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大量的监测数据和全球气候变化模型表明降雨时空格局正在发生改变,由此引起的干旱频度、强度以及持续时间的增加将会成为许多地区面临的问题[1],尤其是在亚热带地区[2-3]。干旱的增加将会对森林生态系统的群落多样性、生存以及生产力产生巨大影响[4],进而可能影响局域或全球的碳水平衡。
叶片对环境变化较为敏感且可塑性较大,其性状特征与植物生存策略和利用资源的能力密切相关[5-6]。因此,加强叶片性状的研究有助于更好地了解植物对干旱的响应与适应。目前,国内外学者从叶片形态、解剖以及生理等性状探究了植物对干旱的响应与适应[7-11]。然而,这些研究大多数是短期的控制实验且缺乏对于叶片解剖结构和光合生理同步研究。因此,对于叶片解剖结构和光合生理对于相对长期干旱胁迫的适应以及二者之间关系的了解依然不足。
研究降雨减少对于叶片解剖结构和光合生理的影响有助于揭示树木对于干旱的适应机制[11]。不同树种对于干旱的敏感性和响应时间存在差异,其叶片解剖结构和光合生理对干旱的响应和适应可能存在物种特异性[11]。对不同水分胁迫条件下地中海2种观赏灌木(马缨丹(Lantana camara L.)和女贞(Ligustrum lucidum W.T.))的研究发现:2种灌木叶片厚度对于水分胁迫响应存在明显差异并指出叶片光合能力与水分胁迫的强度有关[8]。Gratani等[12]发现,干旱条件下叶片厚度的增加可以减少蒸腾失水,维持细胞内较高的水势,从而保证正常的光合速率,提高植物的水分利用效率。然而,Binks等[13]和Rowland等[14]对模拟干旱条件下热带雨林树种的研究证实,叶片解剖结构对于干旱的响应与物种的抗旱性有关,且发现由干旱引起的叶片解剖结构改变未对光合性状产生影响[13]。因此,到目前为止,叶片解剖结构和光合生理对降雨减少的响应认知仍然存在很大争议,这使得理解树木生长对于气候变化的响应具有较大的不确定性。
马尾松(Pinus massoniana Lamb.)和红锥(Castanopsis hystrix Miq.)是我国亚热带地区的典型造林树种。马尾松具有适应性强、生长迅速、生产力高等特点[15],红锥材质优良,具有多种用途[16]。红锥和马尾松的叶片习性、木材解剖和水力学性状存在明显差异,导致2树种拥有不同的生活史特征和环境需求[17-18],这为探究具有不同生活史特征的物种对于干旱的响应和适应提供了绝佳的机会。前人研究已经认识到马尾松和红锥人工林生长对于干旱的响应存在差异,穿透雨减少后马尾松径向生长未发生变化而红锥径向生长出现显著下降[19]。然而,马尾松和红锥叶片解剖结构和光合生理如何响应干旱目前尚未可知。为了进一步探究叶片解剖结构、养分和光合生理对于干旱的响应,解析三者之间的关系,本研究在南亚热带选取马尾松和红锥人工纯林设置穿透雨减少实验,测定了2种树木叶片解剖结构、叶片养分(氮和磷)、气体交换参数和叶绿素荧光参数,旨在探索以下科学问题:(1)马尾松和红锥叶片解剖、叶片养分以及光合生理如何响应干旱?减雨处理后树木是否能够产生更耐旱的叶片?(2)叶片解剖和叶片养分是否能够影响叶片的光合生理?
