种子发芽生态研究
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摘要
生活史对策是植物种群生态研究的重要领域,而种子发芽是植物生活史的重要阶段,因此,研究种子发芽规律及对策,有助于认识和阐明物种进化及其生态适应特征。本论文以松嫩草原区植物为主要对象,研究了种子对恒温和每日增降温的发芽响应,比较了变温梯度模拟的全球增温对C3、C4植物种子发芽的影响,研究了植物发芽格局及在不同温度下的变化,并探讨了植物对盐和水势的发芽响应及盐和温度交互作用的发芽响应机制。另外研究了子叶对幼苗生长的光合贡献。
根据恒温对种子发芽影响的结果,在积温模型的基础上提出了修正积温模型(modified thermal time model),补充了发芽速率恒定部分的拟合公式。C3植物的发芽速率明显低于C4植物。根据积温模型预测的发芽最低温和积温常数,C3植物的平均发芽最低温低于C4植物,但差异不显著,平均积温需要明显高于C4植物,接近显著。本研究指出,种子的发芽格局可以分为五类:速萌型、延迟型、稳定型、正态型、延迟速萌型。多数一年生植物都属于速萌型,分布于草甸草原的多年生植物属于延迟速萌型、正态型或稳定型。大部分植物在不同温度下的发芽格局是有变化的,这是对环境的一种适应对策。
在变温梯度模拟的全球增温条件下,C3和C4植物的发芽特点均发生改变。C3植物对高温敏感,C4植物对低温敏感。从几个变温梯度的比较看,全球增温对种子发芽的影响不是温差变化在起作用,而是高温或低温部分的温度在起作用。野外条件下,松嫩草原区更多的植物种子在春季发芽而不是秋季。室内实验证明,在春季和秋季较低的温度范围5-10和10-15℃内,每日增加的温度下的发芽显著高于每日降低的温度下,植物春季发芽是自然选择下来的一种适应策略。
虎尾草和马唐在不同温度下发芽速率和盐度之间成曲线关系,我们认为可能存在一些生理机制,使得中间盐度和高盐浓度下的渗透胁迫降低,发芽速率明显增加,产生了曲线而非直线的关系。进一步的实验表明,PEG和NaCl两种水势处理相比,种子能在更低的NaCl引起的水势下发芽且发芽比在等渗的PEG处理中快。实验用植物种子的干重随时间降低,NaCl处理中发芽种子的含水量、钠离子浓度和种子溶液钠浓度显著高于PEG处理中。NaCl处理中发芽种子和未发芽种子钠离子浓度和种子溶液钠浓度均随水势降低而增加。种子含水量、钠离子浓度、种子溶液钠浓度随时间变化而线性增加。根据实验结果提出盐度模型:θS = (Sm– S) t,其中θS是盐度常数,Sm是种子能够发芽的最高盐度值,S是外界盐浓度,t是某个盐浓度下的发芽时间。将种子对盐的响应划分为四个阶段。第Ⅰ阶段主要是渗透效应起负作用;第Ⅱ阶段是渗透效应和离子效应共同起作用,离子效用的正作用强于渗透效应的负作用;第Ⅲ阶段等渗的PEG中已经没有种子发芽,离子效应的正作用远远高于渗透效应的负作用;第Ⅳ阶段是离子效应逐渐开始起离子毒害的负作用。
子叶光合贡献研究表明,种子吸胀时去除子叶,真叶不能生长或很快死亡,除了呼吸大于同化外,可能还涉及到激素类型的信号缺失。子叶展开前去除子叶引起幼苗个体发育补偿,根茎比降低。真叶展开后去除子叶延迟了幼苗生长发育,表明子叶在真叶长出后对幼苗仍然有光合贡献。小种子物种对子叶光合作用依赖很强,实现了早期幼苗生长过程中的较高的相对生长速率。本研究量化了子叶光合贡献占总贡献的比例,为物种竞争策略和对不同生境的适合度提供了有用指示。
本论文结果对于预测植物发芽出土、全球变暖对植物生长分布的影响,开发利用野生植物资源策略均有理论指导意义和生产实践价值。
Life-history strategy is the significant research area of plant population ecology. Seed germination is the critical stage of plant life history. Therefore, research on seed germination rule and strategy helps to know and elucidate plant evolution and ecological adaptation traits. Plants in Songnen grassland area were used to test the responses of seed germination to constant temperature and diurnal increased or decreased temperature, the effects of global warming simulated by alternating temperature regimes on seed germination of C3 and C4 species, plant germination pattern and the changes under different constant temperature. The mechanisms of germination responses to NaCl and PEG and the interaction between salt and temperature were also studied. Photosynthetic contribution of cotyledons to seedling growth was evaluated as well.
