Responses of Barringtonia racemosa to Tidal Flooding
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摘要:目的 分析濒危半红树植物玉蕊对潮汐淹浸逆境的应答特性，为玉蕊在城市内河、滨湖、湿地恢复等景观应用中的适宜淹水时长选择提供科学依据。方法 以二年生玉蕊实生幼苗为材料，试验模拟半日潮，研究各幼苗在不同淹水时长的潮汐淹浸逆境中的植株形态、生理生化及矿质元素的应答特性。结果 （1）玉蕊幼苗经过250 d的潮汐淹浸逆境胁迫处理都能存活，其株高、叶片数、叶面积都显著低于对照植株，穿袋株数量、产生气生根及皮孔的数量均显著高于对照组。（2）玉蕊叶片叶绿素a、叶绿素总量（16 h·d−1除外）整体呈下降趋势；MDA含量整体呈上升趋势（14 h·d−1除外），并与脯氨酸、可溶性糖含量在淹水10 h·d−1时达到最大值，且均显著大于CK；（3）淹浸逆境胁迫促进叶片N、P、Fe元素的吸收，但抑制K、Cu元素的吸收。结论 叶面积、叶片磷元素含量、叶绿素总含量、侧根表面积、CAT活性、POD活性、可溶性蛋白含量是玉蕊应答淹浸胁迫的主要指标；玉蕊对淹水时长≤20 h·d−1的半日潮淹浸逆境有很强的耐受性和适应性，可配植于城市内河易淹水的岸域、滨湖绿带及淡水湿地等景观环境中。Abstract:Objective Responses of the endangered semi-mangrove Barringtonia racemosa to submergence in tides were studied to determine the adequacy of their planting for urban inland river, lakeside, and wetland restorations or other similar landscape projects.Method Two-year-old B. racemosa seedlings were used in the simulated semi-diurnal tide experiment. Morphology, physiology, biochemistry, and mineral contents of the plants in response to tidal submergence for varied durations were monitored.Result (1) The seedlings survived 250 d under the flooding treatments, but the plant height, leaf count, and leaf area were significantly lower, while the numbers of aerial roots, piercing plants, and lenticels significantly higher, than those of control. (2) The chlorophyll a and total chlorophyll in the leaves decreased continuously under flooding, except the 16 h·d−1 submergence treatment; MDA increased, except the 14 h·d−1 treatment; and, MDA, proline, and soluble sugars became significantly higher than those of CK and peaked under the 10 h·d−1 treatment. (3) The flooding increased the absorption of N, P, and Fe but inhibited that of K and Cu in the leaves.Conclusion The area, the contents of P, total chlorophyll, and soluble protein, and the activities of CAT and POD of leaves as well as the surface area of lateral roots were the major indicators that reflected the responses of B. racemosa to the flooding stress. The plants showed strong tolerance and adaptability to the adversity brought about by the semi-diurnal tides that lasted 20 h·d−1. Thus, B. racemosa could be adequately planted for landscaping at waterfront, lakeshore green belt, and/or freshwater wetland.
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Keywords:
- Barringtonia racemosa /
- semi-mangrove plant /
- endangered /
- tidal flooding /
- stress
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图 1 淹浸逆境对玉蕊株高、地径、叶片数及叶面积的影响
注：不同处理间玉蕊幼苗叶片的株高总增量（A）、地径总增长量（B）、叶片数变化量（C）及叶面积（D）。不同小写字母表示差异显著（P＜0.05），差异性分析为不同处理间的差异性比较，图2~7同。
Figure 1. Effects of flooding on plant height, underground trunk girth, leaf number, and leaf area of B. racemose
Note: Increases on plant height (A), increases on underground trunk girth (B), changes on leaf count (C), and changes on leaf area (D) of B. racemosa seedlings under treatments. Data with different lowercase letters indicate significant differences at P＜0.05 on different treatments. Same for the following.
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图 2 淹浸逆境对玉蕊气生根数、穿袋株数、皮孔数及主侧根表面积的影响
注：不同处理间玉蕊幼苗叶片的气生根数（A）、穿袋株数（B）、皮孔数（C）、主根表面积（D）及侧根表面积（E）。
Figure 2. Effects of flooding on numbers of aerial roots, piercing plants, and lenticels and surface area of main and lateral roots of B. racemose
Note: Number of aerial roots (A), number of plants penetrated breeding bag (B), number of pores (C), surface area of main roots (D), and surface area of lateral roots (E) of B. racemosa seedlings under treatments.
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图 5 淹浸逆境对玉蕊叶片渗透物质及细胞膜的影响
注：不同处理间玉蕊幼苗叶片的脯氨酸（A）、可溶性糖（B）、可溶性蛋白（C）及丙二醛的含量（D）。
Figure 5. Effects of flooding on osmotic substances and cell membrane of B. racemosa leaves
Note: Contents of proline (A), soluble sugar (B), soluble protein (C) and MDA (D) of B. racemosa seedlings under treatments.