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土壤温湿度存在明显的季节差异,湿季(5—9月)的土壤温湿度大于干季(10月—次年4月)的土壤温湿度(p < 0.05)。穿透雨减少处理对马尾松和红锥表层(5 cm)土壤温度无影响(p > 0.05),但降低了2树种的表层(5 cm)和深层(10、30、50、70、90 cm)土壤湿度(p < 0.05;图1),其中,穿透雨减少使50 cm土层的土壤湿度下降最大,马尾松和红锥分别下降了15.96%和16.76%。
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由图2得出,马尾松和红锥叶片解剖结构明显不同。马尾松的叶片横切面呈半圆形,叶肉不分化栅栏组织和海绵组织,细胞排列不规则但排列紧密,叶肉细胞间存在多个树脂道。红锥叶片叶肉存在明显分化的栅栏组织和海绵组织,细胞排列规则但不紧密,海绵组织发达。穿透雨减少8 a后马尾松针叶的上、下表皮厚度、内皮层厚度、针叶厚度以及气孔密度分别显著增加了28.35%、29.94%、8.46%、8.13%和10.55%,但叶肉厚度未发生显著改变。对于红锥来说,穿透雨减少不仅明显增加了上表皮厚度(24.43%),而且显著的降低了海绵组织的厚度(23.90%)。穿透雨减少使红锥叶片栅栏和海绵组织比值增加明显,但未改变其叶片厚度和气孔密度(表1)。
图 2 对照样地和减雨样地马尾松(a, b)和红锥(c, d)叶片解剖
Figure 2. Leaf anatomical structure of Pinus massoniana (a, b) and Castanopsis hystrix (c, d) in the control and the rainfall-reduced plots
表 1 马尾松和红锥叶片解剖结构
Table 1. Leaf anatomical structure of Pinus massoniana and Castanopsis hystrix
树种
Tree species叶片指标
Index of leaf对照
Control处理
Treatment马尾松
Pinus massoniana上表皮厚度 Thickness of upper epidermis/μm 12.77 ± 0.21 b 16.39 ± 0.67 a 下表皮厚度 Thickness of lower epidermis /μm 21.41 ± 0.21 b 27.82 ± 0.75 a 叶肉厚度 Mesophyll thickness/μm 104.87 ± 8.91 a 99.79 ± 12.30 a 内皮层厚度 Endothelium thickness/μm 13.83 ± 0.43 b 15.00 ± 0.31 a 树脂道数 Resin canals number 11.33 ± 0.67 a 11.00 ± 0.65 a 针叶厚度 Needle thickness/μm 507.94 ± 3.71 b 549.22 ± 30.94 a 针叶宽 Needle width/μm 875.49 ± 22.11 a 854.03 ± 14.81 a 气孔密度 Stomata density/(number·mm−2) 119.96 ± 10.48 b 132.61 ± 10.21 a 红锥
Castanopsis hystrix上表皮厚度 Thickness of upper epidermis /μm 17.44 ± 0.37 b 21.7 ± 0.54 a 下表皮厚度 Thickness of lower epidermis /μm 11.86 ± 0.37 a 13.27 ± 0.66 a 栅栏组织 Palisade tissue thickness/μm 53.43 ± 1.78 a 53.64 ± 1.38 a 海绵组织 Spongy tissue thickness/μm 44.26 ± 1.5 a 33.68 ± 1.3 b 栅栏/海绵 Ratio of Palisade tissue and spongy tissue 1.22 ± 0.04 b 1.60± 0.04 a 叶片厚度 Leaf thickness/μm 140.10 ± 2.44 a 136.02 ± 2.02 a 气孔密度 Stomata density /(number·mm−2) 468.71 ± 16.68 a 489.58 ± 21.67 a 注:不同的小写字母代表对照和减雨样地之间存在显著差异(p < 0.05)
Note: Lowercase letters indicate significant differences between the control plots and treatment plots (p < 0.05) -
由表2得出:马尾松和红锥的气体交换参数存在季节差异和种间差异(p < 0.05);湿季马尾松和红锥的净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)显著高于干季( p < 0.05);与马尾松对比,红锥受到的气孔限制(Ls)更大,水分利用效率(WUE)更高,尤其是在干季。穿透雨减少对马尾松和红锥气体交换参数和叶绿素荧光参数均无影响(p > 0.05;表2、3和图3)。
表 2 对照样地和减雨样地马尾松和红锥气体交换参数
Table 2. Gas exchange parameters of Pinus massoniana and Castanopsis hystrix in control and rainfall-reduced plots
树种
Tree species气体交换参数
Gas exchange parameters湿季 Wet season 干季 Dry season 对照 Control 处理 Treatment 对照 Control 处理 Treatment 马尾松
Pinus massonianaPn / (μmol·m−2·s−1) 4.17 ± 0.18 Aa 3.92 ± 0.20 Aa 2.10 ± 0.19 Ba 1.72 ± 0.03 Ba Gs / (mol·m−2·s−1) 0.063 ± 0.006 Aa 0.058 ± 0.004 Aa 0.025 ± 0.002 Ba 0.023 ± 0.003 Ba Ci / (μmol·m−2·s−1) 295.95 ± 12.71 Aa 289.42 ± 10.22 Aa 305.44 ± 8.03 Aa 308.22 ± 22.37 Aa Tr / (mmol·m−2·s−1) 0.89 ± 0.76 Aa 0.86 ± 0.06 Aa 0.48 ± 0.05 Ba 0.34 ± 0.064 Ba WUE 4.88 ± 0.47 Aa 4.67 ± 0.41 Aa 4.79 ± 0.52 Aa 6.08 ± 1.18 Aa Ls 0.29 ± 0.07 Ba 0.30 ± 0.02 Ba 0.33 ± 0.044 Aa 0.36 ± 0.065 Aa 红锥
