克隆植物的无性与有性繁殖对策

许多植物同时具有克隆生长与有性繁殖,两种繁殖方式间的平衡在不同物种间以及同一物种内不同种群间变化很大。旺盛的克隆生长可能会从多方面影响生活史进化。首先,许多克隆植物的有性繁殖与更新程度都很低,甚至有一些植物由于克隆生长而几乎完全放弃了有性过程,从而影响到克隆植物对局域环境的适应和地理范围进化。其次,克隆生长增大花展示进而增加了对传粉者的吸引,同时也增加了同株异花授粉的风险,而同株异花授粉往往会导致植物雄性和雌性适合度的下降。因此,克隆植物的空间结构与交配方式间可能存在着协同进化关系。最后,克隆生长与有性繁殖间可能存在着权衡关系:对克隆生长的资源投入将会减少对有性繁殖的资源投入。这种权衡关系可能是由环境条件、竞争力度、植物寿命和遗传等因素决定的。如果不同的繁殖方式是植物在不同环境下采取的适应性对策,那么我们可以预期:在波动和竞争力度大的生境中,植物应将大部分的繁殖资源分配给有性繁殖;而在相对稳定的环境中,克隆繁殖应该占据优势地位。但是自然选择对两种繁殖方式的选择结果是什么,以及控制这两种方式间平衡的生态和遗传因子究竟有哪些,到底是克隆生长单向地影响了植物的有性繁殖,还是与有性过程相伴随的选择压力同时塑造了植物的克隆习性?目前尚不清楚。同时从无性与有性繁殖两个方面综合考察克隆植物的繁殖对策是今后亟待加强的工作。     

克隆植物对异质生境的适应对策研究进展

生境异质性是自然生态系统的基本特征,植物生长的必需资源和环境胁迫因子均存在着复杂的时间和空间异质性。克隆植物是指在自然条件下具有克隆特性的植物,即可通过与母株相连的芽、根茎、分蘖或枝条等繁殖体产生无性繁殖的植物,这些繁殖体一旦定居便可成为潜在的独立个体。克隆植物具有独特的生境适应策略(如形态可塑性、克隆整合、克隆分工、觅食行为、风险分摊等),面对异质性的生境条件,它可以通过调整自身的生理和形态结构来适应异质生境。目前,对于克隆植物在异质生境适应行为的研究已有很多报道,然而系统性的归纳和总结尚有欠缺。综述了克隆植物在不同资源异质生境(光照、养分、水分)和不同胁迫生境(盐碱胁迫、风沙胁迫、重金属胁迫)下独特的适应对策。最后,针对克隆植物对异质生境的适应对策,进行了总结并对未来的重点研究方向提出建议：(1)时间异质性尺度上的考量;(2)异质性生境中生物因子的调控作用;(3)克隆植物入侵机制;(4)克隆植物在生态修复中的应用潜力。     

草地植物种群繁殖对策研究

植物的繁殖包括有性繁殖和无性繁殖两大类型,克隆繁殖是一种较为特殊的营养繁殖方式。本文综述了草地种子植物的生殖分配及生殖投资,克隆生长以及放牧对草地植物种群繁殖的影响。植物种群生物量、能量和养分生殖分配是植物种群生殖分配的重要内容,不同植物在结实期营养元素及能量的配置上有着显著的区别,这可能是植物在长期进化过程中形成的生殖对策,是适应环境的结果。在种群水平上,中等强度以上的放牧干扰有利于植物的克隆生长,但有性生殖减弱。草原植物发达的营养繁殖或克隆生长方式是对放牧的适应性对策。     

论沙丘植被过程的基本生态关系

沙丘生态系统常常被认为是一种退化生态系统,自然植被恢复和稳定人工植被构建是沙丘生态系统恢复的基本手段。本文基于对沙丘生态系统长期研究结果,依从对立统一、动态变化哲学原理,结合相关生态学理论,从沙丘生态系统的独特性出发,提出了进行植被塑造过程研究需要注意的若干生态关系。论述了尺度-格局-过程的耦联关系和协同-权衡关系的转化在植被塑造过程研究中的必要性,阐释了区分沙丘流动与沙丘固定、干扰与胁迫、风蚀与沙埋在植被过程研究中的重要性,并探究了生态位法则或中性法则在沙丘植被过程研究中的应用价值,从适沙性与耐旱性、生理与繁殖过程、有性与无性繁殖等方面论述了沙丘植被塑造过程研究应关注的问题,旨在为沙丘生态系统植被自然恢复和稳定植被建设搭建理论构架。     

