喜旱莲子草克隆片段的适应策略
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摘要
干扰普遍存在于自然界,而由干扰引起的克隆植物片段化也在频繁地发生。由自然或人为因素导致的干扰能够使克隆植物分裂成由一个或者多个分株相连的克隆片段。克隆片段的传播、存活和生长是决定克隆植物扩散能力的重要因素。克隆片段的定居对水生和海洋植物(包括具有入侵性的湿地植物)的传播尤其重要。因此,研究克隆植物对克隆片段化的适应策略既有理论价值也有实际意义。我们以匍匐茎入侵克隆植物喜旱莲子草为研究对象,在克隆分株和克隆片段两个尺度上,检验片段化干扰和环境因子对克隆片段萌发和生长的影响。结果如下：
1.增加匍匐茎节间长度和保留叶片能够显著提高喜旱莲子草萌发率和生长(包括生物量、叶面积、分株数、匍匐茎长、叶片数)。所有喜旱莲子草的生长指标在片段分株与后端节间相连时比在片段分株与前端节间相连时表现得更好。
2.与其他种植方向相比较,克隆片段的水平放置将促进喜旱莲子草萌发率和生长能力。种植方向的影响在匍匐茎节间长的时候表现得更加显著。种植方向对植物生长的影响比匍匐茎节间长度的影响小；克隆片段的存活率在匍匐茎节间短的时候为60%,在匍匐茎节间长的时候为90%。
3.土壤掩埋深度的增加显著降低喜旱莲子草的萌发率,增加根冠比和匍匐茎长,但不会影响植物的生长。匍匐茎节间的增加显著增加了喜旱莲子草的存活率和生长,但是土壤掩埋深度与匍匐茎节间长度之间并没有交互作用。实验结果说明存储在匍匐茎节间的存储物质能够显著提高喜旱莲子草抗干扰的能力。
4.不管在母代环境还是在子代环境中,遮阴处理都将严重抑制喜旱莲子草的生长。母代效应只发生在克隆片段子代的对照环境中：而没有经历过遮阴处理的母代所产生的克隆片段具有更强生长能力。在克隆片段子代的环境中,去甲基化药剂“5-氮杂胞苷”只对经历过中度遮阴处理的母代所产生的克隆片段有促进作用。
5.不同程度的片段化和匍匐茎顶端的切除不会影响喜旱莲子草整个克隆片段的生长；但是克隆片段化越严重,新生分株的尺寸越小但数量越多。对于不同尺寸的克隆片段而言,由一个分株组成的克隆片段萌发率为85%,而由五个分株组成的克隆片段萌发率为100%。对于不同年龄的克隆片段而言,由年轻分株组成的克隆片段比由年老分株组成的克隆片段生长得更好。
6.水位上升对喜旱莲子草的生长指标具有显著的促进作用,说明外界资源供给可能随着水位的上升而提高。实验结果与概念模型的预测结果相同：匍匐茎切断对喜旱莲子草生长的影响会在水位为0cm的时候表现为正效应或无影响,而在水位为20cm或者40cm的时候表现为无影响或负效应。匍匐茎切断对克隆片段前端分株的抑制作用会随着水位上升而减小；匍匐茎切断对片段后端分株的促进作用也会随着水位上升而减小。
以上实验结果表明喜旱莲子草能够通过克隆存储、克隆整合以及克隆分株数量和分株质量之间的权衡等策略去适应克隆片段化。匍匐茎节间和叶片中的存储物质显著提高了喜旱莲子草克隆片段的萌发和生长能力,而由远端分株组成的克隆片段具有强大的传播和再生能力。对于整个克隆片段而言,克隆分株数量与分株质量之间的权衡则使喜旱莲子草在不同强度的干扰中均保持相同的适合度。随着外界资源水平的提高,克隆整合的促进作用也提高了喜旱莲子草在同质高资源环境中的竞争能力。另一方面,环境因子将影响喜旱莲子草对克隆片段化的适应策略。匍匐茎节间的存储物质对克隆片段的影响依赖于植物的不同种植方向,并且随着节间的增长而表现得更加明显。喜旱莲子草克隆片段能够通过改变根冠比和匍匐茎长去适应不同的土壤掩埋环境,并且使不同尺寸的克隆片段具有相似的适合度。而在光照胁迫中,喜旱莲子草克隆片段则通过母代效应影响喜旱莲子草子代克隆片段的适合度。因此,不管是人为干扰还是自然干扰,由干扰产生的克隆片段并不能影响喜旱莲子草的生长速率,却会增加其在干扰生境中的入侵风险。
Disturbance is common in nature and disturbance-caused fragmentation of clones happens frequently in clonal plants. Natural and human-caused disturbances can fragment the groups of connected ramets formed by clonal plants into smaller groups of one to several ramets. The capacity of these small fragments to disperse, survive, and grow is a major factor in the spread of clonal plants. Establishment of small fragments is particularly important in the spread of aquatic clonal species, including species of introduced, invasive plants in wetlands. Understanding what determines the survival and growth of small clonal fragments is thus of both scientific and practical interest. At both the single-ramet and whole fragment levels, we conducted a series of greenhouse experiments using the introduced, stoloniferous herb Alternanthera philoxeroides to test effects of fragmentation degree and environmental factors on survival and growth of fragments.
These results show that (1) increasing internode length and presence of leaves significantly increased the survival rate and growth (biomass, leaf area, number of ramets, stolon length and number of leaves) of the A. philoxeroides plants. All growth measures of A. philoxeroides at harvest were larger when the ramets were attached with a distal internode than when they were attached with a proximal internode, but the survival rate was lower.(2) Survival and growth of the A. philoxeroides plants were greatest when fragments were positioned horizontally. Contrary to expectations, some of these effects of orientation were stronger when attached stolons were longer. Orientation had smaller effects than stolon length on the performance of fragments; survival of fragments was about60%with shorter stolons and90%with longer stolons.