植物生态学报 ›› 2006, Vol. 30 ›› Issue (1): 25-32.DOI: 10.17521/cjpe.2006.0004
收稿日期:
2004-11-17
接受日期:
2005-06-29
出版日期:
2006-11-17
发布日期:
2006-01-30
通讯作者:
孙书存
作者简介:
*E-mail:shcs@nju.edu.cn基金资助:
Received:
2004-11-17
Accepted:
2005-06-29
Online:
2006-11-17
Published:
2006-01-30
Contact:
SUN Shu-Cun
摘要:
叶物候参数长期以来被认为与植物的碳获取的最大化有关,能反映物种的资源利用策略。温带地区因为寒冷冬天的限制,延长叶寿命成为一些物种进行生长发育和繁衍的基础。为探讨叶寿命延长的可能途径(早出叶、晚落叶,或两者兼有),该研究以南京地区两个落叶栎(Quercus spp.)林为研究对象,观测了其中木本植物的出叶物候、落叶物候,并分析了它们与叶寿命之间的关系。结果发现:1)不同物种的出叶开始时间相差较大,出叶早的物种早结束出叶过程;2)不同物种的落叶开始时间相差较大,早开始落叶的物种,落叶持续时间较长,落叶结束时间则相对集中。3)相关分析和回归分析都表明,叶寿命与出叶时间和落叶时间显著关联,但早出叶对叶寿命的延长可能更为重要,因为早出叶相对于晚落叶在物种资源利用上比较具有优势。4)不同物种的出叶时间和落叶时间没有显著相关,可能因为出叶过程和落叶过程是由不同的启动因子引起。这说明延长叶寿命不一定同时通过早出叶和晚落叶来达到。
朱旭斌, 孙书存. 南京地区落叶栎林木本植物叶物候研究. 植物生态学报, 2006, 30(1): 25-32. DOI: 10.17521/cjpe.2006.0004
ZHU Xu-Bin, SUN Shu-Cun. LEAF PHENOLOGY OF WOODY SPECIES IN DECIDUOUS BROAD-LEAVED OAK FORESTS IN NANJING AREA, EAST CHINA. Chinese Journal of Plant Ecology, 2006, 30(1): 25-32. DOI: 10.17521/cjpe.2006.0004
物种 Species | 出叶开 始时间 Beggi- nning time of leaf emerg- ence | 出叶结 束时间 Ending time of leaf emerg- ence | 出叶持 续时间 Duration of leaf emerg- ence (d) | 落叶开 始时间 Begin- ning time of leaf abscis- sion | 落叶结 束时间 Ending time of leaf abscis- sion | 落叶持 续时间 Duration of leaf abscis- sion (d) | 出叶时 间 Timing of leaf emerg- ence | 落叶 时间 Timing of leaf abscis- sion | 叶寿 命 Leaf life span (d) | ||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
龙王山 | 菝葜Smilax china | 24 | 42 | 18 | 270 | 286 | 16 | 33 | 278 | 245 | |||||
longwangshan | 白栎Quercus fabri | 18 | 35 | 17 | 248 | 280 | 32 | 27 | 264 | 238 | |||||
白檀Symplocos paniculata | 16 | 48 | 32 | 245 | 278 | 33 | 32 | 262 | 230 | ||||||
扁担木Grewia biloba var.parviflora | 25 | 64 | 39 | 256 | 270 | 14 | 45 | 263 | 219 | ||||||
茶条槭Acer ginnala | 7 | 35 | 28 | 238 | 273 | 35 | 21 | 256 | 235 | ||||||
葛藤Pueraria lobata | 28 | 42 | 14 | 235 | 275 | 40 | 35 | 255 | 220 | ||||||
构树Broussonetia papyrifera | 20 | 49 | 29 | 235 | 280 | 45 | 35 | 258 | 223 | ||||||
化香Platycarya strobilacea | 20 | 49 | 29 | 228 | 273 | 45 | 35 | 251 | 216 | ||||||
黄檀Dalbergia hupeana | 35 | 62 | 27 | 265 | 280 | 15 | 49 | 273 | 224 | ||||||
榔榆Ulmus parvifolia | 18 | 35 | 17 | 242 | 274 | 32 | 27 | 258 | 232 | ||||||
茅莓Rubus parvifolius | 0 | 20 | 20 | 239 | 274 | 35 | 10 | 257 | 247 | ||||||
朴树Celtis sinensis | 16 | 42 | 26 | 239 | 286 | 47 | 29 | 263 | 234 | ||||||