Castanopsis hystrixPn / (μmol·m−2·s−1) 4.59 ± 0.38 Aa 4.42 ± 0.39 Aa 3.66 ± 0.51 Ba 3.24 ± 0.32 Ba Gs / (mol·m−2·s−1) 0.07 ± 0.01 Aa 0.06 ± 0.01 Aa 0.02 ± 0.002 Ba 0.017 ± 0.002 Ba Ci / (μmol·m−2·s−1) 289.10 ± 12.87 Aa 273.31 ± 11.09 Aa 253.24 ± 10.46 Aa 262.85 ± 13.63 Aa Tr / (mmol·m−2·s−1) 0.85 ± 0.09 Aa 0.67 ± 0.06 Aa 0.42 ± 0.02 Ba 0.35 ± 0.02 Ba WUE 5.79 ± 0.62 Ba 6.74 ± 0.70 Ba 8.65 ± 1.08 Aa 9.26 ± 0.66 Aa Ls 0.30 ± 0.03 Ba 0.33 ± 0.03 Ba 0.77 ± 0.02 Aa 0.80 ± 0.03 Aa 注:Pn:净光合速率;Gs:气孔导度;Tr:蒸腾速率;WUE:水分利用效率;Ls:气孔限制。不同的小写字母代表对照和减雨样地之间存在显著差异,不同的大写字母表示季节间存在显著差异(p < 0.05)
Notes: Pn: Net photosynthetic rate; Gs: Stomatal conductance; Tr : Transpiration rate; WUE: Water use efficiency; Ls: Stomatal limitation. Lowercase letters indicate significant differences between the control plots and treatment plots. Uppercase letters indicate significant differences among seasons (p < 0.05)图 3 对照样地和减雨样地马尾松(a)和红锥(b)叶绿素荧光参数
Figure 3. Chlorophyll fluorescence parameters of Pinus massoniana (a) and Castanopsis hystrix (b) in the control and the rainfall-reduced plots
表 3 对照样地和减雨样地马尾松和红锥叶绿素荧光参数
Table 3. Chlorophyll fluorescence parameters of Pinus massoniana and Castanopsis hystrix in control and rainfall-reduced plots
叶绿素荧光参数
Chlorophyll fluorescence parameters马尾松 Pinus massoniana 红锥 Castanopsis hystrix 对照 Control 处理 Treatment 对照 Control 处理 Treatment 最大荧光 Fm 1.46 ± 0.08 a 1.31 ± 0.07 a 1.62 ± 0.13 a 1.58 ± 0.10 a 最大可变荧光 Fv 1.04 ± 0.06 a 0.98 ± 0.06 a 1.28 ± 0.11 a 1.25 ± 0.08 a 表观电子传递速率 ETR 28.91 ± 0.99 a 29.70 ± 1.39 a 34.48 ± 2.03 a 34.52 ± 1.28 a 注:不同的小写字母代表对照和减雨样地之间存在显著差异(p < 0.05)
Note: Lowercase letters indicate significant differences between the control plots and treatment plots (p < 0.05) -
由图4得出:叶片氮、磷含量和氮磷比存在明显的种间差异。马尾松的叶片氮含量以及氮磷比显著低于红锥的叶片氮含量和氮磷比,而马尾松的叶片磷含量显著高于红锥的磷含量。穿透雨减少处理未对马尾松和红锥叶片N、P含量以及N:P产生影响(p > 0.05)。
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由图5、6得出:马尾松和红锥的叶片解剖结构之间、叶片氮和磷之间以及气体交换参数之间自身存在明显地相关关系(p < 0.05),但两树种的叶片解剖结构、叶片养分与气体交换参数三种叶片功能性状之间未发现相关关系(p > 0.05)。
马尾松和红锥叶片解剖和光合对穿透雨减少的响应
The Response of Leaf Anatomical Structure and Photosynthetic Physiology of Pinus massoniana Lamb. and Castanopsis hystrix Miq. to Throughfall Reduction
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摘要:
