外来种喜旱莲子草的入侵生态学
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摘要
从南美引入我国的喜旱莲子草是一种水陆两栖的外来入侵植物。该种分布广,常以中空的茎在水面形成漂浮植毡层。喜旱莲子草快速的生长和扩散能力已对生态系统和社会经济发展造成严重影响。然而,目前对于喜旱莲子草的入侵机制研究较少。本文针对喜旱莲子草的无性繁殖、生态适应和其对乡土水生植物的影响进行了研究,以分析喜旱莲子草入侵机理。
     陆生型喜旱莲子草的地下根茎系统非常庞大,繁殖能力强。肉质根数随着土壤深度的增加而减少,0—10cm土层中根数最多;10—20cm深度的肉质根最粗;生物量集中分布在0—30cm深的土层中。肉质根上的不定芽数以0—10cm的最多(1260个)。喜旱莲子草地下茎主要存在于0-20cm深的土层中,地下茎上的腋芽数量在0-10cm土层最多。肉质根萌生幼芽的能力与其直径和生物量成正相关。肉质根浸水后产生了大量的不定芽。绝大部分不定芽在土壤中可以分枝。
     喜旱莲子草漂浮植毡层搁浅在陆地上越冬后,植毡层上层的匍匐茎失水干枯,而下层的匍匐茎能正常越冬。越冬植毡层中的匍匐茎根据含水量分为三类:干枯茎、萎蔫茎和鲜茎。所有的茎浸水4周以后,比较各类型茎的繁殖能力。结果表明:干枯茎已失去繁殖能力,鲜茎繁殖能力最大。喜旱莲子草属于先萌芽后生根型植物,这种机制有利于喜旱莲子草在水体中扩散。
     光照和牧食是影响外来种入侵的两个重要因子。在对喜旱莲子草进行遮荫和刈割处理的试验中发现:遮荫对未刈割植株的叶数和叶生物量没有影响,但明显减少了刈割植株的叶数和叶生物量。遮荫增加了所有植株的叶面积,并使更多的生物量分配到叶上,抑制了肉质根的形成,减少了茎生物量、根茎比(R/S)、相对生长速率(RGR)和肉质根生物量比(TRMR),但对茎生物量比(SMR)和细根生物量比(FRMR)没有明显影响。无论是在遮荫下还是在全日照下,刈割除对茎生物量有明显影响外,对根粗、最大根长、细根重、SLA、LAR、R/S和生物量分配都没有影响。另外刈割使生长于遮荫环境下的植株RGR、节数、茎长度和叶生物量降低。
     许多外来入侵植物的光合能力强,水分利用效率(WUE)和光利用效率(LUE)比乡土植物高。对处于尘长早期、盛花期(或生长中期)和花果期的水尘型与陆生型喜旱莲子草及乡土水生植物李氏禾、菰和芦苇的光合特性进行了测定。结果
Alternanthera philoxeroides (Mart.) Griseb. is an exotic aquatic-terrestrial plant that was introduced into China from South America. This species distributes widely and often extends hollow stems over the surface of the water, forming interwoven, floating mats. The capacity of growing and spreading rapidly of this species have caused many disadvantage to ecosystem and development of society and economy. However, the mechanisms of its invasion are not clear. This paper studied asexual reproduction, ecological adaptation and the effects on native aquatic plants of A. philoxeroides in order to analyze its invasion mechanisms.The system of root and underground stems of terrestrial A. philoxeroides is huge, and the reproduction capacity of which is high. Thick root numbers of A. philoxeroides were decreased with increasing soil depth. The maximal number and diameter of thick root were found in 0-10cm and 10-20cm depths respectively. Most of thick root biomass was found in 0-30cm depths. The maximal number of adventitious bud (1260 buds) was found in 0-10cm depths. Underground stems mainly distributed in 0-20cm depths. However, The axillary buds generated from underground stem were mainly distributed in 0-10cm depths. The positive relation between the capacity of generating bud and the thick root diameter as well as the thick root biomass was found. A large number of adventitious bud generated when thick root sinked into water. Most of adventitious buds branched before they emerged soil.After the floating-mat of A. philoxeroides stayed on the land and experienced the winter, the prostrate stems could be separate into 3 types according to water content: air-dry stems, wither stems and fresh stems. All the stems were sinked into water for 4 weeks in order to study the reproduction capacity of stems. The results showed that: Air-dry stems had no capacity of reproduction, but fresh stems had the maximal capacity. The mechanisms of generating bud earlier than that of root would help A. philoxeroides disperse in the water.Light and grazing are the two important factors that influence the invasion of exotic plants. The work of the effect of shading and mowing on A. philoxeroides showed that: shading had no effect on leaf number and leaf biomass of unmown plants, but on mown plants. SLA, LAR and leaf biomass was increased by shading treatments.