(3) Increasing burial depth significantly reduced survival of the A. philoxeroides plants and increased root to shoot ratio and total stolon length, but did not change growth. Increasing internode length significantly increased survival and growth, but there was no interaction effect with burial depth on any traits measured.(4) Shading severely inhibited growth of the A. philoxeroides plants in both the maternal environment and daughter's environment. Maternal effect existed only in the daughter's control treatment, in which fragments that did not experience shading in the maternal environment grew better than those that suffered shading. Strikingly, in the daughter's control treatment, the demethylating agent5-azacytidine increased growth of fragments that experienced intermediate shading in the maternal environment.(5) Fragmentation and apex removal did not affect total growth of the A. philoxeroides plants, but more highly fragmented plants produced smaller but more ramets. The survival of fragments consisting of one original ramet was about85%, compared to100%survival of fragments with5original ramets. Fragments consisting of the youngest ramets along the original stolon grew more than fragments of the same size that consisted of older ramets.(6) Across connection treatments, water depth had a positive effect on most measures of growth, indicating that resource availability increased with water depth. Largely consistent with the conceptual model, restricting clonal integration by severing the apical and basal parts of an A. philoxeroides fragment had positive to neutral effects on total growth of fragments at0cm water depth, and neutral to negative effects at20and40cm. The effects of severance on the apical part were generally negative and greater at greater depth; effects of severance on the basal part were generally positive and smaller at greater depth.
These results indicate that A. philoxeroides can response to clonal fragmentation via clonal storage, clonal integration and a trade-off between ramet number and ramet mass. Storage in stolons and leaves can contribute greatly to the survival and growth of A. philoxeroides, and detachment of small groups of distal ramets along stolons may be very effective in spreading the species. The trade-off between the number and the size of new ramets produced by fragments may represent an adaptive, plastic response to disturbance. Clonal integration also helps A. philoxeroides fragments become advantageous in homogeneous habitats where resource availability is high. Furthermore, environmental factors also influence the responses of A. philoxeroides to clonal fragmentation. Orientation will affect establishment of small clonal fragments, and such effects can be stronger in larger rather than smaller fragments. Regeneration capacity of fragments will be reduced by sand burial, but A. philoxeroides can maintain the same fitness of the surviving plants by changing biomass allocation and stolon length. A. philoxeroides can also response to shading by the maternal effect. Therefore, These results suggest that disturbance-based fragmentation of clones cannot control the growth rate of A. philaxeroides but can increase the spread rate of this species in disturbed habitats.
