羊草克隆整合对碱环境异质性的响应
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摘要
克隆植物是一个独特的植物类群,它广泛分布于几乎所有类型的生态系统中,并在许多生态系统如草原、湿地、森林中占据优势地位,在植被恢复中发挥着举足轻重的作用。在长期的进化过程中,克隆植物形成了有效利用异质性资源以及抵御各种局部胁迫的生态适应对策—克隆整合。羊草(Leymus chinensis (Trin.) Tzvel.)属于根茎型克隆植物,是松嫩碱化羊草草甸的优势种,也是植被恢复的主要牧草之一,但是对碱影响克隆整合的机制尚不十分清楚。该研究以羊草为研究对象,采用野外试验和室内模拟的方法,研究切断根茎、环境因素(模拟碱胁迫)和不同发育阶段(生根前和生根后)对克隆整合的影响,探讨不同碱环境异质性和植株不同发育阶段克隆整合的强度、方向和克隆整合耗益之间的权衡关系,进一步分析羊草对碱环境的克隆整合机制。主要结论如下：
(1)异质性碱胁迫明显影响植物的克隆整合作用效果。在重度碱胁迫处理条件下,克隆整合发挥正向作用,未受碱胁迫的母株和遭受碱胁迫的子株之间发生明显的水分传递现象,母株对子株的水分整合提高了子株的防御能力,缓解了碱胁迫对子株的生理伤害,从而相应地提高了羊草的生物量。但在同质和低碱异质性处理下,克隆整合的这种帮助作用不明显,说明克隆整合的强度随碱异质性的增大而增大。克隆植物会根据碱生境条件作出应对策略,产生形态和生理上的可塑性反应,在羊草地下器官生物量分配格局的结果表明,羊草根、根茎和芽对克隆整合表现出不同的可塑性响应,克隆整合作用使得芽的存活数明显增加,根茎的伸长不明显和克隆整合并未使植物根重有所提高,甚至出现下降的趋势。通过对母株和子株生物量的分析得出,克隆整合虽然使母株生物量有所降低,但子株从中获得的益处远远大于母株的损耗,因此对母株和子株的组合来说克隆整合是有益的。克隆整合是根茎克隆植物抵御碱胁迫的重要适应对策。
(2)同位素标记试验证明,克隆植物的母株通过连接的根茎向子株输送氮素,从而减缓了碱胁迫对子株的影响,克隆整合增加了子株的高度和地上地下生物量。在高碱胁迫下,子株的不同发育阶段对克隆整合的强度有一定的影响,母株向子株传输的氮营养在子株生根后比生根前有一定减少,但子株的营养供给主要依赖于母株。通过15N丰度的计算结果表明,母株对子株氮营养的贡献占80.5%,子株根对自己的贡献仅为19.5%。
(3)碱化羊草草甸野外试验表明,切根对植物群落和土壤产生了明显的影响。植被密度、盖度和生物量呈先降低后升高的发展趋势,生物量在第二年达到最高,其中密度、盖度和生物量以切根深度10cm为最高。切根后土壤容重显著降低,其中0-10、10-20cm土层深度变化最明显。羊草作为根茎克隆植物,切断根茎后植株之间的克隆整合作用中止,导致羊草的适合度下降,生物量在第一年明显降低。随着时间的延长,切断根茎带来的消极作用慢慢减弱,羊草通过调节自身的可塑性反应适应新的环境,克隆分株加强了分蘖的速度而减少了对根茎空间扩展的资源投入,采取所谓“密集型”的发展策略。从切根改良羊草草甸的综合效果分析,一般切根深度控制在10-20cm较为适宜。
Clonal species are widely distributed in various ecosystems, as one of the unique group, it plays an important role in the vegetation restoration such as grassland, wetland and forest. During their long-term evolutionary processes, they may take the adaptive strategy-clonal integration to cope with the heterogeneous environments. Leymus chiensis (Trin.) Tzvel.(Poaceae) is a perennial rhizome clonal grass, and one of the main forage for vegetation recovery that often form monodominant stands in the L. chinensis meadow of the Songnen Plain. However, how alkalinity affects clonal integration of this species remained unknown. In this study, the clonal plant of L. chinensis was selected as our study materials, we had studied the clonal integration influences of severe rhizome and external environmental factors (soil alkalinities) and internal factors (developmental stage) on clonal plants, in the way of field investigation and simulation test, to determine the integration intensity, direction and benefits and costs under the combined effects of hetetogeneity alkalinities and plant age, to further analyze the mechanism of clonal integration on L. chinensis. Main conclusions are as follows:
(1) Heterogeneity alkalinities would significantly affect the effects of clonal integtation on plants. Within the heavy heterogeneous alkalinities treatments, clonal integration had a positive effect on plant growth, reproduction and tolerance to alkalinities. Support from mother ramets in non-stressful habitats improved the biomass of daughter ramets in stressful conditions through a connecting rhizome on L. chinensis by water transportation. But it do not occur in homogeneous habitats or those with low heterogeneity. The results indicated that higher rates of clonal integration may be selected for in habitats with increased alkaline heterogeneity. Clonal integration affected the allocation pattern of below-ground organs, with the roots, rhizomes and buds of daughter ramets showing different responses to clonal integration. Under the heterogeneous high alkaline conditions in this study, the biomass allocation to buds markedly increased as did rhizome length, but these changes were not significant and had negative effects on roots. Cost-benefit analysis based on biomass measures showed that the mother ramet incurred a cost to enhance daughter ramet survival but clonal integration confers overall benefits to the entire clonal fragment. Therefore, clonal integration is an important adaptive strategy by which rhizomatous clonal plants can withstand local alkaline stress.