群落中克隆植物的重要性

综述了群落水平上克隆植物的重要性,克隆植物的生态习性,克隆植物的竞争关系等方面的研究进展,并试图在克隆植物竞争关系的背景下,结合其生境状况,探讨植物克隆性与植物物种多样性的关系。克隆植物的等级系统（基株—分株系统—分株）使其具有克隆性所赋予的多样的生活史、资源利用以及空间占据方式。列举了克隆植物的特性及其在群落中的作用。这些例子表明,克隆性强烈地影响和制约着植物群落的空间格局与竞争关系。然而克隆性也能通过权衡活动性与局部持久性来缓解对物种共存的抑制。克隆植物具有在时空尺度上分株间相互调节的机制,直接体现在群落中小尺度（个体与个体间）与大尺度（种群与种群间）间的相互作用上。丰富了传统的竞争和生态位划分理论,为群落中物种共存提供了合理解释。因此克隆植物在群落中的出现拓宽了群落系统潜在机制的范围。     

青藏高原高寒草甸植物花寿命与花吸引特征的关系及其对雌性繁殖成功的影响

花寿命指花保持开放且具有功能的时间长度, 是开花植物繁殖成功的一个重要功能性状。可塑性延长花寿命是植物在不利的传粉环境中保障繁殖的一种策略, 但延长花寿命也会增加繁殖成本。花寿命的可塑性变异不仅受传粉环境的影响, 而且还受资源分配权衡的影响。花寿命的理论模型指出, 植物的花寿命与花吸引特征之间存在资源分配权衡。为了研究在花粉限制环境中, 植物花寿命与花吸引特征之间的资源权衡及其对雌性适合度的相对重要性。该研究以青藏高原高寒草甸不同海拔(2 900和3 600 m)的11种开花植物为研究对象, 分析了不同植物群落中, 物种水平上: (1)花寿命与花吸引特征(花大小以及开花数目)之间的相关关系; (2)花寿命与花吸引特征对植物雌性适合度的相对贡献。结果表明, 无论是低海拔还是高海拔植物群落, 植物的花寿命与开花数目之间均存在权衡关系, 且长的花寿命增加了植物的雌性适合度。但在高海拔环境中, 植物的雌性适合度只与花寿命有关。这说明相对于低海拔植物, 花寿命对高海拔植物的雌性繁殖成功更为重要。     

根茎克隆植物生态学研究进展

根茎在植物的无性繁殖、克隆分株间信息交流和物质交换、预测资源斑块的质量等方面具有重要意义,并且根茎克隆植物的研究涉及生物入侵、全球变化等诸多生态学前沿领域。作为一种重要的克隆植物类型,根茎克隆植物在资源异质性生境中表现出特有的适应方式,这种方式可以通过形态可塑性、觅食行为、生理整合以及适合度来具体表征。着眼于根茎克隆植物,总结和分析了国内外近年来的研究案例,并对形态可塑性起源与多样性的限制假说和适应假说、觅食行为中的强度觅食和广度觅食策略、克隆分株间间隔子保持和断裂的利益权衡等热点内容进行了讨论。最后联系生态学学科前沿,提出了本领域在未来需要重视的研究方向。     

入侵植物的繁殖策略以及对本土植物繁殖的影响

理解入侵生物的繁殖策略是阐明生物入侵机制的一个重要方面。入侵植物常表现出一些共同的繁殖特征, 如以两性花为主的性系统、自动自交为主的繁育系统或不依赖传粉媒介的无融合生殖和无性繁殖以及高生殖投资的资源配置策略等。成功入侵的外来植物通过影响本土的传粉者, 在种群和群落水平上影响本土植物的有性繁殖, 甚至促使某些本土植物在繁殖对策和表型性状上发生快速转变。目前, 入侵植物繁殖策略及其生态效应的研究多侧重于入侵种的快速演化, 而有关外来植物与本土植物间的相互影响及其可能存在的协同适应研究还较为缺乏。探讨本土植物在外来种入侵压力下的繁殖对策和响应机制, 将丰富人们对物种间竞争、共存及群落构建等机制的深入了解。从繁殖和适应的角度探求入侵植物与本土植物之间的复杂关系, 将有助于解析生物入侵的机制及人类干扰下的物种演化规律, 也为预测和防控入侵植物提供科学依据。     