1.增加匍匐茎节间长度和保留叶片能够显著提高喜旱莲子草萌发率和生长(包括生物量、叶面积、分株数、匍匐茎长、叶片数)。所有喜旱莲子草的生长指标在片段分株与后端节间相连时比在片段分株与前端节间相连时表现得更好。
2.与其他种植方向相比较,克隆片段的水平放置将促进喜旱莲子草萌发率和生长能力。种植方向的影响在匍匐茎节间长的时候表现得更加显著。种植方向对植物生长的影响比匍匐茎节间长度的影响小；克隆片段的存活率在匍匐茎节间短的时候为60%,在匍匐茎节间长的时候为90%。
3.土壤掩埋深度的增加显著降低喜旱莲子草的萌发率,增加根冠比和匍匐茎长,但不会影响植物的生长。匍匐茎节间的增加显著增加了喜旱莲子草的存活率和生长,但是土壤掩埋深度与匍匐茎节间长度之间并没有交互作用。实验结果说明存储在匍匐茎节间的存储物质能够显著提高喜旱莲子草抗干扰的能力。
4.不管在母代环境还是在子代环境中,遮阴处理都将严重抑制喜旱莲子草的生长。母代效应只发生在克隆片段子代的对照环境中：而没有经历过遮阴处理的母代所产生的克隆片段具有更强生长能力。在克隆片段子代的环境中,去甲基化药剂“5-氮杂胞苷”只对经历过中度遮阴处理的母代所产生的克隆片段有促进作用。
5.不同程度的片段化和匍匐茎顶端的切除不会影响喜旱莲子草整个克隆片段的生长；但是克隆片段化越严重,新生分株的尺寸越小但数量越多。对于不同尺寸的克隆片段而言,由一个分株组成的克隆片段萌发率为85%,而由五个分株组成的克隆片段萌发率为100%。对于不同年龄的克隆片段而言,由年轻分株组成的克隆片段比由年老分株组成的克隆片段生长得更好。
6.水位上升对喜旱莲子草的生长指标具有显著的促进作用,说明外界资源供给可能随着水位的上升而提高。实验结果与概念模型的预测结果相同：匍匐茎切断对喜旱莲子草生长的影响会在水位为0cm的时候表现为正效应或无影响,而在水位为20cm或者40cm的时候表现为无影响或负效应。匍匐茎切断对克隆片段前端分株的抑制作用会随着水位上升而减小；匍匐茎切断对片段后端分株的促进作用也会随着水位上升而减小。
以上实验结果表明喜旱莲子草能够通过克隆存储、克隆整合以及克隆分株数量和分株质量之间的权衡等策略去适应克隆片段化。匍匐茎节间和叶片中的存储物质显著提高了喜旱莲子草克隆片段的萌发和生长能力,而由远端分株组成的克隆片段具有强大的传播和再生能力。对于整个克隆片段而言,克隆分株数量与分株质量之间的权衡则使喜旱莲子草在不同强度的干扰中均保持相同的适合度。随着外界资源水平的提高,克隆整合的促进作用也提高了喜旱莲子草在同质高资源环境中的竞争能力。另一方面,环境因子将影响喜旱莲子草对克隆片段化的适应策略。匍匐茎节间的存储物质对克隆片段的影响依赖于植物的不同种植方向,并且随着节间的增长而表现得更加明显。喜旱莲子草克隆片段能够通过改变根冠比和匍匐茎长去适应不同的土壤掩埋环境,并且使不同尺寸的克隆片段具有相似的适合度。而在光照胁迫中,喜旱莲子草克隆片段则通过母代效应影响喜旱莲子草子代克隆片段的适合度。因此,不管是人为干扰还是自然干扰,由干扰产生的克隆片段并不能影响喜旱莲子草的生长速率,却会增加其在干扰生境中的入侵风险。
Disturbance is common in nature and disturbance-caused fragmentation of clones happens frequently in clonal plants. Natural and human-caused disturbances can fragment the groups of connected ramets formed by clonal plants into smaller groups of one to several ramets. The capacity of these small fragments to disperse, survive, and grow is a major factor in the spread of clonal plants. Establishment of small fragments is particularly important in the spread of aquatic clonal species, including species of introduced, invasive plants in wetlands. Understanding what determines the survival and growth of small clonal fragments is thus of both scientific and practical interest. At both the single-ramet and whole fragment levels, we conducted a series of greenhouse experiments using the introduced, stoloniferous herb Alternanthera philoxeroides to test effects of fragmentation degree and environmental factors on survival and growth of fragments.
