林业科学 ›› 2021, Vol. 57 ›› Issue (4): 63-72.doi: 10.11707/j.1001-7488.20210407
黄贝佳,朱静,何晨阳,李伟煌,陈志为,樊月,洪滔*
收稿日期:
2020-07-21
出版日期:
2021-04-25
发布日期:
2021-05-21
通讯作者:
洪滔
基金资助:
Beijia Huang,Jing Zhu,Chenyang He,Weihuang Li,Zhiwei Chen,Yue Fan,Tao Hong*
Received:
2020-07-21
Online:
2021-04-25
Published:
2021-05-21
Contact:
Tao Hong
摘要:
目的: 明确秋茄叶表型及叶经济谱对自然潮汐生境的响应特征,以期掌握秋茄在周期性浸淹下的适应机制。方法: 对福州市罗源湾秋茄成熟林满潮线上及线下叶片进行研究,以上层未受淹叶片为对照,测定上、下层叶表型性状,并对各性状间关系及叶经济谱进行分析。结果: 与满潮线上层叶片相比,满潮线下层叶片干质量、叶柄长及叶干物质含量较小(P < 0.01),叶形指数较小(P < 0.05),比叶面积较大(P < 0.01);在叶片气孔特性中,满潮线下层叶片气孔长度、宽度、面积、周长、开度、开口长度、开口面积、开口周长均大于上层,而气孔密度则小于上层(P < 0.01);满潮线下层叶片的净光合速率、蒸腾速率及气孔导度较小,而胞间CO2浓度较大(P < 0.01),羧化速率较小(P < 0.01),水分利用效率在满潮线上、下层叶片中差别不显著。满潮线上层秋茄叶片气孔特性与光合功能之间相关性比下层高,而满潮线下层叶气孔特性与叶形态性状相关性高于上层。结论: 秋茄满潮线上、下层的叶表型差异明显,下层叶片气孔功能适应性较强,但光合功能受到一定限制。上层叶气孔特性与光合功能具有协同进化表现,而下层叶气孔特性与叶形态性状存在趋同适应。满潮线上、下层秋茄叶为适应自然潮汐生境,存在明显生存策略选择。叶经济谱表现为:满潮线下层叶片总体上向"缓慢投资-收益型"一端偏移,具有较小的叶干物质含量、气孔密度,较低的净光合速率及较大的比叶面积、气孔面积和气孔开度,以适应周期性低光、缺氧的淹水极端环境;上层叶片则向"快速投资-收益型"一端偏移以应对海岸带大风、海浪拍打等特殊生境。
中图分类号:
黄贝佳,朱静,何晨阳,李伟煌,陈志为,樊月,洪滔. 潮汐对秋茄叶表型塑造及叶经济谱的影响[J]. 林业科学, 2021, 57(4): 63-72.
Beijia Huang,Jing Zhu,Chenyang He,Weihuang Li,Zhiwei Chen,Yue Fan,Tao Hong. Effects of Tides on Leaf Phenotypic Modeling and Leaf Economics Spectrum of Kandelia obovata[J]. Scientia Silvae Sinicae, 2021, 57(4): 63-72.