According to the results of constant temperature effects, we proposed the modified thermal time model on the basis of thermal time model and supplemented an equation describing germination rate constant. Germination rate of C3 species was lower than C4 species. According to thermal time model, the average base temperature of C3 species was lower than C4 species, but not significantly. The average thermal time constant of C3 species was higher than C4 species, which approached significant difference. In this study, plant seed germination pattern was divided into five categories: quick germination, delayed germination, steady germination, normal germination and delayed-quick germination. Most annual species belonged to quick germination pattern. Perennial C3 species, which distributed to meadow, belonged to delayed-quick germination pattern, normal germination pattern or steady germination pattern. Seed germination pattern of most species changed with temperature, which was an adaptive strategy to environments.
Under global warming simulated by alternating temperature regimes conditions, germination traits of C3 and C4 species changed. C3 species was sensitive to high temperature and C4 species was sensitive to low temperature. By comparison of germination results in all the alternating temperature regimes, we found that high or low temperature, not amplitude acted an important role in the effects of global warming on seed germination. Field experiment showed that more plants germinated in spring, not autumn in Songnen grassland. Germination experiment exhibited that significant differences in germination between diurnal increased and decreased temperature in the spring and autum low ranges 5-10 and 10-15oC, with the former higher. Germination in spring is an adaptive strategy by natural selection. The relationship between germination rate and salinity under different temperatures of C. virgata and D. sanguinalis was curvilinear. We thought there may be some physiological mechanisms, which reduced osmotic stress in the middle and high salinities. This led to the obviously increasing of the germination rate. Further experiments showed that compared with PEG treatment, seeds can germinate in lower water potential induced by NaCl and germinate faster. Seed dry weight decreased with the time. Water content, seed sodium concentration and seed solution sodium concentration of NaCl treatments were significantly higher than the isotonic PEG treatments. Seed sodium concentration and seed solution sodium concentration of germinating and un-germinating seeds in NaCl treatments increased as the water potential decreased. Water content, seed sodium concentration and seed solution sodium concentration increased linearly with time.
According to the results, we proposed salinity model:θS = (Sm– S) t, in whichθS is salinity constant, Sm is the maximum salinity above which seed can’t germinate, S is the external salinity, and t is germination time. The germination response of seed to salt was divided into four stages. The first stage mainly involved negative osmotic effect. In the second stage, ionic effect and osmotic effect existed together, with ionic effect stonger. In the third stage, no seed germinated in PEG treatment and ionic positive effect was largely higher than osmotic effect. In the fourth stage, ionic effect gradually began to harm the seed.
Cotyledon photosynthetic contribution experiments showed that loss of cotyledons in seed imbibition stage, first leaf development did not proceed following early loss of cotyledons, but the reason did not appear to be an excess of respiration over assimilation, so a hormonal type signal may be involved; loss of cotyledons prior to cotyledon expansion, and so presumably before any substantive photosynthesis, induced seedling ontogenetic compensation (increase in shoot:root ratio); loss of cotyledons at the first leaf stage of seedling growth delayed seedling development without changing seedling ontogeny, indicating an ongoing photosynthetic contribution of cotyledons. The data highlighted the way in which almost total reliance on the photosynthetic role of cotyledons in small seeded species confered the possibility to achieve a very high relative growth rate during early seedling development, and raised the possibility that quantitative determination of the ratio between the photosynthetic contribution of cotyledons to seedling development and total (photosynthetic plus reserve) contribution may provide a useful indicator of species competitive strategy and fitness for different habitats.
