竹叶眼子菜居群遗传多样性和克隆结构

采用ISSR技术对长江中游南岸豹澥湖和大冶湖不同生境中的竹叶眼子菜(Potamogeton malaianus)居群的遗传多样性及克隆结构进行了研究.结果表明,在两居群的106株个体中,利用6条ISSR引物共得到40条符合3/N标准并无连锁不平衡的清晰位点,竹叶眼子菜具有较高的遗传多样性,其多态位点百分率为75.0％,Shannon多样性指数为0.3736, 两居群的遗传分化很小.竹叶眼子菜的克隆多样性很高(D=0.9917),两居群间的克隆分化很大,不具有共有的基因型.竹叶眼子菜的基株分布为游击型构型,位于湖心的大冶居群克隆距离(3.0～31.5 m)明显地大于湖岸豹澥居群(2.4～6.7 m).     

Interactions between root and shoot competition vary among species

Understanding how the competition varies with productivity is essential for differentiating among alternative models of plant community organization. Prior attempts to explain shifts in root and shoot competition along gradients have generally assumed an additive interaction between the two competitive forms, using an experimental design which does not fully separate both above‐ and belowground processes. At the most basic level, few field studies have separated root and shoot competition, and we have limited knowledge about both the relative importance of these processes, and how they interact to affect plant growth in the field. Presented here are findings from a field study in which root and shoot competition were experimentally separated by using root exclusion tubes and neighbor tiebacks in an early successional community. Individuals of four species (Abutilon theophrasti, Amaranthus retroflexus, Rumex crispus, and Plantago lanceolata) were grown at two levels of fertilization with full competition, aboveground competition only, belowground competition only, or neither above‐ nor belowground competition. Competition was measured as competitive response, which is the natural log of the relative biomass of a target plant grown with competition compared to growth without competition. In contrast to predictions from current models of productivity‐competition relationships, but in agreement with other experimental studies, there was no change in the strengths or root, shoot, or total competition with a modest increase in productivity. Despite no effect of fertilization on the strength of competition, the form of interaction between root and shoot competition varied both as a function of species identity and fertilization. For both of the rosette forming species, the combined effects of root and shoot competition were less than predicted assuming no interaction (a “negative interaction”), with one species switching from a negative to an additive interaction with fertilization. The fact that fertilization caused a shift in the root‐shoot interaction, but not in the total strength of root and shoot competition, suggests that the root‐shoot interaction is itself a highly labile variable. If root‐shoot interactions are common in natural systems, then simply measuring the strength of one form of competition in no way provides any information about the overall importance of that competitive form to plant growth.     

Disturbance,competition and the maintenance of clonal diversity in Daphnia pulex

Laboratory microcosm experiments tested the intermediate disturbance hypothesis, which states that the highest level of diversity (e.g. species diversity) will be maintained at intermediate scales of disturbance. The effects of disturbance on the maintenance of clonal diversity and on competitive interactions among clones of the obligately parthenogenetic freshwater cladoceran, Daphnia pulex were examined. No significant effect of disturbance size (i.e. dilution volume) on clonal diversity was noted. However, frequency of disturbance had a pronounced effect on clonal diversity, with the highest clonal diversity maintained at low to intermediate disturbance frequencies. Competitive hierarchies among clones were often invariant within a given experiment. Generally, one or two clones dominated, with several less abundant clones persisting throughout an experiment. Results suggest that low to intermediate disturbances could be important in the maintenance of genetic variation in natural populations (i.e. through pre-emption of competitive exclusion between genotypes). This could have a direct bearing on the maintenance of both intra- and interspecific diversity.     

