不同生境下空心莲子草响应模拟昆虫采食的生长和化学防御策略

外来植物往往可以入侵多种生境并受到多种昆虫的采食,而不同生境条件将可能会影响这些入侵植物对昆虫采食的防御策略。以入侵我国的克隆植物——空心莲子草为研究对象,分别选择生长在水生生境、水陆两栖生境和陆生生境中的无性个体(分株),通过50%去叶处理模拟昆虫采食,分析不同生境下空心莲子草对模拟昆虫采食处理的生长及化学防御响应的差异。模拟昆虫采食处理显著抑制了陆生生境、水陆两栖生境以及水生生境下空心莲子草的根、茎、叶和总生物量,但对3种生境下空心莲子草的生物量分配(根冠比、根生物量分配、茎生物量分配和叶生物量分配)均无显著影响。陆生生境下空心莲子草根、茎和总生物量显著高于水陆两栖生境和水生生境,根冠比显著低于水陆两栖生境和水生生境。模拟昆虫采食处理显著降低了空心莲子草的木质素含量,而对单宁和总酚含量影响不显著。生境对木质素含量无显著影响,但陆生生境下空心莲子草单宁含量显著高于水陆两栖生境和水生生境,且总酚含量显著高于水陆两栖生境,表明陆生生境中空心莲子草具有更强的防御能力。空心莲子草木质素含量与总生物量无显著相关性,但在模拟采食情况下,其总酚含量与总生物量呈显著负相关,而无论模拟昆虫采食处理存在与否,空心莲子草单宁含量与总生物量均呈显著正相关。因此,空心莲子草存在昆虫介导的生长和化学防御之间的权衡,在昆虫采食的情况下可通过减少生长来增加对化学防御物质的投入,但生境对空心莲子草这种生长-防御权衡的影响十分有限。     

Effects of simulated herbivory and resource availability on the invasive plant, Alternanthera philoxeroides in different habitats

In biological control programs, the insect natural enemy’s ability to suppress the plant invader may be affected by abiotic factors, such as resource availability, that can influence plant growth and reproduction. Understanding plant tolerance to herbivory under different environmental conditions will help to improve biocontrol efficacy. The invasive alligator weed (Alternanthera philoxeroides) has been successfully controlled by natural enemies in many aquatic habitats but not in terrestrial environments worldwide. This study examined the effects of different levels of simulated leaf herbivory on the growth of alligator weed at two levels of fertilization and three levels of soil moisture (aquatic, semi-aquatic, and terrestrial habitats). Increasing levels of simulated (manual) defoliation generally caused decreases in total biomass in all habitats. However, the plant appeared to respond differently to high levels of herbivory in the three habitats. Terrestrial plants showed the highest below–above ground mass ratio (R/S), indicating the plant is more tolerant to herbivory in terrestrial habitats than in aquatic habitats. The unfertilized treatment exhibited greater tolerance than the fertilized treatment in the terrestrial habitat at the first stage of this experiment (day 15), but fertilizer appears not to have influenced tolerance at the middle and last stages of the experiment. No such difference was found in semi-aquatic and aquatic habitats. These findings suggest that plant tolerance is affected by habitats and soil nutrients and this relationship could influence the biological control outcome. Plant compensatory response to herbivory under different environmental conditions should, therefore, be carefully considered when planning to use biological control in management programs against invasive plants.     

Flooding compromises compensatory capacity of an invasive plant: implications for biological control

Plant compensatory growth is proposed to be insidious to biological control and known to vary under different environmental conditions. However, the effects of microsite conditions on compensation capacity and its indirect impacts on biological control of plant invaders have received little attention. Alligator weed, Alternanthera phioxeroides, is an invasive plant worldwide, growing in both aquatic and terrestrial habitats that are often affected by flooding. Biological control insects have been successful in suppressing the plant in many aquatic habitats but have failed in terrestrial habitats. To evaluate the impact of flooding on compensation capacity, we conducted common garden and greenhouse experiments in which plants were grown under different moisture conditions (aquatic versus terrestrial). Our results show that plants were able to fully recover from continued herbivory in the terrestrial habitat, but failed in the aquatic habitat, indicating a flooding-regulated plant compensatory capacity. Also, the grazed plants increased below-ground growth and reproductive root bud formation in the terrestrial habitat, but there was no such difference in the aquatic habitat. Our findings suggest that the differing plant compensatory capacity, affected by flooding, may explain the different biological control efficacy of alligator weed in aquatic and terrestrial habitats. Understanding mechanisms in plant invader compensation in different microsite conditions is important for improving management efficiency.     