表1
满潮线上、下层秋茄叶片的形态性状(平均值±标准差)①"
叶形态性状 Leaf morphological traits | 满潮线上层叶片 Leaf above high tide line | 满潮线下层叶片 Leaf under high tide line |
鲜质量 Fresh weight(FW)/g | 1.65±0.41 | 1.75±0.51 |
干质量 Dry weight(DW)/g | 0.51±0.12** | 0.45±0.13** |
叶厚 Leaf thickness(LT)/mm | 0.66±0.10 | 0.64±0.13 |
叶柄厚 Petiole thickness(PT)/mm | 2.25±0.29 | 2.24±0.34 |
叶柄长 Petiole length(PL)/cm | 1.21±0.30** | 1.01±0.25** |
叶长 Leaf length(LL)/cm | 9.97±1.16 | 9.97±1.09 |
叶宽 Leaf width(LW)/cm | 3.90±0.48 | 4.06±0.51 |
叶面积 Leaf area(LA)/cm2 | 25.03±5.57 | 26.64±5.78 |
叶周长 Leaf perimeter(LP)/cm | 24.06±3.06 | 24.56±3.56 |
比叶面积 Specific leaf area(SLA)/(cm2·g-1) | 49.28±6.72** | 61.39±10.31** |
叶干物质含量 Leaf dry matter content(LDMC)/(g·g-1) | 0.32±0.26** | 0.11±0.06** |
叶形指数 Leaf shape index(LSI) | 2.58±0.33* | 2.48±0.26* |
表2
满潮线上、下层秋茄叶的气孔特征(平均值±标准差)"
气孔特征 Stomatal characteristics | 满潮线上层叶片 Leaf above high tide line | 满潮线下层叶片 Leaf under high tide line |
气孔长度 Stomatal length(SL)/μm | 40.14±3.61** | 42.39±3.30** |
气孔宽度 Stomatal width(SW)/μm | 18.72±2.64** | 20.31±2.48** |
气孔长宽比 Stomatal length-width ratio(SLWR) | 2.19±0.36** | 2.12±0.30** |
气孔面积 Stomatal area(SA)/μm2 | 566.53±103.77** | 636.81±101.22** |
气孔周长 Stomatal perimeter(SP)/μm | 94.77±7.88** | 100.21±7.69** |
气孔开口长度 Stomatal opening length(SOL)/μm | 33.79±3.69** | 36.10±3.38** |
气孔开度 Stomatal aperture(Sa)/μm | 9.28±2.64** | 10.20±2.33** |
气孔开口长宽比 Stomatal opening length-width ratio(SOLWR) | 3.92±1.11** | 3.73±0.96** |
气孔开口面积 Stomatal opening area(SOA)/μm2 | 229.23±79.28** | 261.33±69.90** |
气孔开口周长 Stomatal opening perimeter(SOP)/μm | 72.91±10.26** | 77.71±8.96** |
气孔密度 Stomatal density(SD)/mm-2 | 146.98±18.65** | 132.86±14.66** |
气孔面积指数 Stomatal area index(SAI)(%) | 8.26±1.27 | 8.41±1.07 |
气孔形状指数 Stomatal shape index(SSI) | 1.13±0.03 | 1.12±0.03 |
表3
满潮线上、下层秋茄叶的光合特性(平均值±标准差)"
光合参数 Photosynthetic parameters | 满潮线上层叶片 Leaf above high tide line | 满潮线下层叶片 Leaf under high tide line |
净光合速率 Net photosynthetic rate(Pn)/(μmol·m-2 s-1) | 8.29±2.58** | 6.08±2.13** |
气孔导度 Stomatal conductance(Gs)/(mol·m-2 s-1) | 0.11±0.04* | 0.09±0.03* |
胞间CO2浓度 Intercellular CO2 concentration(Ci)/(μmol·mol-1) | 273.77±29.88** | 307.46±32.39** |
蒸腾速率 Transpiration rate(Tr)/(mmol·m-2 s-1) | 1.50±0.62** | 1.20±0.48** |
水分利用效率 Water use efficiency(WUE)/(μmol·mmol-1) | 6.20±2.58 | 5.46±1.69 |
羧化速率 Carboxylation efficiency(CE)/(μmol·m-2 s-1) | 0.03±0.00** | 0.02±0.00** |
表4
满潮线上层与下层秋茄叶形态性状与气孔特征间的Pearson相关系数①"
