德夯岩溶河谷异质生境中半蒴苣苔的水分生理生态适应性

王晨瑶; 张婷; 曾德武; 李龙; 叶庆子; 田向荣

doi:10.11932/karst2024y031

德夯岩溶河谷异质生境中半蒴苣苔的水分生理生态适应性

doi: 10.11932/karst2024y031

王晨瑶^{1, 2,},
张婷^{1, 2},
曾德武¹,
李龙¹,
叶庆子³,
田向荣^{1, 2, 4, ,}

1.
吉首大学生物资源与环境科学学院, 湖南吉首 416000
2.
武陵山区植物资源保护与利用湖南省高校重点实验室, 湖南吉首 416000
3.
湘西世界地质公园, 湖南吉首 416000
4.
杜仲综合利用技术国家地方联合工程实验室, 湖南吉首 416000

基金项目: 国家自然科学基金项目（31660119）。

详细信息

作者简介:
王晨瑶（1998－），女，硕士研究生，研究方向为植物生态学；E-mail：120270772@qq.com

通讯作者:
田向荣（1979－），男，教授，博士，研究方向为植物生理生态，种子生物学；E-mail：dna_rna_pro@126.com。

中图分类号: Q948
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出版历程
- 收稿日期: 2023-10-13
- 录用日期: 2024-02-18
- 修回日期: 2024-01-18
- 网络出版日期: 2024-08-15
- 刊出日期: 2024-10-31

Water eco-physiological adaptability of Hemiboea subcapitata in heterogeneous habitats in the Dehang karst valley

WANG Chenyao^{1, 2
,},
ZHANG Ting^{1, 2},
ZENG Dewu¹,
LI Long¹,
YE Qingzi³,
TIAN Xiangrong^{1, 2, 4
, ,}

1.
College of Biology and Environmental Sciences, Jishou University, Jishou, Hunan 416000, China
2.
Key Laboratory of Plant Resources Conservation and Utilization in Colleges and Universities of Hunan Province, Jishou, Hunan 416000, China
3.
Xiangxi UNESCO Global Geopark, Jishou, Hunan 416000, China
4.
National & Local United Engineering Laboratory of Integrative Utilization Technology of Eucommia Ulmoides, Jishou, Hunan 416000, China