喜旱莲子草对喀斯特三种不同生境的可塑性反应

喜旱莲子草自入侵以来在我国各地广为传播蔓延,对人们生产生活产生了重要影响。该研究通过野外采样和室内理化指标分析,探讨喀斯特不同生境中喜旱莲子草的可塑性。结果表明:(1)在三种生境中喜旱莲子草的节间长度、叶面积、单株重和叶干重差异显著。(2)在岩石环境中其无性繁殖器官——茎的投入比例占其生物量的比例最大,其次是行道生境和湘江河道。(3)节间距随环境因子从岩石环境、绿化行道和水生生境的变化,依次增加,叶质比(叶面积/质量)也随水分环境的增加而增加;(4)在岩石环境中,喜旱莲子草通过缩短节间距并增加无性繁殖器官——茎的质量来增加其在水分匮乏生境的繁殖能力。该研究结果为喜旱莲子草今后的防治对策提供了科学依据。     

水生空心莲子草入侵群落数量分类及其物种分布的环境解释

生境异质性在不同生态层次上影响物种分布与扩散,进而可能影响外来植物的入侵态势。该研究通过在中国21°N～31°N范围内设置23个10m×10m的样地,利用双向聚类(TWINSPAN)和除趋势对应分析(DCA)等数量生态学方法,研究水生型空心莲子草(Alternanthera philoxeroides)入侵群落的结构特征并划分其群丛类型,并采用冗余分析法(RDA)分析空心莲子草种群特征、群落物种分布与环境因子(经度、纬度、海拔、氨态氮、硝态氮、电导率、溶解氧)的关系,以期为生物入侵防治及本土生物多样性保护提供参考。结果显示:(1)23个样地共记录植物48种,隶属于21科42属,空心莲子草优势地位显著,主要伴生种为双穗雀稗(Paspalum paspaloides)、李氏禾(Leersia hexandra)和水蓼(Polygonum hydropiper)。(2)23个样地群落可划分为6组群丛类型,其中空心莲子草+双穗雀稗+水芹(Oenanthe javanica)是最主要的类型。(3)48种植物可划分为4类功能群,分别为先锋种型、优势伴生种型、次要伴生种型和逃避型,功能群的DCA分异性高于群丛类型。(4)群落物种分布状况主要由氨态氮和电导率决定,而物种丰富度及空心莲子草种群特征则受到电导率、经度、海拔和溶解氧等多个环境变量的影响;水生空心莲子草的入侵重要值取决于其无性分枝数和种群盖度。研究表明,水生空心莲子草入侵群落的结构相对简单,应加大对其入侵监测力度并维持较高的生境异质性,以延缓水域生态系统的生物入侵进程。     

The interspecific competition between Humulus scandens and Alternanthera philoxeroides

Bioinvasion has become a serious environmental problem in the world in general and is considered the second biggest threat to biodiversity. Alternanthera philoxeroides is widely distributed and causes the most serious threat to biodiversity in China. The traditional physical or biological control methods are not effective in controlling the invasion and extension of A. philoxeroides. In the present paper, some physiological characteristics of Humulus scandens and A. philoxeroides were investigated in the field and laboratory. The results showed that H. scandens is more competitive than A. philoxeroides, the competitive rate (CR) of H. scandens against A. philoxeroides was 9.834. Additionally, the leaf, stem, and root biomass of A. philoxeroides decreased significantly when the two species co-occurred. Thus, the invasive abilities of these two invasive plants are different and H. scandens strongly inhibited the growth of A. philoxeroides. Moreover, as an annual herb, H. scandens can be easily eliminated by harvesting before its seeds mature. The result suggests that sowing seeds of H. scandens in the habitats invaded by A. philoxeroides could be an ideal biological control method.     