Parasites, sex, and clonal diversity in natural snail populations

Under the Red Queen hypothesis, host-parasite coevolution selects against common host genotypes. Although this mechanism might underlie the persistence of sexual reproduction, it might also maintain high clonal diversity. Alternatively, clonal diversity might be maintained by multiple origins of parthenogens from conspecific sexuals, a feature in many animal groups. Herein, we addressed the maintenance of overall genetic diversity by coevolving parasites, as predicted by the Red Queen hypothesis. We specifically examined the contribution of parasites to host clonal diversity and the frequency of sexually reproducing individuals in natural stream populations of Potamopyrgus antipodarum snails. We also tested the alternative hypothesis that clonal diversity is maintained by the input of clones by mutation from sympatric sexuals. Clonal diversity and the frequency of sexual individuals were both positively related to infection frequency. Surprisingly, although clones are derived by mutation from sexual snails, parasites explained more of the genotypic variation among parthenogenetic subpopulations. Our findings thus highlight the importance of parasites as drivers of clonal diversity, as well as sex.     

Variation in growth rate in shoot populations of the clonal dwarf shrub Linnaea borealis

This study was based on two years of field observations of a Linnaea borealis population in a coniferous forest in Sweden. Detailed information on performance and survivorship of shoots were gathered from consecutively made drawings of 90 individually marked shoot systems of Linnaea distributed among 15 plots in the forest. Branching of the shoot systems was correlated to growth of the main shoot. Increasing light flux negatively affected growth of Linnaea . A tendency for a competitive effect on Linnaea of Vaccinium spp. was also found. In contrast, the surrounding mosses, and intraspecific density of Linnaea had no significant effect on the shoot systems. The growth rate of the shoot population was analysed by matrix models. Population growth rate was most dependent on survivorship and growth of main shoots in Linnaea , while the survivorship of lateral shoots was of less importance. In spite of an average decrease in the population size as revealed by the matrix model simulation, the incorporation of the observed spatial variation in shoot growth increment, branching and shoot survivorship, yielded varying population growth rates. Parts of the population increased in size. The importance of choice of spatial scale in population analyses of forest understory plants is emphasized.     

Comparison of clonal diversity in mountain and Piedmont populations of Trillium cuneatum (Melanthiaceae-Trilliaceae), a forest understory species

The balance between clonal and sexual reproduction can vary widely among plant populations, and the extent of clonality may be influenced by the combined effects of historical land use and variation in environmental conditions. We investigated patterns of clonal spread in five Trillium cuneatum populations, two in the Appalachian Mountains characterized by mesic, cooler conditions, and three at lower elevations experiencing warmer, drier conditions and greater disturbance. Using a new measure of the genet effective number and innovative orthogonal contrast methods, we quantified genet structure, contrasting clonal growth in the mountains with that in the Piedmont. Asexual propagation was more common in the Piedmont, where 25% of the sampled ramets were clonally derived, but was much less frequent in the mountains (7% clonal replicates). Hierarchical partitioning of variation in genet diversity showed that the majority (75.8%) of the variation resulted from more vegetative replication in the Piedmont. Most of the remaining variation (21.6%) was attributable to differences between urban and rural Piedmont populations, and a small, statistically nonsignificant fraction of the variation (2.6%) was due to interpopulation differences within the mountains. Higher frequency of cloning may enhance both genetic and demographic population viability in fragmented Piedmont habitats.     

Genetic and clonal diversity in Korean populations ofVitex rotundifolia (Verbenaceae)

Vitex rotundifolia L.f. is a woody perennial and has sexual and asexual modes of reproduction. Allozyme study was conducted on 550 plants in 13 Korean populations. The levels of genetic variability and divergence within and among populations, respectively, are considerably lower and higher than the mean values for woody plants with similar life history tralts. Mean percentage of polymorphic loci (P _P), mean number of alleles per locus (A _P), and mean genetic diversity (He _P) within populations ofV. rotundifolia were: 16.7%, 1.21, and 0.047. On average, about 79% of the total variation inV. rotundifolia was common to all populations (meanG _ST=0.208). In addition, significant differences in allele frequencies among populations were found in all polymorphic loci examined (P<0.001). On the other hand, levels of genotypic diversity within and among populations were moderate. About 44% (18/41) of multilocus genotypes were “local genotypes” (genotypes occurring in only one population), whereas only one “widespread genotype” (genotypes occurring in more than 75% of the populations) were detected. The mean number of multilocus genotypes per population (G) and mean genotypic diversity index (D _G) were 8.4 and 0.74, respectively. Most common multilocus genotypes found in populations were homozygous for five polymorphic loci. The abundance of ramets of these genets is responsible for the low levels of expected heterozygosity within populations. The results indicate that clonal reproduction may act as an enhancer of genetic drift by reducing effective size of local populations ofV. rotundifolia.     