叶片位置 Leaf position | 鲜质量 FW | 干质量 DW | 叶厚 LT | 叶柄厚 PT | 叶柄长 PL | 叶长 LL | 叶宽 LW | 叶面积 LA | 叶周长 LP | 比叶面积 SLA | 叶干物质含量 LDMC | 叶形指数 LSI | |
气孔长度 SL | 上层 Above | -0.190 | -0.133 | -0.083 | -0.341 | -0.019 | -0.105 | -0.330 | -0.259 | -0.134 | -0.225 | -0.056 | 0.217 |
下层 Under | 0.369 | 0.343 | 0.308 | 0.114 | -0.132 | 0.221 | 0.077 | 0.212 | 0.342 | -0.440* | 0.037 | 0.169 | |
气孔宽度 SW | 上层 Above | 0.129 | 0.141 | 0.237 | 0.056 | 0.257 | 0.112 | -0.254 | -0.128 | 0.183 | -0.499** | -0.157 | 0.341 |
下层 Under | 0.252 | 0.255 | 0.022 | -0.103 | -0.061 | 0.285 | -0.094 | 0.168 | 0.330 | -0.252 | 0.078 | 0.379 | |
气孔面积 SA | 上层 Above | -0.045 | 0.040 | 0.135 | -0.131 | 0.181 | 0.012 | -0.342 | -0.224 | 0.059 | -0.494** | -0.086 | 0.330 |
下层 Under | 0.443* | 0.394* | 0.299 | 0.119 | -0.043 | 0.357 | 0.053 | 0.276 | 0.419* | -0.435* | 0.036 | 0.330 | |
气孔开口长度 SOL | 上层 Above | -0.288 | -0.242 | -0.036 | -0.362 | -0.075 | -0.244 | -0.356 | -0.354 | -0.270 | -0.182 | -0.099 | 0.124 |
下层 Under | 0.425* | 0.347 | 0.348 | 0.189 | -0.093 | 0.290 | 0.146 | 0.277 | 0.297 | -0.387* | -0.011 | 0.180 | |
气孔开度 Sa | 上层 Above | -0.062 | -0.073 | 0.169 | 0.023 | 0.058 | -0.155 | -0.157 | -0.214 | -0.090 | -0.246 | -0.232 | 0.046 |
下层 Under | 0.288 | 0.203 | 0.067 | 0.081 | 0.160 | 0.419* | 0.132 | 0.359 | 0.282 | 0.096 | -0.050 | 0.294 | |
气孔开口面积 SOA | 上层 Above | -0.159 | -0.140 | 0.159 | -0.131 | 0.026 | -0.222 | -0.260 | -0.312 | -0.164 | -0.294 | -0.221 | 0.071 |
下层 Under | 0.453* | 0.339 | 0.314 | 0.213 | 0.093 | 0.453* | 0.186 | 0.398* | 0.376 | -0.174 | -0.050 | 0.301 | |
气孔密度 SD | 上层 Above | 0.128 | -0.047 | 0.091 | 0.191 | -0.270 | -0.185 | 0.334 | 0.081 | -0.179 | 0.269 | -0.248 | -0.440* |
下层 Under | -0.119 | 0.019 | -0.057 | 0.150 | 0.090 | 0.049 | -0.214 | -0.028 | -0.075 | 0.099 | 0.169 | 0.202 | |
气孔面积指数 SAI | 上层 Above | 0.138 | 0.055 | 0.260 | 0.096 | -0.042 | -0.103 | -0.012 | -0.104 | -0.037 | -0.275 | -0.326 | -0.048 |
下层 Under | 0.359 | 0.440* | 0.284 | 0.268 | 0.024 | 0.414* | -0.155 | 0.249 | 0.376 | -0.403* | 0.213 | 0.542** |
表5
满潮线上层与下层秋茄叶形态性状与光合特性间的Pearson相关系数"
叶片位置 Leaf position | 鲜质量 FW | 干质量 DW | 叶厚 LT | 叶柄厚 PT | 叶柄长 PL | 叶长 LL | 叶宽 LW | 叶面积 LA | 叶周长 LP | 比叶面积 SLA | 叶干物质含量 LDMC | 叶形指数 LSI | |
净光合速率 Pn | 上层 Above | 0.054 | -0.156 | -0.135 | 0.015 | -0.324 | -0.277 | 0.108 | -0.070 | -0.316 | 0.166 | -0.164 | -0.330 |
下层 Under | 0.056 | 0.028 | -0.261 | -0.227 | 0.097 | 0.034 | -0.085 | 0.039 | 0.181 | -0.032 | -0.096 | 0.083 | |
气孔导度 Gs | 上层 Above | 0.112 | -0.119 | 0.040 | 0.017 | -0.101 | -0.206 | -0.030 | -0.173 | -0.242 | -0.077 | -0.227 | -0.154 |