Phenotypic variation and water selection potential in the stem structure of invasive alligator weed

The morphological and anatomical characteristics of stems have been found to be related to drought resistance in plants. Testing the phenotypic selection of water availability on stem anatomical traits would be useful for exploring the evolutionary potential of the stem in response to water availability. To test the phenotypic variation of the stem anatomical traits of an invasive plant in response to water availability, we collected a total of 320 individuals of Alternanthera philoxeroides from 16 populations from terrestrial and aquatic habitats in 8 plots in China and then analyzed the variation, differentiation, plasticity and selection potential of water availability on the stem anatomical traits. We found that except for the thickness of the cortex, all of the examined phenotypic parameters of the A. philoxeroides stem were significantly and positively correlated with soil water availability. The phenotypic differentiation coefficient for all of the anatomical structural parameters indicated that most of the variation existed between habitats within the same plot, whereas there was little variation among plots or among individuals within the same habitat except for variation in the thickness of the cortex. A significant phenotypic plasticity response to water availability was found for all of the anatomical traits of A. philoxeroides stem except for the thickness of the cortex. The associations between fitness and some of the anatomical traits, such as the stem diameter, the cortex area-to-stem area ratio, the pith cavity area-to-stem area ratio and the density of vascular bundles, differed with heterogeneous water availability. In both the aquatic and terrestrial habitats, no significant directional selection gradient was found for the stem diameter, the cortex area-to-stem area ratio or the density of vascular bundles. These results indicated that the anatomical structure of the A. philoxeroides stem may play an important role in the adaptation to changes in water availability.     

水淹胁迫诱导喜旱莲子草茎发生类环割效应

在水淹环境中,喜旱莲子草水面上未受淹的茎段常表现出增粗膨大的现象。遭受水淹的植物被淹没的组织和器官会面临氧气缺乏和能量供应不足的问题。植物体内碳水化合物的运输需要消耗能量,当因水淹而使植株被部分淹没（即地下部分全部和地上部分的一部分被淹没）时,由于水淹缺氧导致能量供应不足,碳水化合物在植物被淹组织内的运输可能受限从而在水面上的未淹茎段中积累并对未淹茎段的径向形态产生影响。为探究水淹环境中喜旱莲子草未受淹茎段增粗膨大是否与碳水化合物积累有关,对茎被水淹和茎不受水淹的喜旱莲子草进行对比研究,结果发现：（1）水淹的喜旱莲子草位于水面上的未淹茎段节间平均直径显著大于水面下受淹茎段节间平均直径,未淹茎段与受淹茎段相比发生显著的膨大现象;未水淹的喜旱莲子草其茎的上部茎段节间平均直径与下部茎段节间平均直径相比并无显著差异,上部茎段也无明显膨大现象。（2）水淹的喜旱莲子草位于水面上的未淹茎段碳水化合物含量显著高于未水淹的喜旱莲子草对应茎段的碳水化合物含量。本研究表明,水淹胁迫下喜旱莲子草位于水面上的未受淹茎段中碳水化合物发生积累,导致植株位于水面上的未受淹茎段发生与物理环割后类似的茎膨大现象。     

Response of an invasive liana to simulated herbivory: implications for its biological control

Pre-release evaluation of the efficacy of biological control agents is often not possible in the case of many invasive species targeted for biocontrol. In such circumstances simulating herbivory could yield significant insights into plant response to damage, thereby improving the efficiency of agent prioritisation, increasing the chances of regulating the performance of invasive plants through herbivory and minimising potential risks posed by release of multiple herbivores. We adopted this approach to understand the weaknesses herbivores could exploit, to manage the invasive liana, Macfadyena unguis-cati. We simulated herbivory by damaging the leaves, stem, root and tuber of the plant, in isolation and in combination. We also applied these treatments at multiple frequencies. Plant response in terms of biomass allocation showed that at least two severe defoliation treatments were required to diminish this liana's climbing habit and reduce its allocation to belowground tuber reserves. Belowground damage appears to have negligible effect on the plant's biomass production and tuber damage appears to trigger a compensatory response. Plant response to combinations of different types of damage did not differ significantly to that from leaf damage. This suggests that specialist herbivores in the leaf-feeding guild capable of removing over 50% of the leaf tissue may be desirable in the biological control of this invasive species.     