摘要

摘要: 以湘西世界地质公园德夯岩溶河谷内克隆植物半蒴苣苔（Hemiboea subcapitata）为材料，测定其生长、形态与光合水分生理指标，探究其对德夯岩溶台地河谷演化中形成的三类异质生境(峡谷岩壁表面、风化碎裂岩表和河岸有机质层)的水分生理生态适应性。结果显示：(1)异质生境中半蒴苣苔单(分)株叶片自然含水量仅有较小差异(< 2%)，而相对含水量和自然饱和亏则无显著差异；异质生境中叶片水分需求随基质含水量降低而明显增加，但叶片水势也随之显著降低。(2)单株生长差异大但群体累积生物量无明显差异。如峡谷岩壁表面单株的株高和叶面积均显著高于河岸有机质层单株，但群体累积株高、累积叶面积差异不显著。(3)单株外部形态变化显著，匍匐茎长度、直径和根着生密度均呈现峡谷岩壁表面>风化碎裂岩表>河岸有机质层的趋势；峡谷岩壁表面单株叶片比河岸有机质层单株平均增厚近60 μm，比叶面积则降低至其78.2%，气孔密度则显著降低至其66%。(4)光合生理指标中，叶片气孔限制值随叶片水势降低而显著增加，同时气孔导度明显提升，故而胞间CO₂浓度并无明显变化，但峡谷岩壁表面单株叶片的净光合速率仅达到河岸有机质层单株叶片的69.6%。(5)水分亏缺最多的峡谷岩壁表面单株叶片水分利用效率却最低(3.029 ± 0.461 μmol CO₂·mmol⁻¹ H₂O)，仅为河岸有机质层单株的73%。表明，半蒴苣苔可通过自身水分维持机制来保证岩溶异质生境中单株水分的相对稳定，并以增加根系密度、匍匐茎长度、叶片厚度和重量，减少气孔密度等形态可塑性来适应干旱的岩溶岩壁生境，通过增加气孔导度保持蒸腾拉力来维持胞间CO₂浓度，以高耗水来维持生长状态相对稳定的方式达成对岩溶河谷水分异质性生境的适应。
- 岩溶河谷 /
- 半蒴苣苔 /
- 水分利用效率 /
- 水分生理生态适应
Abstract: The Dehang canyon of Xiangxi UNESCO Global Geopark is cut by the runoff of streams. As a unique karst landform, it is an ideal area for studying plant species diversity and ecological adaptability. Collecting the clone plants, Hemiboea subcapitata, as samples from the Dehang karst valley in Xiangxi Global Geopark, this study measured their growth, morphology, and photosynthetic and water physiological indicators. On this basis, this study explored the water eco-physiological adaptability of the plants to three types of heterogeneous habitats formed during the evolution of the Dehang karst valley. These three habitats are canyon karst walls slightly weathering after water erosion without soil covering, bank weathered rocks with little soil covering and bank soil organic horizons highly weathering with soil covering.The results show as follows. (1) There was only a small difference (< 2%) in the natural water content in ramet leaves of H. subcapitata in heterogeneous habitats, while there is no significant difference in terms of relative water content and natural saturation deficit. It is proved that they not only have a strong water maintenance mechanism but also have no obvious difference in their recovery ability after drought. The lowest water potential of canyon karst walls was -1.630 ± 0.047 Mpa, and the highest water potential of bank soil organic horizons was −0.705 ± 0.025 Mpa. It is obvious that, in heterogeneous habitats, the water demand of ramet leaves increased significantly with the decrease of the matrix water content, but the water potential of leaves decreased significantly. (2) There was a significant difference in ramet growth but no significant difference in total biomass. For example, the plant heights and leaf areas of ramets on the surfaces of canyon karst walls were significantly higher than those in bank soil organic horizons, but there was no significant difference in the cumulative height and leaf area of total population. (3) The morphological changes of ramets were significant. Values of lengths, diameters and root densities of stolons were listed as: canyon karst walls>bank weathered rocks>bank soil organic horizons. The leaf thickness on canyon karst walls was nearly 60 μm thicker than that in bank soil organic horizons, while the specific leaf area (SLA) decreased to 78.2%, and the stomatal density significantly reduced to 66%. The highest stomatal density was 2,299 ± 158 mm² in the habitats of bank soil organic horizons, and the lowest stomatal density was 1,518 ± 98 mm² in the habitats of canyon karst walls. (4) In terms of photosynthetic parameters, stomatal limit values of leaves increased significantly with the decrease of leaf water potential in different habitats, but stomatal conductance increased significantly at the same time, so the intercellular CO₂ concentration did not change significantly. However, the net photosynthesis of ramet leaves on bank soil organic horizons only reached 69.6% of that in bank soil organic horizons. (5) The water use efficiency (WUE) of the habitats of bank soil organic horizons was the highest (4.134 ± 0.333 μmol CO₂·mmol⁻¹ H₂O). When the water deficit of leaves on canyon karst walls was the highest, the water use efficiency was the lowest (3.029 ± 0.461 μmol CO₂·mmol⁻¹ H₂O), only 73% of that in bank soil organic horizons. These results indicate that H. subcapitata can ensure the relative water stability of ramets in karst heterogeneous habitats through its own water maintenance mechanism. H. subcapitata can be adapted to the habitats of arid karst rock walls by increasing root density, stolon length, leaf thickness and weight, and by reducing stomatal density and other morphological plasticity. It can also keep intercellular CO₂ concentration by increasing stomatal conductance to maintain transpiration pull. At last, it can be adapted to water heterogeneous habitats in karst river valleys by relatively stable growth with higher water consumption.
- karst valley /
- Hemiboea subcapitata /
- water use efficiency (WUE) /
- water eco-physiological adaptability

HTML

图 1 三种生境半蒴苣苔的含水量及水分饱和亏缺

Figure 1. Water content and water saturation deficit of H. subcapitata in three habitats

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图 2 三种生境半蒴苣苔的平均植株高度、密度及累积植株高度

Figure 2. Average plant height, density and cumulative plant height of H. subcapitata in three habitats

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图 3 峡谷岩壁表面生境半蒴苣苔及三种生境半蒴苣苔1-5层叶片