(2) Our study has also found that the plant developmental stage is an important factor that affects the extent of clonal integration of L. chinensis, which has mother ramets that produce vegetative bulbils (clonal offspring) that pass through distinct pre-and post-rooting stages. The amount of15N translocated from mother to daughter ramets becoming increasing apparent in pre-rooting than post-rooting. In high alkaline environment, the nitrogen demands of the young ramets are supplied mainly from the mother ramets through clonal integration. The largest contribution portion of mother ramets applied was80.5%while that root of daughter ramets was19.5%.
(3) The results of field test showed that cutting roots has a significant influence on community characteristics and soil nature. The total community coverage and density and biomass levels decreased firstly and increased subsequently, and the10cm cutting root depth was highest. This decline of production mainly because of clonal integration loss interconnect ramets. While the time was prolonged, the influence to weaken slowly. Cutting root significantly increased the soil bulk density and water moisture content. The number of tiller was significantly increased and had little effects on rhizome length. On the basis of comprehensive control on alkalinization L. chinensis meadow in Songnen Plain, the cutting root depth in the10-20cm more appropriate.
(1)异质性碱胁迫明显影响植物的克隆整合作用效果。在重度碱胁迫处理条件下,克隆整合发挥正向作用,未受碱胁迫的母株和遭受碱胁迫的子株之间发生明显的水分传递现象,母株对子株的水分整合提高了子株的防御能力,缓解了碱胁迫对子株的生理伤害,从而相应地提高了羊草的生物量。但在同质和低碱异质性处理下,克隆整合的这种帮助作用不明显,说明克隆整合的强度随碱异质性的增大而增大。克隆植物会根据碱生境条件作出应对策略,产生形态和生理上的可塑性反应,在羊草地下器官生物量分配格局的结果表明,羊草根、根茎和芽对克隆整合表现出不同的可塑性响应,克隆整合作用使得芽的存活数明显增加,根茎的伸长不明显和克隆整合并未使植物根重有所提高,甚至出现下降的趋势。通过对母株和子株生物量的分析得出,克隆整合虽然使母株生物量有所降低,但子株从中获得的益处远远大于母株的损耗,因此对母株和子株的组合来说克隆整合是有益的。克隆整合是根茎克隆植物抵御碱胁迫的重要适应对策。
(2)同位素标记试验证明,克隆植物的母株通过连接的根茎向子株输送氮素,从而减缓了碱胁迫对子株的影响,克隆整合增加了子株的高度和地上地下生物量。在高碱胁迫下,子株的不同发育阶段对克隆整合的强度有一定的影响,母株向子株传输的氮营养在子株生根后比生根前有一定减少,但子株的营养供给主要依赖于母株。通过15N丰度的计算结果表明,母株对子株氮营养的贡献占80.5%,子株根对自己的贡献仅为19.5%。
(3)碱化羊草草甸野外试验表明,切根对植物群落和土壤产生了明显的影响。植被密度、盖度和生物量呈先降低后升高的发展趋势,生物量在第二年达到最高,其中密度、盖度和生物量以切根深度10cm为最高。切根后土壤容重显著降低,其中0-10、10-20cm土层深度变化最明显。羊草作为根茎克隆植物,切断根茎后植株之间的克隆整合作用中止,导致羊草的适合度下降,生物量在第一年明显降低。随着时间的延长,切断根茎带来的消极作用慢慢减弱,羊草通过调节自身的可塑性反应适应新的环境,克隆分株加强了分蘖的速度而减少了对根茎空间扩展的资源投入,采取所谓“密集型”的发展策略。从切根改良羊草草甸的综合效果分析,一般切根深度控制在10-20cm较为适宜。
Clonal species are widely distributed in various ecosystems, as one of the unique group, it plays an important role in the vegetation restoration such as grassland, wetland and forest. During their long-term evolutionary processes, they may take the adaptive strategy-clonal integration to cope with the heterogeneous environments. Leymus chiensis (Trin.) Tzvel.