A.Understory light profiles in temperate deciduous forests:Recovery process following selection cutting.Journal of Ecology,2010,92:328-338
[27] Hong Y (红玉),Wang Y (王耀),Jin X (金鑫),et al.Effects of tending managements on stand structure and soil properties of poplar-birch secondary forests.Journal of Northwest Forestry University (西北林学院学报),2018,33(3):61-66 (in Chinese)
[28] Liang J (梁晶),Wang Q-C (王庆成),Xu L-J (许丽娟),et al.Effect of thinning on respiration rate and soil carbon density of young-age poplar-birch stands in western Changbai Mountains.Bulletin of Botanical Research (植物研究),2015,35(1):110-116 (in Chinese)
[29] Zhu J-J (朱教君).Method for measurement of optical stratification porosity (OSP) and its application in stu-dies of management for secondary forests.Chinese Journal of Applied Ecology (应用生态学报),2003,14(8):1229-1233 (in Chinese)
[30] Meng X-Y (孟宪宇).Forest Mensuration.Beijing:China Forestry Press,2006 (in Chinese)
[31] Fotis AT,Morin TH,Fahey RT,et al.Forest structure in space and time:Biotic and abiotic determinants of canopy complexity and their effects on net primary productivity.Agricultural and Forest Meteorology,2018,250:181-191
[32] Makinen H,Isomaki A.Thinning intensity and growth of Scots pine stands in Finland.Forest Ecology and Mana-gement,2004,201:311-325
[33] Huang H-M (黄慧敏),Dong R (董蓉),He D-N (何丹妮),et al.Effects of temporal and spatial variaton of canopy structures and light conditons on population characteristics of Fargesia decurvata.Chinese Journal of Applied Ecology (应用生态学报),2018,29(7):2129-2138 (in Chinese)
[34] Primicia I,Artazcoz R,Imbert JB,et al.Influence of thinning intensity and canopy type on Scots pine stand and growth dynamics in a mixed managed forest.Forest Systems,2016,25:1-10
[35] Teresa V,Jonathan S.Canopy closure rate and forest structure.Ecology,1997,78:1555-1562
[36] Zhang T (张甜),Zhu Y-J (朱玉杰),Dong X-B (董希斌).Effects of thinning and pruning on the growth and canopy of larch forest.Journal of Northeast Forestry University (东北林业大学学报),2017,45(12):8-11 (in Chinese)
[37] Cao Y-H (曹永慧),Chen C-J (陈存及),Li S (李生).Impacts of different intermediate cutting on crown structure of Manglietia yuyuanensis in mixed stands with Cunninghamia lanceolata.Forest Research (林业科学研究),2004,17(5):646-653 (in Chinese)
[38] Wen Y-G (温远光),Lu W-K (卢文科),Zhou X-G (周晓果),et al.Spatial heterogeneity dynamics of canopy structure in evergreen broadleaved forest after a severe ice storm disturbance in Damingshan,Southern China.Guangxi Sciences (广西科学),2017,24(2):150-158 (in Chinese)
[39] Meer PV,Bongers F.Formation and closure of canopy gaps in the rainforest at Nouragues,French Guiana.Vegetatio,1996,126:167-179
[40] Rikimaru A,Roy PS,Miyatake S.Tropical forest cover density mapping.Tropical Ecology,2002,43:39-47
[2] Nicotra AB,Chazdon RL,Iriarte S.Spatial heterogeneity of light and woody seedling regeneration in tropical wet forests.Ecology,1999,80:1908-1926
[3] Li X (李祥),Zhu Y-J (朱玉杰),Dong X-B (董希斌).Canopy structural parameters of Larix gmelini after cutting.Journal of Northeast Forestry University (东北林业大学学报),2015,43(2):1-5 (in Chinese)
[4] Law BE,Cescatti A,Baldocchi DD.Leaf area distribution and radiative transfer in open-canopy forests:Implications for mass and energy exchange.Tree Physiology,2001,21:777-787
[5] Rich PM,Clark DB,Clark DA,et al.Long-term study of solar radiation regimes in a tropical wet forest using quantum sensors and hemispherical photography.Agricultural and Forest Meteorology,1993,65:107-127