Figure 3. H. subcapitata on canyon karst walls and 1-5 layers of leaves of H. subcapitata in three habitats

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图 5 三种生境半蒴苣苔的叶片及根茎

Figure 5. Roots, stems and leaves of H. subcapitata in three habitats

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图 4 三种生境半蒴苣苔的叶片解剖结构和根茎形态

注：A. 峡谷岩壁表面半蒴苣苔叶片的横切 B. 风化碎裂岩表半蒴苣苔叶片的横切 C. 河岸有机质层半蒴苣苔叶片的横切 D. 峡谷岩壁表面半蒴苣苔叶片的气孔密度 E. 风化碎裂岩表半蒴苣苔叶片的气孔密度 F. 河岸有机质层半蒴苣苔叶片的气孔密度 G. 峡谷岩壁表面半蒴苣苔的根茎形态 H. 风化碎裂岩表半蒴苣苔的根茎形态 I. 河岸有机质层半蒴苣苔的根茎形态

Figure 4. Leaf anatomy and rhizome morphology of H. subcapitata in three habitats

Note: A. transection of H. subcapitata leaf on canyon karst walls; B. transection of H. subcapitata leaf on bank weathered rocks; C. transection of H. subcapitata leaf in bank soil organic horizons; D. stomatal density of H. subcapitata leaf on canyon karst walls; E: stomatal density of H. subcapitata leaf on canyon karst walls; F. stomatal density of H. subcapitata leaf on canyon karst walls; G. rhizome morphology of H. subcapitata on canyon karst walls; H. rhizome morphology of H. subcapitata on bank weathered rocks; I. rhizome morphology of H. subcapitata in bank soil organic horizons

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表 1 不同生境环境因子的差异

Table 1. Differences of environmental factors in different habitats

	光照强度 Light illumination /Lux	空气温度 Air temperature / ℃	空气湿度 Air humidity /%	基质含水量 Moisture content of matrix /%
峡谷岩壁表面	9 053.52±10.03a	22.83 ± 0.34a	64.02 ± 1.60c	0.36 ± 0.05c
风化碎裂岩表	9 054.26±9.96a	23.03 ± 0.34b	70.47 ± 1.02b	5.81 ± 0.49b
河岸有机质层	850.30±8.63b	24.04 ± 0.30c	76.04 ± 1.12a	32.54 ± 0.38a
注：同列不同小写字母表示差异显著性（P<0.05），其中基质含水量均为表层基质（0~2 cm）含水量。
Note: Different lowercase letters in the same column indicate significant difference (P<0.05), and the matrix water content is surface matrix (0–2 cm) water content.

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表 2 三种生境半蒴苣苔光合气体交换参数及水分利用效率

Table 2. Photosynthetic gas exchange parameters and water use efficiency of H. subcapitata in three habitats

指标 Index	峡谷岩壁表面	风化碎裂岩表	河岸有机质层
水势/MPa	−1.630 ± 0.047c	−1.015 ± 0.041b	−0.705 ± 0.025a
气孔密度/mm²	1518.00 ± 98.00c	2198.24 ± 102.53b	2299.00 ± 158.00a
气孔导度/mol H₂O·m⁻²·s⁻¹	0.178 ± 0.045a	0.175 ± 0.069a	0.154 ± 0.085b
蒸腾速率/mmol H₂O·m⁻²·s⁻¹	1.601 ± 0.108a	1.739 ± 0.173a	1.704 ± 0.217a
胞间CO₂浓度/μmol CO₂·mol⁻¹	420.988 ± 48.439a	414.436 ± 36.612a	394.325 ± 40.597a
气孔限制值/Ls	0.936 ± 0.031a	0.876 ± 0.086b	0.804 ± 0.080b
净光合速率/μmol CO₂·m⁻²·s⁻¹	4.820 ± 0.537b	6.075 ± 0.468a	6.923 ± 1.663a
水分利用效率/μmol CO₂·mmol⁻¹ H₂O	3.029 ± 0.461b	3.519 ± 0.377ab	4.134 ± 0.333a
注：同行不同小写字母表示差异显著性（P<0.05）。 Note: Different lowercase letters in the same line indicate significant difference (P<0.05).

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