克隆植物对种间竞争的适应策略

克隆植物种群因其寿命的持久性、空间上的可移动性和繁殖方式的多样化等特征与非克隆植物有很大区别, 在自然生态系统中占有重要地位, 甚至成为优势种或者建群种。该文通过归纳有关克隆植物的种间竞争适应策略研究案例, 阐述了克隆植物的竞争能力差异和影响竞争力的因素; 论述了克隆植物在构件形态、克隆构型、繁殖对策等方面对种间竞争的响应, 以及生理整合作用与种间竞争的关系; 分析了导致某些同类研究的结论不一致的原因, 认为实验对象差异、实验设计、生境条件与克隆植物形态及生理上的时空动态变化等都可能影响实验结果; 提出了全球变化背景下的克隆植物种间竞争及其分子生态学机制等可能是今后需要重点关注的问题。     

异质光照环境下克隆整合对南美蟛蜞菊和蟛蜞菊种间关系的影响

很多入侵植物具有克隆性,克隆整合对入侵克隆植物生长和繁殖具有重要的贡献。自然界中,植物生长和繁殖所需的各种资源如光照、水分和矿质养分等在空间上分布通常是异质的,但关于异质环境下克隆整合对入侵植物和本土同属植物种间关系影响的研究相对缺乏。通过温室控制实验,将入侵植物南美蟛蜞菊(Wedelia trilobata)和同属本土植物蟛蜞菊(W.chinensis)的分株对单独种植或者混合种植在异质性光照条件下,同时通过保持或者切断分株之间的连接来控制克隆整合效应的有无,研究异质光照环境下克隆整合对南美蟛蜞菊和蟛蜞菊种间关系的影响。克隆整合对南美蟛蜞菊和蟛蜞菊的生长和繁殖都有促进作用,且南美蟛蜞菊比蟛蜞菊从克隆整合中获益更多。与单独种植相比,两者混种对南美蟛蜞菊的叶生物量有显著影响,而对本地种蟛蜞菊的根生物量有显著影响。克隆整合和种间关系对南美蟛蜞菊的总生物量和叶生物量产生了显著的交互作用,而对蟛蜞菊各指标无显著影响。克隆整合状态显著影响了南美蟛蜞菊和蟛蜞菊的种间关系。这些结果表明,异质环境下克隆整合可以改变入侵植物南美蟛蜞菊和本土植物蟛蜞菊的生长性状及种间关系。     

Asexual and sexual reproductive strategies in clonal plants

Most plants can reproduce both sexually and asexually (or vegetatively),and the balance between the two reproductive modes may vary widely between and within species.Extensive clonal growth may affect the evolution of life history traits in many ways.First,in some clonal species,sexual reproduction and sex ratio vary largely among populations.Variation in sexual reproduction may strongly affect plant's adaptation to local environments and the evolution of the geographic range.Second,clonal growth can increase floral display,and thus pollinator attraction,while it may impose serious constraints and evolutionary challenges on plants through geitonogamy that may strongly influence pollen dispersal.Geitonogamous pollination can bring a cost to plant fitness through both female and male functions.Some co-evolutionary interactions,therefore,may exist between the spatial structure and the mating behavior of clonal plants.Finally,a trade-off may exist between sexual reproduction and clonal growth.Resource allocation to the two reproductive modes may depend on environmental conditions,competitive dominance,life span,and genetic factors.If different reproductive modes represent adaptive strategies for plants in different environments,we expect that most of the resources should be allocated to sexual reproduction in habitats with fluctuating environmental conditions and strong competition,while clonal growth should be dominant in stable habitats.Yet we know little about the consequence of natural selection on the two reproductive modes and factors which control the balance of the two reproductive modes.Future studies should investigate the reproductive strategies of clonal plants simultaneously from both sexual and asexual perspectives.     

Asexual and sexual reproductive strategies in clonal plants

Most plants can reproduce both sexually and asexually (or vegetatively), and the balance between the two reproductive modes may vary widely between and within species. Extensive clonal growth may affect the evolution of life history traits in many ways. First, in some clonal species, sexual reproduction and sex ratio vary largely among populations. Variation in sexual reproduction may strongly affect plant’s adaptation to local environments and the evolution of the geographic range. Second, clonal growth can increase floral display, and thus pollinator attraction, while it may impose serious constraints and evolutionary challenges on plants through geitonogamy that may strongly influence pollen dispersal. Geitonogamous pollination can bring a cost to plant fitness through both female and male functions. Some co-evolutionary interactions, therefore, may exist between the spatial structure and the mating behavior of clonal plants. Finally, a trade-off may exist between sexual reproduction and clonal growth. Resource allocation to the two reproductive modes may depend on environmental conditions, competitive dominance, life span, and genetic factors. If different reproductive modes represent adaptive strategies for plants in different environments, we expect that most of the resources should be allocated to sexual reproduction in habitats with fluctuating environmental conditions and strong competition, while clonal growth should be dominant in stable habitats. Yet we know little about the consequence of natural selection on the two reproductive modes and factors which control the balance of the two reproductive modes. Future studies should investigate the reproductive strategies of clonal plants simultaneously from both sexual and asexual perspectives. Translated from Acta Phytoecologica Sinica, 2006, 20(1): 174–183 [译自: 植物生态学报]     

