The impacts of submergence duration and salinity on the germination and growth process of Aegiceras corniculatum radicles
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摘要: 潮位与盐度是影响桐花树幼苗生长发育的重要环境因子，在一定程度上决定桐花树胚胎能否成功着床与快速成苗。本文采用室内控制实验观测桐花树胚胎萌动期根系发育过程，进而探讨盐度、淹水时长、盐度−淹水时长交互作用对胚根萌发及生长的影响。结果表明：（1）模拟低低潮位淹水环境的完全浸泡胚胎24 h/d培养处理组，桐花树胚胎无法萌根。模拟中低潮位淹水环境的完全浸泡胚胎6 h/d + 浸泡胚根18 h/d培养处理组，胚胎平均萌根率最高，胚根最长。模拟高高潮位胚胎基部浸泡水体环境的浸泡胚根培养处理组，胚胎萌根率与平均萌根率最低，胚根数量最少。模拟高高潮位胚胎基部扎入潮滩环境的蛭石浅插培养处理组，胚根萌发最慢，萌根率最高，胚根最短、数量最多。（2）在盐度为0的环境中，培养13 d后胚根最长。在盐度为10的环境中，培养后的前11 d胚根最长、数量最多。在盐度为20的环境中，萌根率、平均萌根率以及胚根长度和数量明显偏低，胚根萌发与生长受到抑制。（3）淹水时长−盐度交互作用对桐花树胚根生长影响显著，若淹水时长、盐度处于桐花树胚胎萌根耐受范围内，淹水时长−盐度复合胁迫下胚胎依然能够正常萌根。人工培育桐花树幼苗，采用完全浸泡胚胎6 h/d + 浸泡胚根18 h/d，盐度为10培养胚根生长发育最优。研究结果可为潮间带桐花树幼苗人工培育与保护提供部分理论依据。Abstract: Tide level and salinity are crucial environmental factors that significantly impact the growth and development of Aegiceras corniculatum seedlings and to a certain extent, determine if their embryos could land successfully and develop into seedling rapidly. In this paper, the process of A. corniculatum embryos developing to seedings was observed by the indoor controlled experiments, during which, the effects of salinity, submergence duration, as well as the interaction between salinity and submergence duration on the germination and growth of radicles were discussed. The results showed that: (1) the embryos of the A. corniculatum were unable to sprout in the treatment group that the embryos were soaked completely for 24 h/d in a simulated lower low tidal level of flooded environment. For the treatment group that the embryos were soaked completely for 6 h/d and the radicles soaked for 18 h/d in a simulated low to medium tidal level of flooded environment, the highest mean rooting rates and the longest radicle of the embryos were produced. In the treatment group that the base of embryos were soaked in a simulated higher high tide flooded environment, the rooting rates and mean rooting rates of embryos were the lowest, accompanied by the least quantities of radicles. However, in the treatment group where the vermiculites were shallowly inserted into the bottom, which simulated a scene that the base of embryo was immersed within the tidal environment, it was found that the radicle germination was the slowest with the highest rooting rates, and the radicles were the shortest, along with the largest quantities. (2) In the salinity environment of 0, the radicles were the longest after a 11 days cultivation. In the salinity environment of 10, highest levels in both the length and quantities of radicles were observed in the first 13 days of embryo culture. In contrast, the rooting rate and the mean rooting rate of the embryos, as well as the length and quantity of radicle were significantly smaller in the salinity environment of 20, which indicated the germination and growth of radicle were inhibited in such salinity environment. (3) The interaction between submergence duration and salinity had a significant impact on the growth of the radicles of A. corniculatum. If both the submergence duration and salinity were within the tolerance ranges of the radicles of A. corniculatum, its embryos could still sprout normally under the stresses of different combination of submergence duration and salinity. Hence, the optimal way to artificially cultivatethe A. corniculatum seedlings, was that the embryo was completely soaked for 6 h/d and the radicle was soaked for 18 h/d in the salinity environment of 10. The study can provide some theoretical basis for the artificial cultivation and protection of A. corniculatum seedlings in the intertidal zone.