This study has theoretical significances and practical value for predicting plant germination and emergence, testing effects of global warming on plant distribution characteristics and developing and utilizing wild plant resources.
根据恒温对种子发芽影响的结果,在积温模型的基础上提出了修正积温模型(modified thermal time model),补充了发芽速率恒定部分的拟合公式。C3植物的发芽速率明显低于C4植物。根据积温模型预测的发芽最低温和积温常数,C3植物的平均发芽最低温低于C4植物,但差异不显著,平均积温需要明显高于C4植物,接近显著。本研究指出,种子的发芽格局可以分为五类:速萌型、延迟型、稳定型、正态型、延迟速萌型。多数一年生植物都属于速萌型,分布于草甸草原的多年生植物属于延迟速萌型、正态型或稳定型。大部分植物在不同温度下的发芽格局是有变化的,这是对环境的一种适应对策。
在变温梯度模拟的全球增温条件下,C3和C4植物的发芽特点均发生改变。C3植物对高温敏感,C4植物对低温敏感。从几个变温梯度的比较看,全球增温对种子发芽的影响不是温差变化在起作用,而是高温或低温部分的温度在起作用。野外条件下,松嫩草原区更多的植物种子在春季发芽而不是秋季。室内实验证明,在春季和秋季较低的温度范围5-10和10-15℃内,每日增加的温度下的发芽显著高于每日降低的温度下,植物春季发芽是自然选择下来的一种适应策略。
虎尾草和马唐在不同温度下发芽速率和盐度之间成曲线关系,我们认为可能存在一些生理机制,使得中间盐度和高盐浓度下的渗透胁迫降低,发芽速率明显增加,产生了曲线而非直线的关系。进一步的实验表明,PEG和NaCl两种水势处理相比,种子能在更低的NaCl引起的水势下发芽且发芽比在等渗的PEG处理中快。实验用植物种子的干重随时间降低,NaCl处理中发芽种子的含水量、钠离子浓度和种子溶液钠浓度显著高于PEG处理中。NaCl处理中发芽种子和未发芽种子钠离子浓度和种子溶液钠浓度均随水势降低而增加。种子含水量、钠离子浓度、种子溶液钠浓度随时间变化而线性增加。根据实验结果提出盐度模型:θS = (Sm– S) t,其中θS是盐度常数,Sm是种子能够发芽的最高盐度值,S是外界盐浓度,t是某个盐浓度下的发芽时间。将种子对盐的响应划分为四个阶段。第Ⅰ阶段主要是渗透效应起负作用;第Ⅱ阶段是渗透效应和离子效应共同起作用,离子效用的正作用强于渗透效应的负作用;第Ⅲ阶段等渗的PEG中已经没有种子发芽,离子效应的正作用远远高于渗透效应的负作用;第Ⅳ阶段是离子效应逐渐开始起离子毒害的负作用。
子叶光合贡献研究表明,种子吸胀时去除子叶,真叶不能生长或很快死亡,除了呼吸大于同化外,可能还涉及到激素类型的信号缺失。子叶展开前去除子叶引起幼苗个体发育补偿,根茎比降低。真叶展开后去除子叶延迟了幼苗生长发育,表明子叶在真叶长出后对幼苗仍然有光合贡献。小种子物种对子叶光合作用依赖很强,实现了早期幼苗生长过程中的较高的相对生长速率。本研究量化了子叶光合贡献占总贡献的比例,为物种竞争策略和对不同生境的适合度提供了有用指示。
本论文结果对于预测植物发芽出土、全球变暖对植物生长分布的影响,开发利用野生植物资源策略均有理论指导意义和生产实践价值。
Life-history strategy is the significant research area of plant population ecology. Seed germination is the critical stage of plant life history. Therefore, research on seed germination rule and strategy helps to know and elucidate plant evolution and ecological adaptation traits. Plants in Songnen grassland area were used to test the responses of seed germination to constant temperature and diurnal increased or decreased temperature, the effects of global warming simulated by alternating temperature regimes on seed germination of C3 and C4 species, plant germination pattern and the changes under different constant temperature. The mechanisms of germination responses to NaCl and PEG and the interaction between salt and temperature were also studied. Photosynthetic contribution of cotyledons to seedling growth was evaluated as well.