    Shading led to lower stem biomass. thick root biomass, R/S. RGR and TRMR. but had no effects on SMR and FRMR. Mowing had no effects on thick root thickness, maximal root length, fine root biomass, SLA. LAR, R/S and biomass allocation, but on stem biomass of plants that both grew in shading and full sunlight habitats. In addition, mowing led to lower RGR, internodes number, shoot length and leaf biomass of plants grew in shading habitats.Photosynthetic capacity, water use efficiency (WUE) and light use efficiency (LUE) of many invasive exotic plants were greater than native plants. Photosynthetic parameters of aquatic A. philoxeroides, terrestrial A. philoxeroides, L. hexandra, Z. latifolia and P. communis during early growth stages, florescence (or middle growth stages) and flower-fruit stages were measured. The results showed that: Midday depressions of aquatic A. philoxeroides and terrestrial A. philoxeroides were not found in April, but found in June. Midday depressions of terrestrial A. philoxeroides also occurred in October, and which of L. hexandra, Z. latifolia and P. communis did not occur at any growth stage except in April. The mean value of photosynthetic rate, 7r and LUE of aquatic A. philoxeroides and terrestrial A. philoxeroides expressed on a day were greater than that of three native aquatic plants during every growth stage. The value of maximal photosynthetic rate of A. philoxeroides was similar to C4 plants, and which was greater than 3 native aquatic plants. The mean value of WUE of aquatic A. philoxeroides expressed on a day was the maximal in April, and which only lower than Z latifolia in June. The values of apparent quantum yield (AQY) and carboxylation efficiency (CE) of aquatic A. philoxeroides and terrestrial A. philoxeroides were higher than that of 3 native aquatic plants. On the other hand, there were no differences in dark respiration rates (Rd) and light compensation point (LCP) between A. philoxeroides and 3 native aquatic plants. Light saturation point (LSP) of aquatic A. philoxeroide and terrestrial A. philoxeroides was similar to that of Z latifolia and P. communis, but greater than that of L. hexandra. CO2 compensation point (CCP) of aquatic A. philoxeroides was the minimal. CO2 saturation point (CSP) of aquatic A. philoxeroides and terrestrial A. philoxeroides was similar to that of Z latifolia, but lower than that of L. hexandra and P. communis, respectively.There were 13 native species (including 3 life forms: free-floating, emergent and submersed) co-existed in the floating mats of A. philoxeroides in Changgang cannal, and which belonged to 12 genera, 10 families. There were 2 species belong to Hydrocharitaceae, Najadaceae and Gramineae respectively, and only 1 species belong
    to other families. The biomass of emergent plants was the maximal and that o\' submersed plants was minimal, respectively.There were significant effects on the leaf width, spacer length, root length, plant height and leaf number of//, dubia caused by the mats of A. philoxeroides. However, the mat of A. philoxeroides had no effect on the leaf length, biomass per plant and clones number of H. dubia. The branch level, branches number, plant biomass and plant height of P. malaianus were decreased significantly when grew under the mats of A. philoxeroides. Plant density, winter bud density, leaf number and plant biomass of V. spinulosa were decreased with increasing the biomass of A. philoxeroides. Furthermore, no significant variation in plant height of V. spinulosa that grew under and outside the mat was found. The biomass per winter bud was increased when plants grew under the mat.
引文
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