[6] Bianchi S,Cahalan C,Hale S,et al.Rapid assessment of forest canopy and light regime using smartphone hemispherical photography.Ecology and Evolution,2017,7:10556-10566
[7] Pang Y (庞勇),Li Z-Y (李增元),Chen E-X (陈尔学),et al.Lidar technology and its application in forestry.Scientia Silvae Sinicae (林业科学),2005,41(3):129-136 (in Chinese)
[8] Chianucci F,Cutini A.Digital hemispherical photography for estimating forest canopy properties:Current controversies and opportunities.iForest Biogeosciences and Forestry,2012,5:290-295
[9] Ou Y-D (区余端),Su Z-Y (苏志尧).Dynamics of canopy structure and understory light in montane evergreen broadleaved forest following a natural disturbance in North Guangdong.Acta Ecologica Sinica (生态学报),2012,32(18):5637-5645 (in Chinese)
[10] Zhang T (张甜),Zhu Y-J (朱玉杰),Dong X-B (董希斌).Canopy structure and light characters after tending felling in Daxing’an Mountains.Journal of Northeast Forestry University (东北林业大学学报),2016,44(10):1-7 (in Chinese)
[11] Fan S-H (范少辉),Liu W-Q (刘蔚漪),Su W-H (苏文会),et al.A comparison of light environmental chara-cteristics of different Phyllostachys heterocycla cv.pubescens forest tapes in Northern Fujian.Journal of Bamboo Research (竹子研究汇刊),2012,31(4):7-11 (in Chinese)
[12] Beaudet M,Messier C.Variation in canopy openness and light transmission following selection cutting in northern hardwood stands:An assessment based on hemispherical photographs.Agricultural and Forest Meteo-rology,2002,110:217-228
[13] Zhu J-J (朱教君),Kang H-Z (康宏樟),Hu L-L (胡理乐).Estimation on optical porosity or canopy closure for a forest stand with hemispherical images.Chinese Journal of Ecology (生态学杂志),2005,24(10):1234-1240 (in Chinese)
[14] Hu L-L (胡理乐),Zhu J-J (朱教君),Li J-S (李俊生),et al.Reviews on methods of measuring light intensities with in forest gaps.Acta Ecologica Sinica (生态学报),2009,29(9):5056-5065 (in Chinese)
[15] Liu Z-L (刘志理),Jin G-Z (金光泽).Estimation of leaf area index of three forest types in Xiaoxing’an Mountains of Northeast China.Chinese Journal of Applied Ecology (应用生态学报),2012,23(9):2437-2444 (in Chinese)
[16] Jia X-R (贾小容),Su Z-R (苏志尧),Ou Y-D (区余端),et al.Canopy structural parameters and understory light regimes of 3 artificial forest stands in South China.Guihaia (广西植物),2011,31(4):473-478 (in Chinese)
[17] Zhu JJ,Matsuzaki T,Lee FQ,et al.Effect of gap size created by thinning on seedling emergency,survival and establishment in a coastal pine forest.Forest Ecology and Management,2003,182:339-354
[18] Canham CD.Growth and canopy architecture of shade-tolerant trees:Response to canopy gaps.Ecology,1988,69:786-795
[19] Lieffers VJ,Messier C,Stadt KJ,et al.Predicting and managing light in the understory of boreal forests.Canadian Journal of Forest Research,1999,29:796-811
[20] Ameztegui A,Coll L,Benavides R,et al.Understory light predictions in mixed conifer mountain forests:Role of aspect-induced variation in crown geometry and openness.Forest Ecology and Management,2012,276:52-61
[21] Zhang SY,Chauret G,Swift DE,et al.Effects of precommercial thinning on tree growth and lumber quality in a jack pine stand in New Brunswick,Canada.Canadian Journal of Forest Research,2006,36:945-952
[22] Lei X-D (雷向东),Lu Y-C (陆元昌),Zhang H-R (张会儒),et al.Effects of thinning on mixed stands of Larix olgensis,Abies nephrolepis and Picea jazoensis.Scientia Silvae Sinicae (林业科学),2005,41(4):78-85 (in Chinese)