According to the results of constant temperature effects, we proposed the modified thermal time model on the basis of thermal time model and supplemented an equation describing germination rate constant. Germination rate of C3 species was lower than C4 species. According to thermal time model, the average base temperature of C3 species was lower than C4 species, but not significantly. The average thermal time constant of C3 species was higher than C4 species, which approached significant difference. In this study, plant seed germination pattern was divided into five categories: quick germination, delayed germination, steady germination, normal germination and delayed-quick germination. Most annual species belonged to quick germination pattern. Perennial C3 species, which distributed to meadow, belonged to delayed-quick germination pattern, normal germination pattern or steady germination pattern. Seed germination pattern of most species changed with temperature, which was an adaptive strategy to environments.
Under global warming simulated by alternating temperature regimes conditions, germination traits of C3 and C4 species changed. C3 species was sensitive to high temperature and C4 species was sensitive to low temperature. By comparison of germination results in all the alternating temperature regimes, we found that high or low temperature, not amplitude acted an important role in the effects of global warming on seed germination. Field experiment showed that more plants germinated in spring, not autumn in Songnen grassland. Germination experiment exhibited that significant differences in germination between diurnal increased and decreased temperature in the spring and autum low ranges 5-10 and 10-15oC, with the former higher. Germination in spring is an adaptive strategy by natural selection. The relationship between germination rate and salinity under different temperatures of C. virgata and D. sanguinalis was curvilinear. We thought there may be some physiological mechanisms, which reduced osmotic stress in the middle and high salinities. This led to the obviously increasing of the germination rate. Further experiments showed that compared with PEG treatment, seeds can germinate in lower water potential induced by NaCl and germinate faster. Seed dry weight decreased with the time. Water content, seed sodium concentration and seed solution sodium concentration of NaCl treatments were significantly higher than the isotonic PEG treatments. Seed sodium concentration and seed solution sodium concentration of germinating and un-germinating seeds in NaCl treatments increased as the water potential decreased. Water content, seed sodium concentration and seed solution sodium concentration increased linearly with time.
According to the results, we proposed salinity model:θS = (Sm– S) t, in whichθS is salinity constant, Sm is the maximum salinity above which seed can’t germinate, S is the external salinity, and t is germination time. The germination response of seed to salt was divided into four stages. The first stage mainly involved negative osmotic effect. In the second stage, ionic effect and osmotic effect existed together, with ionic effect stonger. In the third stage, no seed germinated in PEG treatment and ionic positive effect was largely higher than osmotic effect. In the fourth stage, ionic effect gradually began to harm the seed.
Cotyledon photosynthetic contribution experiments showed that loss of cotyledons in seed imbibition stage, first leaf development did not proceed following early loss of cotyledons, but the reason did not appear to be an excess of respiration over assimilation, so a hormonal type signal may be involved; loss of cotyledons prior to cotyledon expansion, and so presumably before any substantive photosynthesis, induced seedling ontogenetic compensation (increase in shoot:root ratio); loss of cotyledons at the first leaf stage of seedling growth delayed seedling development without changing seedling ontogeny, indicating an ongoing photosynthetic contribution of cotyledons. The data highlighted the way in which almost total reliance on the photosynthetic role of cotyledons in small seeded species confered the possibility to achieve a very high relative growth rate during early seedling development, and raised the possibility that quantitative determination of the ratio between the photosynthetic contribution of cotyledons to seedling development and total (photosynthetic plus reserve) contribution may provide a useful indicator of species competitive strategy and fitness for different habitats.
This study has theoretical significances and practical value for predicting plant germination and emergence, testing effects of global warming on plant distribution characteristics and developing and utilizing wild plant resources.
引文
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