The invasive plant Alternanthera philoxeroides benefits from clonal integration in response to defoliation

Many notorious alien invasive plants have the capacity for vigorous clonal growth, and clonal integration may contribute to their invasiveness in response to various disturbances. Here, it is hypothesized that clonal integration affects the growth, biomass allocation, physiology, and compensatory response of the alien invasive clonal plant Alternanthera philoxeroides when faced with defoliation. To test these hypotheses, a growth experiment was conducted to investigate the effect of clonal integration on the responses of A. philoxeroides to different levels of defoliation. Daughter ramets that had been grown with stolon connections that were either severed from or connected to the mother plant were subjected to four defoliation levels: 0 (control), 30% (mild), 60% (moderate) and 90% (heavy) removal of leaf tissue. Defoliation greatly decreased growth (total biomass, number of ramets and total stolon length) but increased the maximum quantum yield of photosystem II (F_v/F_m) of daughter ramets. Clonal integration significantly increased growth, F_v/F_m and contents of non-structural carbohydrates (soluble sugars and total non-structural carbohydrates) of A. philoxeroides, and these effects were larger under heavier defoliation. Moreover, clonal integration markedly reduced the shoot/root ratio of A. philoxeroides, and these effects tended to increase with increasing levels of defoliation. These results support our hypothesis that A. philoxeroides benefits from clonal integration in response to defoliation, suggesting that clonal integration may be closely related to the invasiveness of A. philoxeroides in natural habitats with frequent disturbances.     

Impact of Galerucella birmanica (Coleoptera: Chrysomelidae) on growth and seed production of Trapa natans

Success in biological weed control programs depends upon the ability of host-specific herbivores to suppress populations of their host plant. While pre-release predictions of field host range (i.e., specificity) appear widely accurate, predictions about which agent or agent combination may suppress plant populations have lately been compared to predictions in a lottery. The history of weed biocontrol does not offer immediately obvious approaches to improve the lottery model, however, pre-release assessments of the impact of different herbivore densities on the invasive plant may provide an opportunity to improve predictions of success. In this paper, we report on the impact of the leaf beetle Galerucella birmanica on growth and reproduction of water chestnut, Trapa natans, in the native range in China. At low herbivore densities (10–50 larvae/rosette), plants compensated for leaf herbivory by increasing leaf production at the expense of reproductive effort. Inoculating >50 first instar larvae per rosette greatly suppressed biomass production and plants were unable to grow when three or more G. birmanica pairs were released per seven rosettes. In the native range, similar densities are found in the field, resulting in complete defoliation of T. natans. Our study indicates that G. birmanica feeding has significant negative impacts on T. natans. This chrysomelid species appears to be a promising biological control agent and we would predict that the species will be able to attain sufficiently high populations to control its host plant—if approved for release in North America.     

Effect of simulated herbivory on growth of the invasive weed Hygrophila polysperma: Experimental and predictive approaches

Herbivory simulation studies, through mechanical removal of leaf tissue, provide valuable insight about plant compensation and tolerance to defoliation. A mesocosm experiment was conducted to examine the effects of defoliation on growth and biomass accumulation of Hygrophila polysperma and thereby determine the critical level of herbivory necessary to achieve significant reduction in growth of this invasive plant. The data collected during the experiment were used to develop an empirical plant growth model to examine the usefulness of a model-based approach for a priori understanding of plant response to defoliation. The results of the mesocosm experiment showed that defoliation significantly influenced growth and biomass accumulation of hygrophila. The empirical plant growth model accurately simulated plant growth response to herbivory across treatments. Based on the results of the mesocosm experiment, an insect defoliator that causes complete defoliation of hygrophila at least at monthly intervals may be able to reduce biomass and growth of hygrophila. The ability of the mathematical model to predict the effects of defoliation on hygrophila suggest that it could be a useful tool for the selection of effective biological control agents.     

