Response of leaf functional traits and anatomical structure to altitude in Crataegus songarica K. Koch in Tianshan wild fruit forest
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摘要:
为揭示准噶尔山楂（Crataegus songarica K. Koch）对海拔高度变化的响应，以新疆伊犁果子沟不同海拔（1100~1700 m）的准噶尔山楂作为研究对象，分析其叶片功能性状及解剖结构差异。结果显示：（1）随着海拔的升高，准噶尔山楂的叶长、叶宽、叶面积、比叶面积、叶绿素含量呈降低趋势，叶长宽比、比叶重、叶干物质含量呈上升趋势，比叶重可塑性最强。（2）随着海拔升高，叶厚、栅栏组织厚度、上下表皮厚度呈上升趋势；主脉厚度、突起度呈下降趋势，主脉突起度可塑性最强。（3）叶面积与叶长、叶长宽比、比叶面积呈正相关；叶厚与上下表皮厚度、主脉突起度等呈正相关。研究结果表明，准噶尔山楂在高海拔区域主要通过增加叶厚和叶干物质含量以提高抗逆性，减少叶面积、叶绿素含量来降低高光强对叶片的伤害；在低海拔区域主要通过增加叶面积、叶绿素含量来促进有机物的积累。
Abstract:This study investigated the adaptive responses of Crataegus songarica K. Koch to varying altitudes (1 100-1 700 m) in Guozigou, Yili, Xinjiang, China, focusing on leaf functional traits and anatomical structure. Results showed that: (1) An increase in altitude led to a reduction in leaf length, leaf width, leaf area, specific leaf area, and chlorophyll content, but an increase in the leaf length to width ratio, specific leaf weight, and leaf dry matter content, with specific leaf weight showing the strongest plasticity. (2) Higher altitudes were associated with increasing leaf thickness, palisade tissue thickness, and upper and lower epidermis thickness, but decreasing central vein thickness and protrusion, with central vein protrusion showing the strongest plasticity. (3) Leaf area was positively correlated with leaf length, length-width ratio, and specific leaf area, while leaf thickness was positively correlated with upper and lower epidermis thickness and central vein protrusion. Overall, our findings indicated that Crataegus songarica adapts to high-altitude conditions by enhancing leaf thickness and dry matter content for better stress tolerance and by diminishing leaf area and chlorophyll content to mitigate leaf damage from intense sunlight. Conversely, at lower altitudes, the plant increases leaf area and chlorophyll content to facilitate organic matter accumulation.
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图 1 准噶尔山楂叶片功能性状间相关性分析
Figure 1. Correlation analysis of leaf functional traits in Crataegus songarica
*：P<0.05；**：P<0.01. Same below.
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图 2 准噶尔山楂叶片解剖结构随海拔梯度的变化趋势
不同字母表示不同海拔各参数差异显著（P<0.05）。
Figure 2. Variations in leaf anatomical structures in Crataegus songarica with altitude
Different letters indicate significant differences in parameters at different altitudes (P<0.05).
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图 3 准噶尔山楂叶片解剖结构相关性分析
Figure 3. Correlation analysis of leaf anatomical structures in Crataegus songarica
表 1 样地基本信息
Table 1 Basic information of sample plots
样地号
Sampling site海拔
Altitude / m纬度
Latitude / N经度
Longitude / E坡度
Slope / °1 1 156 44°22′20.19″ 80°57′43.15″ 13.89 2 1 180 44°22′20.08″ 80°57′40.26″ 16.13 3 1 195 44°22′20.23″ 80°57′41.51″ 17.98 4 1 209 44°22′20.22″ 80°57′44.35″ 18.85 5 1 236 44°22′20.08″ 80°57′44.25″ 19.56 6 1 274 44°22′20.94″ 80°57′45.13″ 20.23 7 1 312 44°22′20.57″ 80°57′49.05″ 20.63 8 1 358 44°22′21.26″ 80°57′50.12″ 21.05 9 1 376 44°22′21.27″ 80°57′51.26″ 22.47 10 1 410 44°22′22.28″ 80°57′52.41″ 23.50 11 1 430 44°22′26.43″ 80°58′15.16″ 24.65 12 1 483 44°22′23.30″ 80°57′54.02″ 24.78 13 1 510 44°22′22.31″ 80°57′55.33″ 25.01 14 1 538 44°22′23.32″ 80°57′56.21″ 25.58 15 1 580 44°22′21.29″ 80°57′59.58″ 27.83 16 1 612 44°22′22.80″ 80°57′43.92″ 29.92 17 1 623 44°22′23.59″ 80°57′45.83″ 30.97 18 1 630 44°23′13.15″ 80°57′37.41″ 31.65 
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表 2 不同海拔叶片功能性状变化特征
Table 2 Changes of leaf functional traits at different altitudes
海拔
A / m叶长
LL / cm叶宽
LW / cm叶长宽比
LL/LW叶面积
LA / cm2比叶面积
SLA / cm2/g比叶重
SLW / mg/cm2叶干物质含量
LDMC / g/g叶绿素含量
SPAD1 100~1 200 5.92±0.15a 5.73±0.16a 1.00±0.04c 17.56±0.97a 170.58±10.22a 6.03±0.47c 0.37±0.03c 42.01±1.51a 1 200~1 300 5.77±0.16ab 5.57±0.12a 1.02±0.04bc 16.19±0.81ab 140.34±8.75b 6.08±0.43c 0.46±0.03bc 40.34±0.87a 1 300~1 400 5.68±0.13ab 5.38±0.17ab 1.10±0.05abc 14.56±0.84bc 130.82±7.88bc 6.75±0.38c 0.49±0.04ab 39.73±0.81a 1 400~1 500 5.59±0.18ab 5.38±0.21ab 1.14±0.04ab 13.77±0.61c 118.24±10.37bcd 7.57±1.23c 0.49±0.03ab 39.28±1.33a 1 500~1 600 5.42±0.19ab 4.98±0.17bc 1.16±0.05a 13.25±0.90c 113.81±7.68cd 11.12±1.08b 0.50±0.05ab 38.93±1.15a 1 600~1 700 5.39±0.18b 4.61±0.12c 1.18±0.04a 7.88±0.59d 96.10±6.81d 14.17±0.69a 0.58±0.03a 34.89±0.89b 注：数据显示为平均值±标准误，同列不同字母表示不同海拔各参数差异显著（P<0.05）。 Notes: Data is shown as Mean±Standard error, and different letters in the same column indicate significant differences in parameters at different altitudes (P<0.05).
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表 3 准噶尔山楂叶片解剖结构可塑性指数
Table 3 Plasticity index of leaf anatomical structures in Crataegus songarica
指标 Index 可塑性指数 PI 指标 Index 可塑性指数 PI 栅栏组织厚度 0.26 角质层厚度 0.45 海绵组织厚度 0.28 叶片组织结构紧密度 0.29 主脉厚度 0.40 叶片组织结构疏松度 0.30 叶片厚度 0.27 栅海比 0.33 上表皮厚度 0.33 主脉突起度 0.49 下表皮厚度 0.38
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