山胡椒Lindera glauca | 16 | 35 | 19 | 245 | 274 | 29 | 26 | 260 | 234 | ||||||
长叶鼠李Rhamnus crenata | 23 | 55 | 32 | 249 | 275 | 26 | 39 | 262 | 223 | ||||||
狭叶山胡椒Lindera angustifolia | 20 | 48 | 28 | 265 | 280 | 15 | 34 | 273 | 239 | ||||||
一叶 | 20 | 42 | 22 | 228 | 272 | 44 | 31 | 250 | 219 | ||||||
柘树Cudrania tricuspidata | 20 | 42 | 22 | 248 | 278 | 30 | 31 | 263 | 232 | ||||||
紫藤Wisteria sinensis | 28 | 42 | 14 | 240 | 274 | 34 | 35 | 257 | 222 | ||||||
藏经阁 | 白栎Quercus fabri | 17 | 37 | 20 | 236 | 274 | 38 | 27 | 255 | 228 | |||||
Cangjingge | 白檀Symplocos paniculata | 23 | 57 | 34 | 260 | 286 | 26 | 40 | 273 | 233 | |||||
糙叶树Aphananthe aspera | 24 | 57 | 33 | 237 | 262 | 25 | 41 | 250 | 209 | ||||||
茶条槭Acer ginnala | 17 | 43 | 26 | 246 | 285 | 39 | 30 | 266 | 236 | ||||||
刺楸Kalopanax septemlobus | 20 | 50 | 30 | 225 | 285 | 60 | 35 | 255 | 220 | ||||||
豆腐柴Premna microphylla | 31 | 62 | 31 | 225 | 264 | 39 | 47 | 245 | 198 | ||||||
枫香Liquidambar formosana | 17 | 47 | 30 | 249 | 270 | 21 | 32 | 260 | 228 | ||||||
胡颓子Elaeagnus pungens | 10 | 37 | 27 | 246 | 290 | 44 | 24 | 268 | 245 | ||||||
黄檀Dalbergia hupeana | 31 | 241 | 269 | 28 | 255 | ||||||||||
荚迷Viburnum dilatatum | 10 | 37 | 27 | 255 | 285 | 30 | 24 | 270 | 247 | ||||||
榔榆Ulmus parvifolia | 22 | 43 | 21 | 246 | 270 | 24 | 33 | 258 | 226 | ||||||
六月雪Serissa serissoides | 10 | 269 | 291 | 22 | 280 | ||||||||||
白玉兰Magnolia denudata | 20 | 43 | 23 | 240 | 273 | 33 | 32 | 257 | 225 | ||||||
木莓Rhbus swinhoei | 10 | 43 | 33 | 233 | 291 | 58 | 27 | 262 | 236 | ||||||
三角枫Acer buergerianum | 14 | 34 | 20 | 246 | 270 | 24 | 24 | 258 | 234 | ||||||
栓皮栎Quercus variabilis | 19 | 42 | 23 | 225 | 268 | 43 | 31 | 247 | 216 | ||||||
卫矛Euonymus alatus | 15 | 33 | 18 | 232 | 285 | 53 | 24 | 259 | 235 | ||||||
梧桐Firmiana simplex | 40 | 65 | 25 | 246 | 270 | 24 | 53 | 258 | 206 | ||||||
小槐花Desmodium caudatum | 31 | 49 | 18 | 232 | 290 | 58 | 40 | 261 | 221 | ||||||
小叶石楠Photinia parvifolia | 19 | 50 | 31 | 242 | 267 | 25 | 35 | 255 | 220 | ||||||
盐肤木Rhus chinensis | 24 | 50 | 26 | 232 | 288 | 56 | 37 | 260 | 223 | ||||||
野柿Diospyros rhombifolia var. sylvestris | 31 | 65 | 34 | 241 | 272 | 31 | 48 | 257 | 209 | ||||||
一叶 | 27 | 43 | 16 | 267 | 285 | 18 | 35 | 276 | 241 | ||||||
圆叶鼠李Rhamnus globosa | 17 | 43 | 26 | 236 | 288 | 52 | 30 | 262 | 232 | ||||||
柘树Cudrania tricuspidata | 23 | 50 | 27 | 241 | 273 | 32 | 37 | 257 | 221 |