(Poaceae) is a perennial rhizome clonal grass, and one of the main forage for vegetation recovery that often form monodominant stands in the L. chinensis meadow of the Songnen Plain. However, how alkalinity affects clonal integration of this species remained unknown. In this study, the clonal plant of L. chinensis was selected as our study materials, we had studied the clonal integration influences of severe rhizome and external environmental factors (soil alkalinities) and internal factors (developmental stage) on clonal plants, in the way of field investigation and simulation test, to determine the integration intensity, direction and benefits and costs under the combined effects of hetetogeneity alkalinities and plant age, to further analyze the mechanism of clonal integration on L. chinensis. Main conclusions are as follows:
(1) Heterogeneity alkalinities would significantly affect the effects of clonal integtation on plants. Within the heavy heterogeneous alkalinities treatments, clonal integration had a positive effect on plant growth, reproduction and tolerance to alkalinities. Support from mother ramets in non-stressful habitats improved the biomass of daughter ramets in stressful conditions through a connecting rhizome on L. chinensis by water transportation. But it do not occur in homogeneous habitats or those with low heterogeneity. The results indicated that higher rates of clonal integration may be selected for in habitats with increased alkaline heterogeneity. Clonal integration affected the allocation pattern of below-ground organs, with the roots, rhizomes and buds of daughter ramets showing different responses to clonal integration. Under the heterogeneous high alkaline conditions in this study, the biomass allocation to buds markedly increased as did rhizome length, but these changes were not significant and had negative effects on roots. Cost-benefit analysis based on biomass measures showed that the mother ramet incurred a cost to enhance daughter ramet survival but clonal integration confers overall benefits to the entire clonal fragment. Therefore, clonal integration is an important adaptive strategy by which rhizomatous clonal plants can withstand local alkaline stress.
(2) Our study has also found that the plant developmental stage is an important factor that affects the extent of clonal integration of L. chinensis, which has mother ramets that produce vegetative bulbils (clonal offspring) that pass through distinct pre-and post-rooting stages. The amount of15N translocated from mother to daughter ramets becoming increasing apparent in pre-rooting than post-rooting. In high alkaline environment, the nitrogen demands of the young ramets are supplied mainly from the mother ramets through clonal integration. The largest contribution portion of mother ramets applied was80.5%while that root of daughter ramets was19.5%.
(3) The results of field test showed that cutting roots has a significant influence on community characteristics and soil nature. The total community coverage and density and biomass levels decreased firstly and increased subsequently, and the10cm cutting root depth was highest. This decline of production mainly because of clonal integration loss interconnect ramets. While the time was prolonged, the influence to weaken slowly. Cutting root significantly increased the soil bulk density and water moisture content. The number of tiller was significantly increased and had little effects on rhizome length. On the basis of comprehensive control on alkalinization L. chinensis meadow in Songnen Plain, the cutting root depth in the10-20cm more appropriate.
引文
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