入侵植物水花生解剖学和组织化学染色研究

水花生(Alternanthera philoxeroides)因其表型可塑性、高生长速率和快速无性繁殖能适应水、陆生境。该文利用光学显微镜和荧光显微镜对水、陆生境的水花生不定根、茎解剖结构、组织化学特征及质外体通透性进行了研究。结果表明:(1)水生境下,其不定根皮层中具较大裂生型通气组织,无次生生长,内皮层具凯氏带且栓质化,皮层和皮下层明显木质化。(2)在陆生环境下,其不定根有次生生长,胞间具通气组织,内皮层具凯氏带且栓质化,皮层和皮下层略木质化;此外,不定根还具额外形成层,产生次生维管束、薄壁组织和不定芽;多年生不定根中具直接分裂的薄壁组织,周皮具凯氏带,且栓质化和木质化。(3)水、陆生境下,其匍匐茎具髓和中空髓腔,发生次生生长,具裂-溶生型通气组织、单层内皮层、厚角组织和木质化且栓质化的角质层,陆生匍匐茎周皮栓质化且木质化。(4)水花生质外体屏障结构组成复杂,黄连素无法穿透质外体屏障结构。水花生的上述解剖学特征,是水花生适应水、陆生境的有力证据。     

空心莲子草叶甲对越冬保护的响应与控害效能

空心莲子草叶甲Agasicles hygrophila是入侵杂草空心莲子草Alternan thera philoxeroide的重要天敌,冬春季低温冰冻是影响其安全越冬的关键胁迫因子。为了探明采取保护措施是否能提高越冬后的种群基数问题,在低温冰冻天气来临前,用塑料膜覆盖网室构建空心莲子草与空心莲子草叶甲保护生境(PH),分低温期(2009年1月21日至4月9日)、升温期(4月15日至5月1日)与自然生境(NH)中叶甲成虫初现期(从6月1日至29日)3个阶段,调查比较PH与NH两类生境中空心莲子草叶甲种群数量、结构、空间生态位分布特征及控害效能。结果表明:在保护生境中,低温期空心莲子草叶甲不休眠,且能安全越冬,当晴天温度较高时,成虫主要在直立茎或嫩枝上活动取食、补充营养,而在阴雨低温天气则停留在匍匐草层或土壤表层、缝隙中,少活动。随着气温回升,成虫转移到直立茎生态位上活动取食,并于4月上旬开始产卵繁殖,种群数量迅速增加,对空心莲子草的地上部分有较好的控制效果。在自然生境中,直到6月初才见少量成虫活动,此时草生长茂密,较低的空心莲子草叶甲种群难以发挥生防因子的作用。比较两种生境中6月1日与6月15日的控草效果显示,空心莲子草株高增长率分别为:PH-13.91%,NH-2.94%,NH中空心莲子草株高极显著高于PH,且单株生物量的控制率为PH(47.56%)极显著高于NH(4.89%)。在6月15日与29日的调查表明,单株叶片控制效果PH极显著优于NH,PH中的空心莲子草茎上均无叶,叶片或被取食殆尽、或被害后干枯脱落。茎被害率、茎蛀孔数与蛀茎率均为PH极显著高于NH。结果说明覆膜保护能显著提高叶甲越冬种群基数,使其提前进入繁殖期,从而能更有效地控制生境中的空心莲子草。     

The Structural Adaptation of Aerial Parts of Invasive <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> to Water Regime