下层 Under | 0.106 | -0.016 | -0.282 | -0.300 | 0.069 | -0.030 | -0.034 | 0.068 | 0.227 | 0.052 | -0.206 | -0.052 | |
胞间CO2浓度 Ci | 上层 Above | 0.208 | 0.034 | 0.223 | 0.036 | 0.180 | 0.104 | -0.064 | -0.064 | 0.131 | -0.142 | -0.223 | 0.159 |
下层 Under | 0.014 | -0.193 | 0.101 | -0.090 | -0.029 | -0.114 | 0.012 | -0.046 | 0.068 | 0.156 | -0.247 | -0.136 | |
蒸腾速率 Tr | 上层 Above | 0.111 | -0.070 | 0.296 | 0.186 | -0.222 | -0.196 | 0.066 | -0.050 | -0.223 | 0.046 | -0.202 | -0.236 |
下层 Under | 0.309 | 0.119 | -0.078 | -0.139 | -0.311 | -0.091 | 0.160 | 0.187 | 0.161 | -0.019 | -0.269 | -0.305 | |
水分利用效率 WUE | 上层 Above | -0.181 | -0.114 | -0.390* | -0.287 | 0.049 | -0.048 | -0.090 | -0.092 | -0.050 | 0.032 | 0.078 | 0.053 |
下层 Under | -0.165 | 0.099 | -0.047 | 0.109 | 0.517** | 0.319 | -0.237 | -0.030 | 0.092 | -0.121 | 0.328 | 0.577** | |
羧化速率 CE | 上层 Above | -0.054 | -0.184 | -0.229 | -0.038 | -0.371 | -0.303 | 0.109 | -0.057 | -0.341 | 0.232 | -0.071 | -0.357 |
下层 Under | 0.026 | 0.050 | -0.190 | -0.198 | 0.046 | 0.029 | -0.130 | -0.005 | 0.113 | -0.093 | 0.002 | 0.138 |
表6
满潮线上层与下层秋茄叶气孔特征与光合特性间的Pearson相关系数"
叶片位置 Leaf position | 气孔长度 SL | 气孔宽度 SW | 气孔面积 SA | 气孔开口长度 SOL | 气孔开度 Sa | 气孔开口面积 SOA | 气孔密度 SD | 气孔面积指数 SAI | |
净光合速率Pn | 上层 Above | -0.257 | -0.355 | -0.394* | -0.299 | -0.408* | -0.425* | 0.272 | -0.192 |
下层 Under | -0.081 | 0.009 | -0.021 | -0.122 | -0.099 | -0.089 | 0.113 | 0.064 | |
气孔导度Gs | 上层 Above | -0.054 | 0.055 | -0.034 | -0.056 | 0.023 | -0.007 | 0.168 | 0.109 |
下层 Under | 0.040 | 0.168 | 0.096 | -0.016 | 0.026 | -0.017 | -0.196 | -0.085 | |
胞间CO2浓度Ci | 上层 Above | 0.247 | 0.489** | 0.389* | 0.242 | 0.488** | 0.453* | 0.029 | 0.476* |
下层 Under | 0.228 | 0.225 | 0.221 | 0.215 | 0.172 | 0.160 | -0.489** | -0.190 | |
蒸腾速率Tr | 上层 Above | -0.139 | 0.076 | -0.022 | -0.044 | 0.127 | 0.080 | 0.098 | 0.069 |
下层 Under | 0.111 | 0.000 | 0.063 | 0.068 | -0.116 | -0.060 | -0.203 | -0.123 | |
水分利用效率 WUE | 上层 Above | 0.136 | -0.203 | -0.085 | 0.014 | -0.282 | -0.209 | 0.030 | -0.078 |
下层 Under | -0.094 | 0.120 | 0.042 | -0.077 | 0.157 | 0.121 | 0.384* | 0.392* | |
羧化速率 CE | 上层 Above | -0.285 | -0.498** | -0.479* | -0.319 | -0.529** | -0.521** | 0.213 | -0.348 |
下层 Under | -0.122 | 0.040 | -0.034 | -0.136 | 0.005 | -0.011 | 0.168 | 0.104 |
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[1] | 黄剑坚, 李际平, 刘素青, 韩维栋. 红树林扩展林窗空间营养指数[J]. 林业科学, 2013, 49(9): 16-22. |
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