Clonal diversity,clonal persistence and rapid taxon replacement in natural populations of species and hybrids of the Daphnia longispina complex

Hybridization is common among cyclical parthenogens, especially in zooplankton species assemblages of the genus Daphnia. To explore hybridization dynamics and the extent of clonal diversity in the Daphnia longispina complex, we analysed population structure in eight permanent lakes. Based on 15 microsatellite loci, three major taxonomic units emerged: two species, D. galeata and D. longispina and their F1 hybrids, supported by factorial correspondence analysis and two Bayesian methods. At the same time, the detection of backcross classes differed between methods. Mean clonal diversity was lowest in the F1 hybrids, as expected from the high rate of asexual reproduction. Within taxa, replicated genotypes were of clonal origin, but clonal lineages persisted in subsequent years in only one of three resampled lakes. In another lake, the taxon composition changed from being dominated by hybrids to complete dominance by one parental taxon. Such a year‐to‐year taxon replacement has not been reported for the D. longispina complex before. Our data on this hybrid complex illustrate that high‐resolution genotyping is essential for the understanding of ecological and evolutionary outcomes of hybridization in partially clonal taxa.     

Genetic diversity in three invasive clonal aquatic species in New Zealand

Background

Elodea canadensis, Egeria densa and Lagarosiphon major are dioecious clonal species which are invasive in New Zealand and other regions. Unlike many other invasive species, the genetic variation in New Zealand is very limited. Clonal reproduction is often considered an evolutionary dead end, even though a certain amount of genetic divergence may arise due to somatic mutations. The successful growth and establishment of invasive clonal species may be explained not by adaptability but by pre-existing ecological traits that prove advantageous in the new environment. We studied the genetic diversity and population structure in the North Island of New Zealand using AFLPs and related the findings to the number of introductions and the evolution that has occurred in the introduced area.

Results

Low levels of genetic diversity were found in all three species and appeared to be due to highly homogeneous founding gene pools. Elodea canadensis was introduced in 1868, and its populations showed more genetic structure than those of the more recently introduced of E. densa (1946) and L. major (1950). Elodea canadensis and L. major, however, had similar phylogeographic patterns, in spite of the difference in time since introduction.

Conclusions

The presence of a certain level of geographically correlated genetic structure in the absence of sexual reproduction, and in spite of random human dispersal of vegetative propagules, can be reasonably attributed to post-dispersal somatic mutations. Direct evidence of such evolutionary events is, however, still insufficient.     

Growth patterns of Juncus gerardi clonal populations in a coastal habitat

Juncus gerardi populations demonstrated a logistic growth curve during the colonization stage. Shoot production by vegetative multiplication was virtually continuous from December to June. Experiments suggested that the stabilisation stage of the demographic curve reflected water deficit. Taller, fertile, winter and early spring cohorts could be distinguished from shorter, infertile end of spring and beginning of summer cohorts. Shoot emergence began in March and terminated at the end of June, when water becomes a limiting factor due to a period of water shortage, typical of the thermo-atlantic climate. Spatial extension of populations was due to rhizome growth, which ceased during flowering.Flowering in May temporarily checked growth in shoot height of all emerged cohorts. No cost of reproduction was demonstrated concerning the rate of appearance of new shoots.Although fertile shoots were taller than vegetative shoots, their growth rates were significantly lower from April onwards. The tallest fertile shoots produced the most capsules.Energy allocation to seed production is the only possible means for long-term establishment of new genotypes, and vegetative multiplication appears as the principal source of recruitment of new modules in Juncus gerardi.Resource allocation patterns in this clonal species are discussed in relation to the ecological background in the concerned marshlands and with theoretical proposals derived from models of spatial colonization strategies in clonal plants.Nomenclature: follows Flora Europaea (Tutin et al., 1964ndash;1980).     