These results show that (1) increasing internode length and presence of leaves significantly increased the survival rate and growth (biomass, leaf area, number of ramets, stolon length and number of leaves) of the A. philoxeroides plants. All growth measures of A. philoxeroides at harvest were larger when the ramets were attached with a distal internode than when they were attached with a proximal internode, but the survival rate was lower.(2) Survival and growth of the A. philoxeroides plants were greatest when fragments were positioned horizontally. Contrary to expectations, some of these effects of orientation were stronger when attached stolons were longer. Orientation had smaller effects than stolon length on the performance of fragments; survival of fragments was about60%with shorter stolons and90%with longer stolons.(3) Increasing burial depth significantly reduced survival of the A. philoxeroides plants and increased root to shoot ratio and total stolon length, but did not change growth. Increasing internode length significantly increased survival and growth, but there was no interaction effect with burial depth on any traits measured.(4) Shading severely inhibited growth of the A. philoxeroides plants in both the maternal environment and daughter's environment. Maternal effect existed only in the daughter's control treatment, in which fragments that did not experience shading in the maternal environment grew better than those that suffered shading. Strikingly, in the daughter's control treatment, the demethylating agent5-azacytidine increased growth of fragments that experienced intermediate shading in the maternal environment.(5) Fragmentation and apex removal did not affect total growth of the A. philoxeroides plants, but more highly fragmented plants produced smaller but more ramets. The survival of fragments consisting of one original ramet was about85%, compared to100%survival of fragments with5original ramets. Fragments consisting of the youngest ramets along the original stolon grew more than fragments of the same size that consisted of older ramets.(6) Across connection treatments, water depth had a positive effect on most measures of growth, indicating that resource availability increased with water depth. Largely consistent with the conceptual model, restricting clonal integration by severing the apical and basal parts of an A. philoxeroides fragment had positive to neutral effects on total growth of fragments at0cm water depth, and neutral to negative effects at20and40cm. The effects of severance on the apical part were generally negative and greater at greater depth; effects of severance on the basal part were generally positive and smaller at greater depth.
These results indicate that A. philoxeroides can response to clonal fragmentation via clonal storage, clonal integration and a trade-off between ramet number and ramet mass. Storage in stolons and leaves can contribute greatly to the survival and growth of A. philoxeroides, and detachment of small groups of distal ramets along stolons may be very effective in spreading the species. The trade-off between the number and the size of new ramets produced by fragments may represent an adaptive, plastic response to disturbance. Clonal integration also helps A. philoxeroides fragments become advantageous in homogeneous habitats where resource availability is high. Furthermore, environmental factors also influence the responses of A. philoxeroides to clonal fragmentation. Orientation will affect establishment of small clonal fragments, and such effects can be stronger in larger rather than smaller fragments. Regeneration capacity of fragments will be reduced by sand burial, but A. philoxeroides can maintain the same fitness of the surviving plants by changing biomass allocation and stolon length. A. philoxeroides can also response to shading by the maternal effect. Therefore, These results suggest that disturbance-based fragmentation of clones cannot control the growth rate of A. philaxeroides but can increase the spread rate of this species in disturbed habitats.
引文
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