目的 探究8 a干旱胁迫条件下树木叶片光合生理特征与叶片解剖结构的响应及二者之间的关系。 方法 在南亚热带选取2种典型人工林(马尾松和红锥)设置林内穿透雨减少的模拟干旱实验,在穿透雨减少8 a后测定叶片气体交换参数、叶绿素荧光参数、叶片解剖特征以及叶片养分(N、P)含量。 结果 (1)穿透雨减少后,马尾松针叶上、下表皮、叶片厚度以及气孔密度显著增加,而红锥仅叶片上表皮厚度显著增加,海绵组织厚度显著降低。(2)马尾松和红锥的气体交换参数(净光合速率Pn、气孔导度Gs、蒸腾速率Tr)、叶绿素荧光参数(光系统Ⅱ最大光化学效率Fv/Fm、实际光化学效率 ΦPSII,表观电子传递速率ETR)以及叶氮、磷含量在穿透雨减少8 a后均未发生显著变化。(3)与马尾松相比,红锥受到的气孔限制(Ls)更大,水分利用效率(WUE)更高,尤其在干季更明显。 结论 马尾松和红锥应对干旱胁迫的策略存在差异,马尾松通过调节叶片解剖结构产生更具耐旱的针叶来应对干旱,红锥则更倾向通过调节气孔(Ls)、提高水分利用效率(WUE)来适应干旱。 Abstract:Objective To explore the response of leaf anatomy and leaf photosynthetic physiology to long-term drought and analyze the relationship between leaf anatomy and leaf photosynthetic physiology under long-term drought conditions. Method We conducted a manipulation drought experiment by reducing the throughfall in the forest and measured the leaf gas exchange, chlorophyll fluorescence, leaf nutrient (N, P) and leaf anatomy of the two typical tree species (Pinus massoniana Lamb. and Castanopsis hystrix Miq.) in subtropical China at the 8th year after the throughfall reduction treatment. Results (1) Significant differences were observed in leaf anatomy between P. massoniana and C. hystrix in response to throughfall reduction; the upper and lower epidermis, leaf thickness and stomatal density of P. massoniana needles increased significantly. However, the thickness of the upper epidermis of the leaves of C. hystrix increased significantly, and the thickness of the spongy tissue decreased significantly. (2) No significant differences were observed in the gas exchange parameters (net photosynthetic rate Pn, stomatal conductance Gs, transpiration rate Tr), chlorophyll fluorescence parameters (photosystem II maximum photochemical efficiency Fv/Fm, actual photochemical efficiency ΦPSII, apparent electron transfer rate ETR) and leaf nutrient contents of both species after 8 years of throughfall reduction. (3) Compared with P. massoniana, the stomatal limitation (Ls) of the C. hystrix was larger and the water use efficiency (WUE) was higher, especially in the dry season. Conclusion There are differences in the strategies of P. massoniana and C. hystrix to cope with drought stress. P. massoniana responds to drought by adjusting leaf anatomy to produce xeromorphic needles, while C. hystrix is more inclined to adjust stomata (Ls) and improve water use efficiency (WUE) to adapt to drought. -
表 1 马尾松和红锥叶片解剖结构
Table 1. Leaf anatomical structure of Pinus massoniana and Castanopsis hystrix
树种
Tree species叶片指标
Index of leaf对照
Control处理
Treatment马尾松
Pinus massoniana上表皮厚度 Thickness of upper epidermis/μm 12.77 ± 0.21 b 16.39 ± 0.67 a 下表皮厚度 Thickness of lower epidermis /μm 21.41 ± 0.21 b 27.82 ± 0.75 a 叶肉厚度 Mesophyll thickness/μm 104.87 ± 8.91 a 99.79 ± 12.30 a 内皮层厚度 Endothelium thickness/μm 13.83 ± 0.43 b 15.00 ± 0.31 a 树脂道数 Resin canals number 11.33 ± 0.67 a 11.00 ± 0.65 a 针叶厚度 Needle thickness/μm 507.94 ± 3.71 b 549.22 ± 30.94 a 针叶宽 Needle width/μm 875.49 ± 22.11 a 854.03 ± 14.81 a 气孔密度 Stomata density/(number·mm−2) 119.96 ± 10.48 b 132.61 ± 10.21 a 红锥