Habitat diversity and species diversity: testing the competition hypothesis with juvenile salmonids

I experimentally tested two predictions of the hypothesis that the positive relationship between habitat diversity and species diversity arises from a reduction in the negative effects of interspecific competition. By allowing species to partition habitat and avoid competition, habitat diversity should 1) facilitate the addition of an ecologically intermediate species into an existing community, and 2) reduce the negative effects of that species on existing members of the community. I tested these predictions with juveniles of three sympatric salmonid species: coho salmon ( Oncorhynchus kisutch ), steelhead trout ( O. mykiss ), and cutthroat trout ( O. clarki ), which in natural streams occupy deep low-velocity pools, shallow high-velocity riffles, and intermediate habitats, respectively. I introduced two (coho and steelhead) and three species communities into each of three artificial stream habitats: pools, riffles, and diverse. The results provide partial support for the predictions. Cutthroat trout grew fastest in the diverse stream habitat. Though habitat diversity did not eliminate the negative effects of competition, in the three species community coho and steelhead grew as fast in the diverse habitat as in either homogeneous habitat. The results are consistent with data on species number and evenness from natural communities, where variation along other niche axes confounds the relationship between habitat diversity, interspecific competition, and species diversity.     

Importance of clonal plants and plant species diversity in the Northeast China Transect

In plant communities, the internal (genet-level) control mechanisms on a spatio-temporal scale of clonal plants impose strong constraints on spatial pattern as well as on competitive relations and, thus, species coexistence. Therefore, the presence of clonal species within a plant community affects spatio-temporal dynamics and plant species diversity. We examined the distribution of plants with different clonal growth forms in the Northeast China Transect (NECT) and correlated plant species diversity with the importance of clonal plants, and the importance of phalanx and guerilla clonal plants. Phalanx clonal plants were more abundant in western communities where the altitude was higher and both the soil nitrogen contents and precipitation were relatively low. Whereas guerilla clonal plants were more abundant in the middle of the NECT where the precipitation, mean annual temperature and photosynthetically active radiation were relatively high. In the relatively productive temperate typical steppe, plant species diversity was negatively correlated with the importance of phalanx clonal plants and positively correlated with the importance of guerilla clonal plants. In relatively unproductive temperate desert steppe, plant species diversity was positively correlated with the importance of both phalanx and guerilla clonal plants.     

Effects of interspecific competition on asexual proliferation and clonal genetic diversity in larval trematode infections of snails

Interactions among different parasite species within hosts can be important factors shaping the evolution of parasite and host populations. Within snail hosts, antagonistic interactions among trematode species, such as competition and predation, can influence parasite abundance and diversity. In the present study we examined the strength of antagonistic interactions between 2 marine trematodes (Maritrema novaezealandensis and Philophthalmus sp.) in naturally infected Zeacumantus subcarinatus snails. We found approximately the same number of snails harbouring both species as would be expected by chance given the prevalence of each. However, snails infected with only M. novaezealandensis and snails with M. novaezealandensis and Philophthalmus sp. co-occurring were smaller than snails harbouring only Philophthalmus sp. In addition, the number of Philophthalmus sp. rediae was not affected by the presence of M. novaezealandensis sporocysts and the within-host clonal diversity of M. novaezealandensis was not influenced by the presence of Philophthalmus sp. Our results suggest that antagonistic interactions may not be a major force influencing the evolution of these trematodes and that characteristics such as host size and parasite infection longevity are shaping their abundance and population dynamics.     

Genet age in marginal populations of two clonal Carex species in the Siberian Arctic

During a Swedish-Russian expedition to northern Siberia 1994, we sampled two marginal populations of two Carex species at two high arctic sites ( C, stans Drej. on Faddeyevsky Island and C. ensifolia V. Krecz ssp. arctisibirica Jurtz. at north-eastern Taymyr Peninsula), both north of previously documented localities in that areas for the two species. These populations were composed of a few distinct patches of ramet colonies, some of them shaped like fairy rings with dead centres. We measured the size of all colonies and collected samples for detailed morphometric analyses of rhizome growth. By using RAPD (random amplified polymorphic DNA) analysis we established that the largest colony at each site consisted of a single genet, based on 41 polymorphic bands amplified with three primers. Pooled samples from each of two additional colonies of C. stans on Faddeyevsky Island were analysed and showed that clones of the same species at the same site were relatively dissimilar (Dice's similarity index 0.26 0,43), We then assumed that each ramet colony represented a single genet. Based on the morphometric data, we developed a deterministic growth model that simulates the clonal growth of these species and enabled estimates of the time since establishment of the genets. The estimated age of the five C. Stans clones varied from 17 to 154 yr and the age of the two C. ensifolia ssp, arctisibirica clones was well over 3000 yr.     