Trade-offs between clonal and sexual reproduction in Sagittaria latifolia (Alismataceae) scale up to affect the fitness of entire clones

? Many plants combine sexual reproduction with vegetative propagation, but how trade-offs between these reproductive modes affect fitness is poorly understood. Although such trade-offs have been demonstrated at the level of individual shoots (ramets), there is little evidence that they scale up to affect genet fitness. For hermaphrodites, reproductive investment is further divided between female and male sexual functions. Female function should generally incur greater carbon costs than male function, which might involve greater nitrogen (N) costs. ? Using a common garden experiment with diclinous, clonal Sagittaria latifolia we manipulated investment in reproduction through female and male sex functions of 412 plants from monoecious and dioecious populations. ? We detected a 1?:?1 trade-off between biomass investment in female function and clonal reproduction. For male function, there was no apparent trade-off between clonal and sexual reproduction in terms of biomass investment. Instead, male function incurred a substantially higher N cost. ? Our results indicate that: trade-offs between investment in clonal propagation and sexual reproduction occur at the genet level in S.?latifolia; and sexual reproduction interferes with clonal expansion, with investment in female function limiting the quantity of clonal propagules produced, and investment in male function limiting the nutrient content of clonal propagules.     

Sexual and vegetative reproduction in the aboveground part of a dioecious clonal plant, <Emphasis Type="Italic">Dioscorea japonica</Emphasis> (Dioscoreaceae)

In dioecious clonal plants, the reproductive effort required to set seeds will be responsible for the larger investment in sexual reproduction by females. If there will be a trade-off in resource allocation between sexual and clonal reproduction, this differential sexual reproduction will lead to sexual differentiation in the relative amount of clonal reproduction. To test this prediction, we studied differences between the sexes in their phenologies and investments in sexual and vegetative reproduction (clonal reproduction by means of bulbils) with respect to ramet size in a dioecious clonal plant, Dioscorea japonica Thunb. The period of bulbil production overlapped the period during which infructescences developed. Females flowered later, produced heavier inflorescences, and fewer flowers per inflorescence than did males. Regression analysis using the size of the individual plants demonstrated that large females made smaller investments in inflorescences and larger investments in sexual reproduction than did large males. In contrast, females invested fewer resources in vegetative reproduction than did males. However, the total investments in sexual and vegetative reproduction did not differ between the sexes. These results supported our hypothesis on the sexual differentiation in sexual and clonal reproduction.     

Modeling the impact of reproductive mode on masting

Masting is defined as the intermittent highly variable production of seed in a plant population. According to reproductive modes, that is, sexual and asexual reproduction, masting species can be separated into three groups, that is, (1) species, for example, bamboo, flower only once before they die; (2) species, for example, Fagus, reproduce sexually; and (3) species, for example, Stipa tenacissima, reproduce both sexually and asexually. Several theories have been proposed to explore the underlying mechanisms of masting. However, to our knowledge, no theory has been found to explain the mechanism of masting species that reproduce both sexually and asexually. Here we refine the Resource Budget Model by considering a trade‐off between sexual and asexual reproduction. Besides the depletion efficient (i.e., the ratio of the cost of seed setting and the cost of flowering), other factors, such as the annual remaining resource (i.e., the rest of the resource from the photosynthetic activity after allocating to growth and maintenance), the trade‐off between sexual and asexual reproduction, and the reproductive thresholds, also affect masting. Moreover, two potential reproductive strategies are found to explain the mechanisms: (1) When the annual remaining resource is relatively low, plants reproduce asexually and a part of the resource is accumulated as the cost of asexual reproduction is less than the annual remaining resource. Plants flower and set fruits once the accumulated resource exceeds the threshold of sexual reproduction; (2) when the annual remaining resource is relatively high, and the accumulated resource surpasses the threshold of sexual reproduction, masting occurs. Remarkably, under certain depletion efficient, more investigation in sexual reproduction will lead plants to reproduce periodically. Additionally, plants investigate less resource to reproduce periodically when depletion efficient keeps increasing as plants can reproduce efficiently. Overall, our study provides new insights into the interpretation of masting, especially for species that reproduce both sexually and asexually.     