Castanopsis hystrix上表皮厚度 Thickness of upper epidermis /μm 17.44 ± 0.37 b 21.7 ± 0.54 a 下表皮厚度 Thickness of lower epidermis /μm 11.86 ± 0.37 a 13.27 ± 0.66 a 栅栏组织 Palisade tissue thickness/μm 53.43 ± 1.78 a 53.64 ± 1.38 a 海绵组织 Spongy tissue thickness/μm 44.26 ± 1.5 a 33.68 ± 1.3 b 栅栏/海绵 Ratio of Palisade tissue and spongy tissue 1.22 ± 0.04 b 1.60± 0.04 a 叶片厚度 Leaf thickness/μm 140.10 ± 2.44 a 136.02 ± 2.02 a 气孔密度 Stomata density /(number·mm−2) 468.71 ± 16.68 a 489.58 ± 21.67 a 注:不同的小写字母代表对照和减雨样地之间存在显著差异(p < 0.05)
Note: Lowercase letters indicate significant differences between the control plots and treatment plots (p < 0.05)表 2 对照样地和减雨样地马尾松和红锥气体交换参数
Table 2. Gas exchange parameters of Pinus massoniana and Castanopsis hystrix in control and rainfall-reduced plots
树种
Tree species气体交换参数
Gas exchange parameters湿季 Wet season 干季 Dry season 对照 Control 处理 Treatment 对照 Control 处理 Treatment 马尾松
Pinus massonianaPn / (μmol·m−2·s−1) 4.17 ± 0.18 Aa 3.92 ± 0.20 Aa 2.10 ± 0.19 Ba 1.72 ± 0.03 Ba Gs / (mol·m−2·s−1) 0.063 ± 0.006 Aa 0.058 ± 0.004 Aa 0.025 ± 0.002 Ba 0.023 ± 0.003 Ba Ci / (μmol·m−2·s−1) 295.95 ± 12.71 Aa 289.42 ± 10.22 Aa 305.44 ± 8.03 Aa 308.22 ± 22.37 Aa Tr / (mmol·m−2·s−1) 0.89 ± 0.76 Aa 0.86 ± 0.06 Aa 0.48 ± 0.05 Ba 0.34 ± 0.064 Ba WUE 4.88 ± 0.47 Aa 4.67 ± 0.41 Aa 4.79 ± 0.52 Aa 6.08 ± 1.18 Aa Ls 0.29 ± 0.07 Ba 0.30 ± 0.02 Ba 0.33 ± 0.044 Aa 0.36 ± 0.065 Aa 红锥
Castanopsis hystrixPn / (μmol·m−2·s−1) 4.59 ± 0.38 Aa 4.42 ± 0.39 Aa 3.66 ± 0.51 Ba 3.24 ± 0.32 Ba Gs / (mol·m−2·s−1) 0.07 ± 0.01 Aa 0.06 ± 0.01 Aa 0.02 ± 0.002 Ba 0.017 ± 0.002 Ba Ci / (μmol·m−2·s−1) 289.10 ± 12.87 Aa 273.31 ± 11.09 Aa 253.24 ± 10.46 Aa 262.85 ± 13.63 Aa Tr / (mmol·m−2·s−1) 0.85 ± 0.09 Aa 0.67 ± 0.06 Aa 0.42 ± 0.02 Ba 0.35 ± 0.02 Ba WUE 5.79 ± 0.62 Ba 6.74 ± 0.70 Ba 8.65 ± 1.08 Aa 9.26 ± 0.66 Aa Ls 0.30 ± 0.03 Ba 0.33 ± 0.03 Ba 0.77 ± 0.02 Aa 0.80 ± 0.03 Aa 注:Pn:净光合速率;Gs:气孔导度;Tr:蒸腾速率;WUE:水分利用效率;Ls:气孔限制。不同的小写字母代表对照和减雨样地之间存在显著差异,不同的大写字母表示季节间存在显著差异(p < 0.05)
Notes: Pn: Net photosynthetic rate; Gs: Stomatal conductance; Tr : Transpiration rate; WUE: Water use efficiency; Ls: Stomatal limitation. Lowercase letters indicate significant differences between the control plots and treatment plots. Uppercase letters indicate significant differences among seasons (p < 0.05)表 3 对照样地和减雨样地马尾松和红锥叶绿素荧光参数
Table 3. Chlorophyll fluorescence parameters of Pinus massoniana and Castanopsis hystrix in control and rainfall-reduced plots
叶绿素荧光参数
Chlorophyll fluorescence parameters马尾松 Pinus massoniana 红锥 Castanopsis hystrix 对照 Control 处理 Treatment 对照 Control 处理 Treatment 最大荧光 Fm 1.46 ± 0.08 a 1.31 ± 0.07 a 1.62 ± 0.13 a 1.58 ± 0.10 a 最大可变荧光 Fv 1.04 ± 0.06 a 0.98 ± 0.06 a 1.28 ± 0.11 a 1.25 ± 0.08 a 表观电子传递速率 ETR 28.91 ± 0.99 a 29.70 ± 1.39 a 34.48 ± 2.03 a 34.52 ± 1.28 a 注:不同的小写字母代表对照和减雨样地之间存在显著差异(p < 0.05)
Note: Lowercase letters indicate significant differences between the control plots and treatment plots (p < 0.05) -
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