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Key words:
- Aegiceras corniculatum /
- submergence duration /
- salinity /
- radicle
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图 1 桐花树胚胎在潮滩上的分布状态
Fig. 1 The distribution of embryos of Aegiceras corniculatum on tidal flat
图 2 室内培养观测
a. 桐花树胚胎室内培养试验设计；b. 胚胎萌根观察
Fig. 2 Observation of indoor culture
a. Design of laboratory culture experiment for the embryos of Aegiceras corniculatum embryo; b. observation of embryo germination
图 3 淹水处理的桐花树胚胎萌根状态
Fig. 3 The state of sprouting roots of Aegiceras corniculatum in flooded treatment
图 4 淹水处理的桐花树胚根生长状态
胚根数量指同一处理胚胎萌根的平均数量，胚根长度指同一处理胚胎全部根系的平均长度
Fig. 4 Growth state of radicle of Aegiceras corniculatum in flooded treatment
The number of radicle refers to the average number of embryonic roots in the same treatment, and the length of radicle refers to the average length of all roots in the same treatment
图 5 盐度处理的桐花树胚胎萌根状态
Fig. 5 The state of sprouting roots of Aegiceras corniculatum in salinity treatment
图 6 盐度处理的桐花树胚根生长状态
胚根数量指同一处理胚胎萌根的平均数量，胚根长度指同一处理胚胎全部根系的平均长度
Fig. 6 Growth state of radicle of Aegiceras corniculatum in salinity treatment
The number of radicle refers to the average number of embryonic roots in the same treatment, and the length of radicle refers to the average length of all roots in the same treatment
图 7 淹水时长−盐度交互处理下胚根生长过程
胚根数量指同一处理胚胎萌根的平均数量，胚根长度指同一处理胚胎全部根系的平均长度
Fig. 7 Interactive treatment of submergence duration and salinity on the growth process of embryonic roots
The number of radicle refers to the average number of embryonic roots in the same treatment, and the length of radicle refers to the average length of all roots in the same treatment
表 1 室内培养盐度梯度试验设计
Tab. 1 Salinity gradient test design for indoor culture
编号 盐度 处理方式 E 0 自来水静放24 h F 10 盐度为10.9海水加自来水调配 G 20 盐度为10.9海水加海盐调配 
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表 2 淹水处理桐花树胚根长度及数量差异性分析
Tab. 2 Divergence analysis on the number and length of Aegiceras corniculatum embryos roots in submergence treatments
处理方式 7 d 14 d 21 d 28 d 34 d 胚根长度/cm B 0.33 ± 0.44b 1.09 ± 1.03b 1.60 ± 1.38a 2.02 ± 1.77a 2.45 ± 2.26a C 0.11 ± 0.30a 0.63 ± 0.72a 1.26 ± 0.97a 1.76 ± 1.22a 2.10 ± 1.40a D 0.06 ± 0.16a 0.52 ± 0.51a 1.14 ± 0.69a 1.55 ± 0.83a 2.05 ± 1.10a 胚根数量 B 0.50 ± 0.58b 1.38 ± 1.02a 1.77 ± 1.07a 2.77 ± 1.70a 3.31 ± 1.85a C 0.17 ± 0.38a 0.92 ± 0.97a 1.58 ± 1.02a 2.17 ± 1.05a 2.50 ± 1.06a D 0.15 ± 0.37a 1.04 ± 1.18a 2.65 ± 1.16b 3.69 ± 1.74b 5.04 ± 2.54b 注：不同小写字母表示不同处理组在相同时间下差异显著（p < 0.05）。
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表 3 盐度处理桐花树胚根长度及数量差异性分析
Tab. 3 Divergence analysis on the number and length of Aegiceras corniculatum embryonic roots in salinity treatments