Alternanthera philoxeroides has successfully invaded diverse habitats with considerably various water availability, threatening biological diversity in many parts of the world. Because its genetic variation is very low, phenotypic plasticity is believed to be the primary strategy for adapting to the diverse habitats. In the present paper, we investigated the plastic changes of anatomical traits of the aerial parts of A. philoxeroides from flooding to wet then to drought habitat; the results are as follows: A. philoxeroides could change anatomical structures sensitively to adapt to water regime. As a whole, effects of water regime on structures in stem were greater than those in leaf. Except for principal vein diameter and stoma density on leaf surfaces, all other structural traits were significantly affected by water regime. Among which, cuticular wax layer, collenchyma cell wall, phloem fiber cell wall, and hair density on both leaf surfaces thickened significantly with decrease of water availability, whereas, pith cavity and vessel lumen in stem lessened significantly; wet habitat is vital for the spread of A. philoxeroides from flooding to drought habitat and vice versa, because in this habitat, it had the greatest structural variations; when switching from flooding to wet then to drought habitat, the variations of cuticular wax layer, collenchyma cell wall, phloem fiber cell wall, pith cavity area ratio, diameter of vessel lumen, and hair density on both leaf surfaces, played the most important role. These responsive variables contribute most to the adaptation of A. philoxeroides to diverse habitats with considerably various water availability.     

Mortality of the terrestrial adult and aquatic nymphal life stages of Baetis vernus and Baetis rhodani in the Breitenbach, Germany (Insecta: Ephemeroptera)

1. The mortality of Baetis vernus Curtis and Baetis rhodani Pictet during the terrestrial-aerial and aquatic life stages was studied at the Breitenbach near Schlitz, Hesse, Germany. The number of females emerging from the stream was recorded with emergence traps. To estimate mortality of females of both species during terrestrial life, numbers of emerging females were compared with numbers of females returning to the stream for oviposition, as shown by numbers of egg masses found in the stream. 2. Mortality of female B. vernus during their terrestrial life stage was 98.8%. It was 91.2 and 96.6%, respectively, during the first and second generations of B. rhodani. 3. To estimate the mortality of both sexes of B. rhodani during the aquatic life stage, the number of eggs laid by the first generation was compared with the number of adults emerging during the second generation. 4. Mortality during the aquatic phase (both sexes combined) of B. rhodani was 91.1%. 5. Mortality during the terrestrial life stages was at least as high, if not higher than during the aquatic stages. Evidently, there is a considerable export of organic material into the terrestrial environment around the stream. Mortality during terrestrial life may be an important regulator of population size.     

Effects of a nematode Meloidogyne incognita and its interaction with above‐ground herbivory on an invasive wetland plant,alligator weed (Alternanthera philoxeroides)

Invasive plants may be attacked both above ground and below ground. Few studies have, however, investigated the simultaneous effects of above‐ground and below‐ground herbivory. In the present study, we report the effects of beetle herbivory and nematode infection on alligator weed, Alternanthera philoxeroides, an invasive plant in China. We found that the root‐knot nematode Meloidogyne incognita widely occurred on the plant in south China. To examine its effect on the plant in conjunction with above‐ground herbivory, we conducted a field common garden experiment with a local insect defoliator, Cassida piperata. We also included the native congener Alternanthera sessilis in our experiments for a comparison of the response of invasive and native species. We found no significant effects on plant biomass of the nematode infection in conjunction with the above‐ground herbivory. Further chemical analysis, however, showed that the water‐soluble carbohydrate content in roots of A. philoxeroides was significantly increased in plants attacked by both the nematode and the herbivore compared with the water‐soluble carbohydrate content in plants attacked by only the nematode or herbivore alone. We found no such change in the native congener A. sessilis. Together these results may suggest that A. philoxeroides tolerates joint above‐ground and below‐ground damage by allocating more resources to below‐ground material.     