Genetic variation and evolutionary trade-offs for sexual and asexual reproductive modes in Allium vineale (Liliaceae)

Populations of Allium vineale commonly include individuals with very different allocation patterns to three modes of reproduction: sexual flowers, aerially produced asexual bulbils, and belowground asexual offsets. If selection is currently acting to maintain these different allocation patterns there must be a genetic basis for variation in allocation to these three reproductive modes. In addition, negative genetic correlations between reproductive traits would imply evolutionary trade-offs among reproductive strategies. We evaluated the heritability of these allocation patterns by growing 16 clones from a single population in the greenhouse at two levels of fertilization. Bulb mass and the mass and number of bulbils, offsets, and flowers were used as response variables, in addition to the proportion allocated to each reproductive mode. We found evidence of substantial heritable variation in allocation to sexual reproduction and in allocation within the two modes of asexual reproduction, indicating high sensitivity of these allocation patterns to natural selection. We also found evidence of strong negative genetic correlations between bulbil and flower traits, as well as between bulbil and offset traits, with one group of genotypes allocating greater resources to aerial asexual bulbils and the second group allocating more resources to belowground asexual offsets and aerial flowers. Phenotypic plasticity in allocation to above- vs. belowground asexual reproduction and sexual vs. asexual aerial reproduction was limited, indicating that plants are unlikely to change reproductive mode in response to nutrient availability. Together, then, we have demonstrated strong heritability for, and trade-offs in, the reproductive allocation patterns within this plant population.     

Evolutionary increase in sexual and clonal reproductive capacity during biological invasion in an aquatic plant Butomus umbellatus (Butomaceae)

To test the hypothesis that increased allocation to reproduction is selected during biological invasion, we compared germination, survival, growth, and reproduction of native vs. introduced populations of the invasive aquatic plant Butomus umbellatus in a common greenhouse environment. Although seedling emergence and establishment did not differ consistently, survival thereafter was twice as high for eight introduced North American than eight native European populations. As predicted, introduced plants were more likely to produce sexual inflorescences and clonal asexual vegetative bulbils, and they invested much more biomass in both reproductive modes. Higher reproductive investment was due to higher proportional allocation of biomass rather than larger plant size. These results are consistent with selection for increased reproduction during range expansion. However, population genetic surveys indicate that recruitment from seed rarely occurs in introduced populations. Hence increased sexual allocation is not an adaptive response to invasion. Although increased clonal reproduction may be advantageous in expanding populations, genetic evidence from introduced populations of B. umbellatus suggests that increased clonal allocation may have arisen via stochastic processes during long-distance transport or a selective filter right at introduction, rather than incremental natural selection during range expansion.     

An experimental demonstration of the cost of sex and a potential resource limitation on reproduction in the moss Pterygoneurum (Pottiaceae)

The cost of sexual reproduction is incurred when the current reproductive episode contributes to a a decline in future plant performance. To test the hypotheses that a trade-off exists between current sexual reproduction and subsequent clonal regeneration and that resources limit reproduction and regeneration, plants of the widespread moss Pterygoneurum ovatum were subjected to induced sporophytic abortion, upper leaf removal, and nutrient amendment treatments. Sexually reproducing plants were slower or less likely to produce regenerative structures (protonemata or shoots) and produced fewer regenerative tissue areas or structures. The ability and the timeline to reproduce sexually and regenerate clonally were unaffected by an inorganic nutrient amendment. However, when leaves subtending the sporophyte were removed, the sporophytes were less likely to mature, tended to take a longer time to mature, and were smaller compared to sporophytes from shoots with a full complement of upper leaves. Our findings indicate that plants investing in sexual reproduction suffer a cost of decreased clonal regeneration and indicate that sporophyte maturation is resource-limited, with upper leaves contributing to the nutrition of the sporophyte. This study represents only the second explicit experimental demonstration of a trade-off between sexual and asexual reproduction in bryophytes.     

Trade-offs between sexual and asexual reproduction in a monoecious species Sagittaria pygmaea (Alismataceae): the effect of different nutrient levels

Available resources could influence the trade-offs among different reproductive components in plants. Here, we created three nutrient levels to test the nutrient effects on trade-offs among sexual reproduction, clonal propagation and vegetative growth in a monoecious clonal herb Sagittaria pygmaea. The results of this study showed that the plant exhibited different trade-off patterns among different nutrient levels. When the nutrient level was low, there were weak trade-offs between sexual reproduction and vegetative growth and between clonal propagation and vegetative growth; when the nutrient level was moderate, we found a strong trade-off between sexual reproduction and clonal propagation; but when the nutrient level was high, we found no trade-offs among these three different reproductive components. These results indicated that the plant could adjust its trade-off patterns to fit the nutrient variation and suggested that trade-offs are unlikely to constrain the evolution of reproductive strategy in this species.