表1 南京地区两个落叶栎林木本植物叶物候观测数据
Table 1 Data on leaf phenology of the woody species of two deciduous broad-leaved oak forests in Nanjing area, East China
物种 Species | 出叶开 始时间 Beggi- nning time of leaf emerg- ence | 出叶结 束时间 Ending time of leaf emerg- ence | 出叶持 续时间 Duration of leaf emerg- ence (d) | 落叶开 始时间 Begin- ning time of leaf abscis- sion | 落叶结 束时间 Ending time of leaf abscis- sion | 落叶持 续时间 Duration of leaf abscis- sion (d) | 出叶时 间 Timing of leaf emerg- ence | 落叶 时间 Timing of leaf abscis- sion | 叶寿 命 Leaf life span (d) | ||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
龙王山 | 菝葜Smilax china | 24 | 42 | 18 | 270 | 286 | 16 | 33 | 278 | 245 | |||||
longwangshan | 白栎Quercus fabri | 18 | 35 | 17 | 248 | 280 | 32 | 27 | 264 | 238 | |||||
白檀Symplocos paniculata | 16 | 48 | 32 | 245 | 278 | 33 | 32 | 262 | 230 | ||||||
扁担木Grewia biloba var.parviflora | 25 | 64 | 39 | 256 | 270 | 14 | 45 | 263 | 219 | ||||||
茶条槭Acer ginnala | 7 | 35 | 28 | 238 | 273 | 35 | 21 | 256 | 235 | ||||||
葛藤Pueraria lobata | 28 | 42 | 14 | 235 | 275 | 40 | 35 | 255 | 220 | ||||||
构树Broussonetia papyrifera | 20 | 49 | 29 | 235 | 280 | 45 | 35 | 258 | 223 | ||||||
化香Platycarya strobilacea | 20 | 49 | 29 | 228 | 273 | 45 | 35 | 251 | 216 | ||||||
黄檀Dalbergia hupeana | 35 | 62 | 27 | 265 | 280 | 15 | 49 | 273 | 224 | ||||||
榔榆Ulmus parvifolia | 18 | 35 | 17 | 242 | 274 | 32 | 27 | 258 | 232 | ||||||
茅莓Rubus parvifolius | 0 | 20 | 20 | 239 | 274 | 35 | 10 | 257 | 247 | ||||||
朴树Celtis sinensis | 16 | 42 | 26 | 239 | 286 | 47 | 29 | 263 | 234 | ||||||
山胡椒Lindera glauca | 16 | 35 | 19 | 245 | 274 | 29 | 26 | 260 | 234 | ||||||
长叶鼠李Rhamnus crenata | 23 | 55 | 32 | 249 | 275 | 26 | 39 | 262 | 223 | ||||||
狭叶山胡椒Lindera angustifolia | 20 | 48 | 28 | 265 | 280 | 15 | 34 | 273 | 239 | ||||||
一叶 | 20 | 42 | 22 | 228 | 272 | 44 | 31 | 250 | 219 | ||||||
柘树Cudrania tricuspidata | 20 | 42 | 22 | 248 | 278 | 30 | 31 | 263 | 232 | ||||||
紫藤Wisteria sinensis | 28 | 42 | 14 | 240 | 274 | 34 | 35 | 257 | 222 | ||||||
藏经阁 | 白栎Quercus fabri | 17 | 37 | 20 | 236 | 274 | 38 | 27 | 255 | 228 | |||||
Cangjingge | 白檀Symplocos paniculata | 23 | 57 | 34 | 260 | 286 | 26 | 40 | 273 | 233 | |||||
糙叶树Aphananthe aspera | 24 | 57 | 33 | 237 | 262 | 25 | 41 | 250 | 209 | ||||||
茶条槭Acer ginnala | 17 | 43 | 26 | 246 | 285 | 39 | 30 | 266 | 236 | ||||||
刺楸Kalopanax septemlobus | 20 | 50 | 30 | 225 | 285 | 60 | 35 | 255 | 220 | ||||||
