无性系植物种群生态学研究进展及有关概念

无性系植物种群生态学研究进展及有关概念刘庆钟章成（中国科学院成都生物研究所,６１００４１）（西南师范大学生物系,重庆６３０７１５）ＡｄｖａｎｃｅｓｉｎＥｃｏｌｏｇｉｃａｌＲｅｓｅａｒｃｈｏｆＣｌｏｎａｌＰｌａｎｔＰｏｐｕｌａｔｉｏｎａｎｄＳｏｍｅＲｅ...     

Fitness and evolution in clonal plants: the impact of clonal growth

Seeds have often been emphasized in estimates of plant fitness because they are the units that carry genes to the next generation, disperse, and found new populations. We contend that clonal growth also needs to be considered when estimating fitness in clonal plants, regardless of whether fitness is measured from a genet or ramet perspective. Clonal growth affects genet fitness through both genet persistence and seed production. It affects ramet fitness through new ramet production, because both seeds and clonal propagants are considered offspring. The differential production of clonal propagants will contribute to fitness differences among individuals which may result in population-level changes in allele frequencies (i.e. microevolution). We describe a form of selection unique to clonal organisms, genotypic selection, that can result in evolution. Genotypic selection occurs when genotypically based traits are associated with differences in the rate of ramet production. It can lead to evolutionary change in quantitative trait means both directly and indirectly. It leads directly to change in the ramet population by increasing the proportion of ramets with more advantageous trait values. From the genet perspective, it leads indirectly to evolution within and among populations whenever significant portions of the genetic effect on a trait are inherited through seed. We argue that under most conditions, clonal growth will play a major role in the microevolution of clonal plants.     

关于无性系植物种群整合作用（Integration）研究的现状及其应用前景

关于无性系植物种群整合作用（Ｉｎｔｅｇｒａｔｉｏｎ）研究的现状及其应用前景王昱生（东北农业大学动物科学系草地教研室,哈尔滨１５００３０）ＩｎｔｅｇｒａｔｉｏｎｉｎＣｌｏｎａｌＰｌａｎｔＰｏｐｕｌａｔｉｏｎｓ：ＣｕｒｒｅｎｔＳｉｔｕａｔｉｏｎｏｆｔｈｅ...     

Genet structure and determinants of clonal structure in a temperate deciduous woodland herb, Uvularia perfoliata

1 We used isozyme variation to examine the genet structure of Uvularia perfoliata patches in gap and closed canopy habitats in a temperate deciduous forest in Maryland, USA.
2 A large patch in a gap habitat was composed of a small number of widely spread genets with many ramets, and a large number of genets with more restricted distribution and few ramets. Genets with many ramets were patchily distributed at a metre scale. Analysis of genet structure on a scale of square centimetres, however, revealed that the genets were highly intermingled with no clear boundaries between them. The presence at both scales of sampling of many genets with unique multilocus genotypes indicated continuing genet recruitment within the population.
3 In the closed canopy habitat, the patches examined were each composed of a single unique multilocus genotype, suggesting that each had developed by asexual propagation following the establishment of a single genet.
4 The clonal structure of U. perfoliata patches in both gap and closed canopy habitats therefore appears to depend on recruitment patterns of genets. Populations in closed canopy habitats are characterized by a 'waiting' strategy, in which asexual ramet production maintains populations until genet recruitment by seed production can occur under the more optimal conditions associated with canopy gaps. Extended sampling suggests that the genetic diversity of U. perfoliata populations is primarily controlled by the disturbance regime of the forest canopy.     

Fine-scale clonal structure and diversity within patches of a clone-forming dioecious shrub, Ilex leucoclada (Aquifoliaceae)

BACKGROUND AND AIMS: The mode of reproduction (sexual vs. asexual) is likely to have important effects on genetic variation and its spatial distribution within plant populations. An investigation was undertaken of fine-scale clonal structure and diversity within patches of Ilex leucoclada (a clone-forming dioecious shrub). METHODS: Six patches were selected in a 1-ha plot previously established in an old-growth beech forest. Two of the selected patches were composed predominantly of stems with male flowers (male patch), and two contained stems with predominantly female flowers (female patch). The remaining two patches contained stems with male flowers and stems with female flowers in more or less equal proportions (mixed patch). Different genets were distinguished using random amplified polymorphic DNA (RAPD) markers. KEY RESULTS: One hundred and fifty-six genets with different RAPD phenotypes were identified among 1928 stems from the six patches. Among the six patches, the male patches had the lowest clonal diversity, and the mixed patches had the highest. Distribution maps of the genets showed that they extended downhill, reflecting natural layering that occurred when stems were pressed to the ground by heavy snow. In every patch, there were a few large genets with many stems and many small genets with a few stems. CONCLUSION: The differences in clonal diversity among patches may be due to differences in seedling recruitment frequencies. The skewed distribution of genet size (defined as the number of stems per genet) within patches may be due to differences in the timing of germination, or age (with early-establishing genets having clear advantages for acquiring resources) and/or intraspecific competition.     

Sexual differences in physiological integration in the dioecious shrub Lindera triloba: a field experiment using girdling manipulation

Background and Aims

It is important to consider the modular level when verifying sexual dimorphism in dioecious plants. Nevertheless, between-sex differences in resource translocation among modules (i.e. physiological integration) have not been tested at the whole-plant level. In this study, sexual differences in physiological integration were examined among ramets, within a genet in the dioecious sprouting shrub Lindera triloba, by a field experiment with girdling manipulation.

Methods

Female and male genets were randomly assigned to girdled or intact groups. Girdling of the main ramets was conducted in May 2009 by removing a ring of bark and cambium approx. 1 cm wide at a height of 80–100 cm. The effects of treatment and sex on ramet dynamics (mortality, recruitment and diameter growth) and inflorescence production during 1 year after girdling were examined.