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图 6 淹浸逆境对玉蕊叶片氮磷钾的影响
注：不同处理间玉蕊幼苗叶片的硝态氮（A）、有效磷（B）和有效钾的含量（C）。
Figure 6. Effects of flooding on N, P, and K in B. racemosa leaves
Note: Contents of nitrate N (A), effective P (B), and effective K (C) of B. racemosa seedlings under treatments.
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图 3 淹浸逆境对玉蕊叶绿素含量的影响
注：不同处理间玉蕊幼苗叶片的叶绿素a的含量（A）、叶绿素b的含量（B）及叶绿素的总含量（C）。
Figure 3. Effect of flooding on chlorophyll content of B. racemose
Note: Chlorophyll a (A), chlorophyll b (B), and total chlorophyll (C) of B. racemosa seedlings under treatments.
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图 4 淹浸逆境对玉蕊叶片酶系统的影响
注：不同处理间玉蕊幼苗叶片的SOD活性（A）、POD活性（B）及CAT活性（C）。
Figure 4. Effect of flooding on enzyme activities of B. racemosa leaves
Note: Activities of SOD (A), POD (B), and CAT (C) of B. racemosa seedlings under treatments.
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图 7 淹浸逆境对玉蕊叶片铁、铜离子的影响
注：不同处理间玉蕊幼苗叶片矿质元素的Fe含量（A）和Cu的含量（B）。
Figure 7. Effects of flooding on Fe and Cu ions in B. racemosa leaves
Note: Contents of Fe (A) and Cu (B) of B. racemosa seedlings under treatments.
表 1 玉蕊对淹浸逆境应答指标的主成分分析
Table 1 Principal component analysis on responses of B. racemosa to flooding
应答指标
Response indicators主成分矩阵
Principal component matrix a1 2 3 4 5 6 7 株高 Plant height 0.780 −0.212 0.119 0.027 −0.016 −0.079 −0.286 地径 Ground diameter −0.121 0.586 0.387 0.149 0.469 0.149 0.188 叶片数 Number of blades 0.770 −0.107 0.202 0.007 0.031 −0.263 0.010 叶面积 leaf area 0.822 0.000 −0.329 0.203 0.007 0.140 0.004 皮孔数 Number of lenticels −0.703 −0.398 −0.103 −0.366 0.025 0.129 −0.196 主根表面积 Surface area of taproot 0.370 0.159 0.269 0.563 −0.430 −0.014 −0.316 侧根表面积 Surface area of lateral roots 0.132 0.250 0.460 0.703 −0.082 0.353 0.090 叶绿素 a Chlorophyll a 0.599 0.125 −0.153 −0.429 0.335 0.160 0.028 叶绿素 b Chlorophyll b −0.287 0.049 −0.820 0.404 −0.074 −0.056 0.098 叶绿素总量 Total chlorophyll 0.398 −0.051 0.728 −0.488 0.081 0.068 −0.095 SOD活性 SOD activity 0.573 0.194 −0.572 −0.082 0.179 0.272 0.093 POD活性 POD activity −0.027 −0.381 0.201 −0.157 −0.604 0.355 0.433 CAT活性 CAT activity −0.334 0.247 0.057 0.242 0.538 −0.497 0.153 脯氨酸含量 Proline content −0.098 0.602 0.130 −0.679 −0.207 0.062 0.076 可溶性糖含量 Soluble sugar content −0.155 0.638 0.493 0.181 0.022 −0.114 −0.216 可溶性蛋白含量 Soluble protein content −0.346 −0.070 0.096 0.123 0.509 0.691 −0.104 丙二醛含量 Malondialdehyde content −0.164 0.535 −0.323 −0.279 −0.139 0.189 −0.589 硝态 N含量 Nitrate N content −0.549 −0.569 0.227 0.270 −0.101 0.145 −0.013 P含量 P content 0.111 0.877 0.009 −0.075 −0.105 0.148 0.289 K含量 K content 0.477 −0.660 0.155 −0.197 0.145 −0.097 0.168 Fe含量 Fe content −0.449 0.561 0.066 −0.235 −0.284 −0.245 0.126 Cu含量 Cu content 0.418 0.585 −0.230 0.146 −0.167 0.057 0.095 特征值 Characteristic value 4.679 4.155 2.71 2.467 1.695 1.385 1.05 贡献率 Contribution rate/% 21.267 18.888 12.32 11.211 7.707 6.297 4.772 累积贡献率 Cumulative contribution rate/% 21.267 40.155 52.475 63.686 71.393 77.691 82.462 注：提取方法：主成分；a：已提取了7个成分。
Note: Extraction method: principal component; a:7 principle components extracted.
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