[23] Rich PM,Clark DB,Clark DA,et al.Long-term study of solar radiation regimes in a tropical wet forest using quantum sensors and hemispherical photography.Agricultural and Forest Meteorology,1993,65:107-127
[24] Davi H,Baret F,Huc R,et al.Effect of thinning on LAI variance in heterogeneous forests.Forest Ecology and Management,2008,256:890-899
[25] Pelt RV,Franklin JF.Influence of canopy structure on the understory environment in tall,old-growth,conifer forests.Canadian Journal of Forest Research,2000,30:1231-1245
[26] Beaudet M,Messier C,Leduc A.Understory light profiles in temperate deciduous forests:Recovery process following selection cutting.Journal of Ecology,2010,92:328-338
[27] Hong Y (红玉),Wang Y (王耀),Jin X (金鑫),et al.Effects of tending managements on stand structure and soil properties of poplar-birch secondary forests.Journal of Northwest Forestry University (西北林学院学报),2018,33(3):61-66 (in Chinese)
[28] Liang J (梁晶),Wang Q-C (王庆成),Xu L-J (许丽娟),et al.Effect of thinning on respiration rate and soil carbon density of young-age poplar-birch stands in western Changbai Mountains.Bulletin of Botanical Research (植物研究),2015,35(1):110-116 (in Chinese)
[29] Zhu J-J (朱教君).Method for measurement of optical stratification porosity (OSP) and its application in stu-dies of management for secondary forests.Chinese Journal of Applied Ecology (应用生态学报),2003,14(8):1229-1233 (in Chinese)
[30] Meng X-Y (孟宪宇).Forest Mensuration.Beijing:China Forestry Press,2006 (in Chinese)
[31] Fotis AT,Morin TH,Fahey RT,et al.Forest structure in space and time:Biotic and abiotic determinants of canopy complexity and their effects on net primary productivity.Agricultural and Forest Meteorology,2018,250:181-191
[32] Makinen H,Isomaki A.Thinning intensity and growth of Scots pine stands in Finland.Forest Ecology and Mana-gement,2004,201:311-325
[33] Huang H-M (黄慧敏),Dong R (董蓉),He D-N (何丹妮),et al.Effects of temporal and spatial variaton of canopy structures and light conditons on population characteristics of Fargesia decurvata.Chinese Journal of Applied Ecology (应用生态学报),2018,29(7):2129-2138 (in Chinese)
[34] Primicia I,Artazcoz R,Imbert JB,et al.Influence of thinning intensity and canopy type on Scots pine stand and growth dynamics in a mixed managed forest.Forest Systems,2016,25:1-10
[35] Teresa V,Jonathan S.Canopy closure rate and forest structure.Ecology,1997,78:1555-1562
[36] Zhang T (张甜),Zhu Y-J (朱玉杰),Dong X-B (董希斌).Effects of thinning and pruning on the growth and canopy of larch forest.Journal of Northeast Forestry University (东北林业大学学报),2017,45(12):8-11 (in Chinese)
[37] Cao Y-H (曹永慧),Chen C-J (陈存及),Li S (李生).Impacts of different intermediate cutting on crown structure of Manglietia yuyuanensis in mixed stands with Cunninghamia lanceolata.Forest Research (林业科学研究),2004,17(5):646-653 (in Chinese)
[38] Wen Y-G (温远光),Lu W-K (卢文科),Zhou X-G (周晓果),et al.Spatial heterogeneity dynamics of canopy structure in evergreen broadleaved forest after a severe ice storm disturbance in Damingshan,Southern China.Guangxi Sciences (广西科学),2017,24(2):150-158 (in Chinese)
[39] Meer PV,Bongers F.Formation and closure of canopy gaps in the rainforest at Nouragues,French Guiana.Vegetatio,1996,126:167-179
[40] Rikimaru A,Roy PS,Miyatake S.Tropical forest cover density mapping.Tropical Ecology,2002,43:39-47