水体溶氧影响陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)对完全水淹的耐受力

溶氧是水环境中一个重要的环境因子,为了探讨水中的溶氧含量水平是否会对陆生植物的耐淹能力造成影响,研究了陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)在遭受不同溶氧含量水体完全淹没后的生长表现、存活情况和非结构碳水化合物的变化。实验结果表明:(1)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的存活。受高溶氧水体完全水淹的喜旱莲子草和牛鞭草主茎的完好程度和存活叶的数量均显著高于遭受低溶氧水体完全水淹的喜旱莲子草和牛鞭草,喜旱莲子草和牛鞭草在高溶氧水体完全水淹后的生物量比低溶氧水体完全水淹后要高;(2)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的生长,受高溶氧水体完全水淹的喜旱莲子草主茎伸长生长和不定根生长显著强于受低溶氧水体完全水淹的喜旱莲子草,在不定根的生长上牛鞭草也具有同样的表现。(3)高溶氧水环境有利于减小被完全淹没的喜旱莲子草和牛鞭草的碳水化合物消耗,两种植物在受高溶氧完全水淹后体内具有的非结构性碳水化合物含量均比受低溶氧完全水淹后高。(4)喜旱莲子草比牛鞭草能更好地耐受完全水淹,当处于低溶氧完全水淹时表现得更为明显,本研究表明入侵物种喜旱莲子草比本地物种牛鞭草具有更强的环境适应能力和水淹耐受能力。     

空心莲子草响应南方菟丝子寄生的生长-防御权衡

为探讨全寄生植物南方菟丝子(Cuscuta australis)防治入侵植物空心莲子草(Alternanthera philoxeroides)的可行性,以二者野外天然生长的种群为研究对象,分析南方菟丝子寄生对空心莲子草生长及防御的影响,阐明空心莲子草在受到寄生胁迫时如何权衡自身生长与防御的关系,进而发展出一套应对南方菟丝子寄生的生长-防御策略。结果显示:(1)南方菟丝子寄生显著改变空心莲子草茎的形态,茎直径和平均节间长均增加,茎直径变化极显著(P0.01);(2)南方菟丝子寄生显著减少空心莲子草叶片数,但同时显著增加后者茎的分枝数,而茎上的节是潜在的无性繁殖体,故有利于空心莲子草的克隆繁殖;此外,南方菟丝子寄生显著降低了空心莲子草的根、茎、叶生物量和总生物量,抑制空心莲子草的生长;(3)南方菟丝子寄生显著增加空心莲子草茎的单宁、总酚、三萜皂苷含量,增强其防御能力;(4)南方菟丝子寄生的空心莲子草的生物量与茎部木质素、三萜皂苷、单宁和总酚含量均呈现显著负相关性(P0.01),对照组则不存在相关性;且寄生组较对照组相比,生物量的相对百分比显著低于对照组(P0.01),而用于防御的次生代谢产物总含量的相对百分比显著高于对照组(P0.01)。以上结果表明,受到南方菟丝子寄生胁迫后,空心莲子草改变自身的生长-防御策略,减少营养生长投入而将更多的资源投向克隆繁殖,同时增强对"防御"物质的投入,增强其防御能力,以利于后代生存和繁衍。     

Nitrogen-15 partitioning within a three generation tiller sequence of the bunchgrass Schizachyrium scoparium: response to selective defoliation

Summary Nitrogen partitioning among three generations of tillers within the bunchgrass Schizachyrium scoparium var. frequens was investigated in a controlled environment as a potential mechanism of herbivory tolerance. Nitrogen-15 was transported from the labelled primary tiller generation to both shoots and roots of nondefoliated secondary and tertiary tiller generations within 24 h. Partial defoliation increased shoot nitrogen concentration of secondary and tertiary generation tillers by 110 and 120%, respectively, 24 h following defoliation. Shoot nitrogen concentration was preferentially increased by partial defoliation of tertiary generation tillers throughout the 120 h experimental period, but diminished to concentrations comparable to nondefoliated tillers within shoots of the secondary generation at 72 h. In contrast to nitrogen concentration, the total amount of nitrogen imported by secondary and tertiary generation tillers decreased 62 and 73%, respectively, 24 h following partial defoliation and did not attain values comparable to respective nondefoliated tillers. Consequently, preferential nitrogen concentration occurred in response to partial tiller defoliation without an increase in total nitrogen import based on the reduction in the total nitrogen requirement per tiller generation associated with defoliation. Estimates of both the total amount of nitrogen import and nitrogen concentration are necessary to accurately interpret the dynamics of intertiller nitrogen allocation.