处理方式 7 d 14 d 21 d 28 d 34 d 胚根长度/cm E 0.04 ± 0.14a 0.62 ± 0.51b 1.23 ± 0.96c 1.76 ± 1.34c 2.19 ± 1.63c F 0.17 ± 0.27b 0.54 ± 0.57b 0.81 ± 0.64b 1.09 ± 0.81b 1.24 ± 0.94b G 0.01 ± 0.05a 0.01 ± 0.06a 0.01 ± 0.06a 0.07 ± 0.18a 0.07 ± 0.19a 胚根数量 E 0.07 ± 0.25a 1.10 ± 1.09b 1.53 ± 1.17b 2.40 ± 1.89b 2.93 ± 2.42b F 0.37 ± 0.56b 1.00 ± 1.11b 2.00 ± 1.23b 2.60 ± 1.54b 3.07 ± 1.82b G 0.03 ± 0.18a 0.03 ± 0.18a 0.03 ± 0.18a 0.43 ± 1.30a 0.43 ± 1.30a 注：不同小写字母表示不同处理组在相同时间下差异显著（p < 0.05）。
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表 4 淹水时长−盐度交互处理下胚胎萌根状态
Tab. 4 Embryonic rooting status under the interaction of submergence duration and salinity
处理组 始萌根日数/d 平均萌根日数/d 萌根率/% 平均萌根率/% BE 6.0 11.9 90.0 70.0 CE 4.0 9.9 90.0 50.0 DE 8.0 11.1 80.0 50.0 BF 1.0 8.0 90.0 60.0 CF 5.0 12.0 80.0 40.0 DF 7.0 14.4 100.0 50.0 BG − − 0.0 0.0 CG 6.0 6.0 10.0 10.0 DG 23.0 24.7 30.0 20.0 注：BE处理组为完全浸泡胚胎6 h/d + 浸泡胚根18 h/d，盐度为0；CE处理组为浸泡胚根，盐度为0；DE处理组为蛭石浅插，盐度为0；BF处理组为完全浸泡胚胎6 h/d + 浸泡胚根18 h/d，盐度为10；CF处理组为处理组为浸泡胚根，盐度为10；DF处理组为蛭石浅插，盐度为10；BG处理组为完全浸泡胚胎6 h/d + 浸泡胚根18 h/d，盐度为20；CG处理组为浸泡胚根，盐度为20；DG处理组为蛭石浅插，盐度为20。“−”代胚根无萌动。
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表 5 淹水时长−盐度对胚根长度特征的双因素方差分析
Tab. 5 A bivariate anove of the effects of submergence duration and salinity on the characteristics of radical length
差异来源 7 d时胚根长度 14 d时胚根长度 21 d时胚根长度 28 d时胚根长度 34 d时胚根长度 F p F p F p F p F p 淹水−盐度处理 2.556 0.024 2.749 0.016 4.972 < 0.001 5.496 < 0.001 5.465 < 0.001
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表 6 淹水时长−盐度对胚根数量特征的双因素方差分析
Tab. 6 Bivariate anova of the effects of submergence duration and salinity on the quantitative characteristics of embryonic roots
差异来源 7 d时胚根数量 14 d时胚根数量 21 d时胚根数量 28 d时胚根数量 34 d时胚根数量 F p F p F p F p F p 淹水−盐度处理 2.566 0.023 1.500 0.185 3.914 < 0.001 2.127 0.056 2.404 0.032
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表 7 淹水时长−盐度处理下桐花树胚根生长信息主成分分析
Tab. 7 Principal component analysis of root growth information of Aegiceras corniculatum under the condition of submergence duration and salinity treatment
项目 第一主成分 第二主成分 特征根 3.87 1.66 方差贡献率/% 64.52 27.62 累积方差贡献率/% 64.52 92.14
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表 8 淹水时长−盐度处理下桐花树胚根生长指标的主成分得分
Tab. 8 Principal component scores of root growth indices of Aegiceras corniculatum under the condition of submergence duration and salinity treatment
处理 第一主成分（F1） 第二主成分（F2） 综合得分（F） AE −1.98 0.13 −1.24 AF −1.98 0.13 −1.24 AG −1.98 0.13 −1.24 BE 1.57 −1.13 0.70 BF 2.71 3.48 2.71 BG −1.98 0.13 −1.24 CE 2.01 −1.43 0.91 CF 1.27 0.03 0.83 CG −1.57 0.58 −0.85 DE 1.26 −1.44 0.41 DF 2.23 −0.54 1.29 DG −1.58 −0.08 −1.04
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