豆腐柴Premna microphylla | 31 | 62 | 31 | 225 | 264 | 39 | 47 | 245 | 198 | ||||||
枫香Liquidambar formosana | 17 | 47 | 30 | 249 | 270 | 21 | 32 | 260 | 228 | ||||||
胡颓子Elaeagnus pungens | 10 | 37 | 27 | 246 | 290 | 44 | 24 | 268 | 245 | ||||||
黄檀Dalbergia hupeana | 31 | 241 | 269 | 28 | 255 | ||||||||||
荚迷Viburnum dilatatum | 10 | 37 | 27 | 255 | 285 | 30 | 24 | 270 | 247 | ||||||
榔榆Ulmus parvifolia | 22 | 43 | 21 | 246 | 270 | 24 | 33 | 258 | 226 | ||||||
六月雪Serissa serissoides | 10 | 269 | 291 | 22 | 280 | ||||||||||
白玉兰Magnolia denudata | 20 | 43 | 23 | 240 | 273 | 33 | 32 | 257 | 225 | ||||||
木莓Rhbus swinhoei | 10 | 43 | 33 | 233 | 291 | 58 | 27 | 262 | 236 | ||||||
三角枫Acer buergerianum | 14 | 34 | 20 | 246 | 270 | 24 | 24 | 258 | 234 | ||||||
栓皮栎Quercus variabilis | 19 | 42 | 23 | 225 | 268 | 43 | 31 | 247 | 216 | ||||||
卫矛Euonymus alatus | 15 | 33 | 18 | 232 | 285 | 53 | 24 | 259 | 235 | ||||||
梧桐Firmiana simplex | 40 | 65 | 25 | 246 | 270 | 24 | 53 | 258 | 206 | ||||||
小槐花Desmodium caudatum | 31 | 49 | 18 | 232 | 290 | 58 | 40 | 261 | 221 | ||||||
小叶石楠Photinia parvifolia | 19 | 50 | 31 | 242 | 267 | 25 | 35 | 255 | 220 | ||||||
盐肤木Rhus chinensis | 24 | 50 | 26 | 232 | 288 | 56 | 37 | 260 | 223 | ||||||
野柿Diospyros rhombifolia var. sylvestris | 31 | 65 | 34 | 241 | 272 | 31 | 48 | 257 | 209 | ||||||
一叶 | 27 | 43 | 16 | 267 | 285 | 18 | 35 | 276 | 241 | ||||||
圆叶鼠李Rhamnus globosa | 17 | 43 | 26 | 236 | 288 | 52 | 30 | 262 | 232 | ||||||
柘树Cudrania tricuspidata | 23 | 50 | 27 | 241 | 273 | 32 | 37 | 257 | 221 |
出叶开始 时间 Begginning time of leaf emergence | 出叶结束 时间 Ending time of leaf emergence | 出叶持续 时间 Duration of leaf emergence (d) | 落叶开始 时间 Beginning time of leaf abscission | 落叶结束 时间 Ending time of leaf abscission | 落叶持续 时间 Duratioin of leaf abscission (d) | 出叶时间 Timing of leaf emergence | 落叶时间 Timing of leaf abscission | |
---|---|---|---|---|---|---|---|---|
叶寿命 Leaf life span (d) | -0.607*** | -0.680*** | -0.306ns | 0.490*** | 0.573*** | -0.140ns | -0.730*** | 0.653*** |
落叶时间 Timing of leaf abscission | -0.143ns | -0.063ns | -0.042ns | 0.757*** | 0.708*** | -0.352* | -0.094ns | |
出叶时间 Timing of leaf emergence | 0.830*** | 0.915*** | 0.352* | -0.013ns | -0.196ns | -0.176ns | ||
落叶持续时间 Duratioin of leaf abscission (d) | -0.240ns | -0.119ns | -0.082ns | -0.839*** | 0.330* | |||
落叶结束时间 Ending time of leaf abscission | -0.267ns | -0.168ns | -0.150ns | 0.151ns | ||||