Key Results

The diameter growth rate of main ramets of both sexes was lower at ground level (D₀) but higher at breast height (dbh) in girdled than in intact groups. In sprouted ramets with a dbh of 0–2 cm, males in girdled groups had lower growth rates at D₀ than those of intact groups, whereas no girdling effect was found for females. The main ramets in girdled groups produced more inflorescences than intact groups, irrespective of sex, but male ramets showed a greater response to the treatment than females.

Conclusions

In L. triloba, physiological integration exists at the whole-plant level, and sprouted ramets are dependent on assimilates translocated from main ramets, but this dependence weakens as sprouted ramets get larger. Female sprouted ramets can grow in a physiologically independent manner from the main ramet earlier than those of males. This study highlights the importance of considering modular structures and physiological integration when evaluating sexual differences in demographic patterns of clonal plants.     

Investigating the community consequences of competition among clonal plants

Although clonal plants comprise most of the biomass of several widespread ecosystems, including many grasslands, wetlands, and tundra, our understanding of the effects of clonal attributes on community patterns and processes is weak. Here we present the conceptual basis for experiments focused on manipulating clonal attributes in a community context to determine how clonal characteristics affect interactions among plants at both the individual and community levels. All treatments are replicated at low and high density in a community density series to compare plant responses in environments of different competitive intensity. We examine clonal integration, the sharing of resources among ramets, by severing ramets from one another and comparing their response to ramets with intact connections. Ramet aggregation, the spacing of ramets relative to each other, is investigated by comparing species that differ in their natural aggregation (either clumped growth forms, with ramets tightly packed together, or runner growth forms, with ramets loosely spread) and by planting individual ramets of all species evenly spaced throughout a mesocosm. We illustrate how to test predictions to examine the influence of these two clonal traits on competitive interactions at the individual and community levels. To evaluate the effect of clonal integration on competition, we test two predictions: at the individual level, species with greater clonal integration will be better individual-level competitors, and at the community level, competition will cause a greater change in community composition when ramets are integrated (connected) than when they are not. For aggregation we test at the individual level: clumped growth forms are better competitors than runner growth forms because of their ability to resist invasion, and at the community level: competition will have a greater effect on community structure when ramets are evenly planted. An additional prediction connects the individual- and community-level effects of competition: resistance ability better predicts the effects of competition on relative abundance in a community than does invasion ability. We discuss additional experimental design considerations as revealed by our ongoing studies. Examining how clonal attributes affect both the individual- and community-level effects of competition requires new methods and metrics such as those presented here, and is vital to understanding the role of clonality in community structure of many ecosystems.     

辽东栎林下和开阔地上短柄五加(Acanthopanax brachpus)无性系种群生长的比较研究

研究了分布于开阔地和辽东栎林下短柄五加无性系种群拆散方式、无性系的形态可塑性和分株的死亡年龄,研究结果表明,开阔地上无性系分株的扩散方式为稀疏线型,根茎长度、无性系分株数目,林下和开阔地差异显著,根茎分枝角度、无性系根茎总长无显著性差异。林下分株的死亡年龄显著小于开阔地上无性系分株的死亡年龄。     

入侵植物剑叶金鸡菊克隆繁殖特性及群落特征

以江西庐山剑叶金鸡菊（Coreopsis lanceolata）入侵群落为研究对象,调查了不同生境剑叶金鸡菊的克隆繁殖特性和群落特征。结果表明,生境资源水平高低对剑叶金鸡菊的克隆构型影响不显著,无论是资源贫乏的岩石还是资源丰富的弃耕地生境,剑叶金鸡菊依靠植株基部节处萌生的小芽体生长成密集丛生状,其游击型克隆构型变化与环境资源可利用水平高低无关。剑叶金鸡菊种群的伴生植物多为一些适应范围广的杂草,由于剑叶金鸡菊的密集型克隆生长的集团化生长策略,使其种群耐贫瘠和干旱能力非常强,因而适应不同生境的能力也非常强,这种生长策略使剑叶金鸡菊个体能快速扩增并排挤本土植物,〖JP〗形成大面积的单一优势种群影响当地的生物多样性。     

A comparison of plant communities on the basis of their clonal growth patterns

This study was performed to analyse how a vegetative propagation pattern of plants affects the coexistence of species and subsequent species richness of the community. We compared community average clonal growth in the herbal communities of forests, wooded meadows, and open meadows in Laelatu, Estonia. The parameters used for the calculation of the community averages and measured for each species were ramet life span, rhizome branching, and clonal mobility. We also examined the intrinsic (i.e. independent of the environment) relationship between community clonal growth and plant species density. We found strong correlations between the environmental factors (productivity, light availability, and mowing regime) and community averages of clonal growth parameters, while species density was (negatively) correlated only with community average of rhizome increment. The community average of ramet life span decreased with the increasing biomass of the herb layer. No evidence was found to support the hypothesis that species-rich communities may consist of species with more contrasting mobility compared with species-poor communities. Independent of the effect of the environmental factors, species density was positively correlated with ramet density. There was intrinsic positive relationship between species density and community average of ramet life span at open meadow sites and intrinsic negative relationship between species density and community average of rhizome increment at wooded meadow sites. We conclude that in forest communities the capability of clonal plants to forage for light is favoured, while in unmown meadows a competitively strong phalanx growth form is advantageous. We established that ramet turnover increases and vegetative mobility decreases with increasing species diversity, although these two relationships depend strongly on the type of the studied community.Co-ordinating editor: J. Tuomi