Intergenotypic competition and shoot production in wheat plants

In field experiments, the competition among plants of two distinct wheat genotypes in binary mixtures, and among three genotypes being grown in adjacent rows was investigated. The competition greatly affected the over-ground dry weight, weight of stem and grain per plant. This effect was undercompensatory, the depression in poor competitor was higher than the enhancement of strong competitor for all these characteristics. From final components of grain yield per plant, the number of fertile stems and number of grains per ear were more affected than weight per grain. More expressively than weight of grain on the main-stem ear, the weight of grain per average secondary ear-bearing stem was affected by competition in binary mixture. The enhancement in standard height—a strongly competitive genotype— reached in binary mixture about 10% for this last characteristic in comparison with the value from the pure stand. Some morphological characteristics of plants being in relation with competition for grain yield per plant are discussed in these experiments.     

AFLP markers reveal high clonal diversity and extreme longevity in four key arctic-alpine species

We investigated clonal diversity, genet size structure and genet longevity in populations of four arctic‐alpine plants (Carex curvula, Dryas octopetala, Salix herbacea and Vaccinium uliginosum) to evaluate their persistence under past climatic oscillations and their potential resistance to future climate change. The size and number of genets were determined by an analysis of amplified fragment length polymorphisms and a standardized sampling design in several European arctic‐alpine populations, where these species are dominant in the vegetation. Genet age was estimated by dividing the size by the annual horizontal size increment from in situ growth measurements. Clonal diversity was generally high but differed among species, and the frequency distribution of genet size was strongly left‐skewed. The largest C. curvula genet had an estimated minimum age of c. 4100 years and a maximum age of c. 5000 years, although 84.8% of the genets in this species were <200 years old. The oldest genets of D. octopetala, S. herbacea and V. uliginosum were found to be at least 500, 450 and 1400 years old, respectively. These results indicate that individuals in the studied populations have survived pronounced climatic oscillations, including the Little Ice Age and the postindustrial warming. The presence of genets in all size classes and the dominance of presumably young individuals suggest repeated recruitment over time, a precondition for adaptation to changing environmental conditions. Together, persistence and continuous genet turnover may ensure maximum ecosystem resilience. Thus, our results indicate that long‐lived clonal plants in arctic‐alpine ecosystems can persist, despite considerable climatic change.     

Resource heterogeneity, soil fertility, and species diversity: effects of clonal species on plant communities

Spatial heterogeneity in soil resources is widely thought to promote plant species coexistence, and this mechanism figures prominently in resource-ratio models of competition. However, most experimental studies have found that nutrient enhancements depress diversity regardless of whether nutrients are uniformly or heterogeneously applied. This mismatch between theory and empirical pattern is potentially due to an interaction between plant size and the scale of resource heterogeneity. Clonal plants that spread vegetatively via rhizomes or stolons can grow large and may integrate across resource patches, thus reducing the positive effect of small-scale resource heterogeneity on plant species richness. Many rhizomatous clonal species respond strongly to increased soil fertility, and they have been hypothesized to drive the descending arm of the hump-shaped productivity-diversity relationship in grasslands. We tested whether clonals reduce species richness in a grassland community by manipulating nutrient heterogeneity, soil fertility, and the presence of rhizomatous clonal species in a 6-year field experiment. We found strong and consistent negative effects of clonals on species richness. These effects were greatest at high fertility and when soil resources were applied at a scale at which rhizomatous clonals could integrate across resource patches. Thus, we find support for the hypothesis that plant size and resource heterogeneity interact to determine species diversity.