落叶开始时间 Beginning time of leaf abscission | 0.011ns | -0.021ns | 0.019ns | |||||
出叶持续时间 Duration of leaf emergence (d) | 0.025ns | 0.671*** | ||||||
出叶结束时间 Ending time of leaf emergence | 0.626*** |
表2 南京地区落叶栎林叶物候参数的相关分析
Table 2 Correlation analysis for leaf phenological parameters of the woody species of deciduous broad-leaved oak forests in Nanjing area, East China
出叶开始 时间 Begginning time of leaf emergence | 出叶结束 时间 Ending time of leaf emergence | 出叶持续 时间 Duration of leaf emergence (d) | 落叶开始 时间 Beginning time of leaf abscission | 落叶结束 时间 Ending time of leaf abscission | 落叶持续 时间 Duratioin of leaf abscission (d) | 出叶时间 Timing of leaf emergence | 落叶时间 Timing of leaf abscission | |
---|---|---|---|---|---|---|---|---|
叶寿命 Leaf life span (d) | -0.607*** | -0.680*** | -0.306ns | 0.490*** | 0.573*** | -0.140ns | -0.730*** | 0.653*** |
落叶时间 Timing of leaf abscission | -0.143ns | -0.063ns | -0.042ns | 0.757*** | 0.708*** | -0.352* | -0.094ns | |
出叶时间 Timing of leaf emergence | 0.830*** | 0.915*** | 0.352* | -0.013ns | -0.196ns | -0.176ns | ||
落叶持续时间 Duratioin of leaf abscission (d) | -0.240ns | -0.119ns | -0.082ns | -0.839*** | 0.330* | |||
落叶结束时间 Ending time of leaf abscission | -0.267ns | -0.168ns | -0.150ns | 0.151ns | ||||
落叶开始时间 Beginning time of leaf abscission | 0.011ns | -0.021ns | 0.019ns | |||||
出叶持续时间 Duration of leaf emergence (d) | 0.025ns | 0.671*** | ||||||
出叶结束时间 Ending time of leaf emergence | 0.626*** |
图1 出叶物候和落叶物候中参数间关系 A:出叶物候中,出叶开始时间与出叶结束时间关系 The beginning time and the ending time of leaf emergence (R2=0.604,p<0.001) B:落叶物候中,落叶开始时间与落叶持续时间关系 The beginning time of leaf abscission and the duration of leaf abscission (R2=0.626,p<0.001) 图中描述叶物候的时间是由2002年3月1日起的天数 The timing for leaf phenology is defined as the days since March 1, 2002
Fig.1 The relationships between phenological parameters of leaf emergence and leaf abscission
图2 叶寿命与出叶物候、落叶物候之间的关系 A:出叶物候中,出叶开始时间与出叶结束时间关系 The beginning time and the ending time of leaf emergence (R2=0.552,p<0.001) B:落叶物候中,落叶开始时间与落叶持续时间关系 The beginning time of leaf abscission and the duration of leaf abscission (R2=0.462,p<0.001) 图中描述叶物候的时间是由2002年3月1日起的天数 The timing for leaf phenology is defined as the days since March 1, 2002
Fig.2 The relationships between leaf life span and leaf phonological parameters of leaf emergence and abscission
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