GEOMETRIC MORPHOMETRICS OF DEVELOPMENTAL INSTABILITY: ANALYZING PATTERNS OF FLUCTUATING ASYMMETRY WITH PROCRUSTES METHODS

Although fluctuating asymmetry has become popular as a measure of developmental instability, few studies have examined its developmental basis. We propose an approach to investigate the role of development for morphological asymmetry by means of morphometric methods. Our approach combines geometric morphometrics with the two-way ANOVA customary for conventional analyses of fluctuating asymmetry and can discover localized features of shape variation by examining the patterns of covariance among landmarks. This approach extends the notion of form used in studies of fluctuating asymmetry from collections of distances between morphological landmarks to an explicitly geometric concept of shape characterized by the configuration of landmarks. We demonstrate this approach with a study of asymmetry in the wings of tsetse flies (Glossina palpalis gambiensis). The analysis revealed significant fluctuating and directional asymmetry for shape as well as ample shape variation among individuals and between the offspring of young and old females. The morphological landmarks differed markedly in their degree of variability but multivariate patterns of landmark covariation identified by principal component analysis were generally similar between fluctuating asymmetry (within-individual variability) and variation among individuals. Therefore there is no evidence that special developmental processes control fluctuating asymmetry. We relate some of the morphometric patterns to processes known to be involved in the development of fly wings.     

EXAMINATION OF TWO DIMENSIONAL SPATIAL PATTERN OF PERIPHYTIC DIATOMS USING AN ADHESIVE SURFICIAL PEEL TECHNIQUE1

A simple surficial peel technique using adhesive tape was developed for quantitative removal of haptobenthic diatom communities from topographically simple substrata. The method combines high removal efficiency with low peel distortion, permitting the use of spatial statistics to test whether populations are distributed in the peel randomly or form aggregated or uniform patterns. Using this technique, the microdistribution of Cocconeis placentula Ehr. on a smooth acrylic rod was examined. Using conventional nearest neighbor analyses, a clonal population of C. placentula. characterized by an indentation of the value margin, was significantly aggregated, whereas the overall C. placentula population was uniform or aggregated depending on whether the method of analysis allowed for cell size. Using refined nearest neighbor analysis, the indented population was aggregated, and the overall population was random at distances greater than cell size. The results suggest that the indented clone was weakly motile following cell division and that its directional bearing was random.     

自然条件下风箱果的克隆构型

在自然条件下,风箱果(Physocarpus amurensis)主要靠克隆繁殖维持种群。植物克隆构型的可塑性变化对于其适应环境异质性具有重要意义。为探求风箱果的克隆构型及根茎生长动态,研究了其地下根茎的构筑型、形态特征、根茎的直径随长度变化的规律和地下根茎间的夹角。结果表明:风箱果的地下根茎的构筑型基本上属于游击型;风箱果无性系平均含有(6±2)个分株和(9.33±3.48)个根茎;根茎的直径随长度变化的曲线为抛物线型;分枝夹角较为稳定,多为30°和70°。风箱果生产大量的根茎系统,每一个克隆片段能够占据一定的空间,以保证自身生存和维持种群繁衍。     

THE ONTOGENY OF ASYMMETRY IN EARWIG FORCEPS

Fluctuating asymmetry may play an important role in the evolution of naturally selected and secondary sexual traits. However, very little is known about how asymmetries arise or how organisms maintain symmetry during development. Here I propose three mutually exclusive patterns for the development of asymmetries through consecutive growth stages: (1) compensatory growth, in which growth of the shorter side is greatest at the following growth stage; (2) persistent growth, in which growth of the longer side is greatest at the following growth stage; and (3) uncorrelated growth in which growth of the following stage is unrelated to the asymmetry at the previous one. I followed the growth in the forceps of male earwigs through four successive instars. Dyar's rule was used as a null model of insect growth. In the molt from the second to third instar, asymmetries increased through uncorrelated growth and with the magnitude but not the sign expected from Dyar's rule. However, following this, at the molts between instars 3–4 and 4–5, compensatory growth maintained asymmetries at a lower level than expected from Dyar's rule. Although there was no reduction in the absolute magnitude of asymmetry, relative asymmetry did decline. The net growth of forceps length did not follow Dyar's rule. The interpretation of patterns of growth were more sensitive and informative than the interpretation of the relations between asymmetries at consecutive instars.     

CLONAL INTEGRATION: NUTRIENT SHARING BETWEEN SISTER RAMETS OF SOLIDAGO ALTISSIMA (COMPOSITAE)

To test whether sharing of resources occurs among connected ramets of the tall goldenrod, Solidago altissima, we examined the extent of clonal integration for nutrients. In a greenhouse experiment, two-ramet clones were grown in a triad of connected pots so that nutrients could be supplied to either sister ramet or to their old rhizome (mother rhizome). Mother rhizomes and their associated roots shared nutrients with daughter ramets; however, any nutrient sharing that occurred between sister ramets was too little to significantly affect their growth. In addition, sister ramets not only competed for nutrients through parental connections, but larger ramets inhibited the growth of smaller ramets. We suggest that, for tall goldenrod, a clonal growth strategy in which nutrients are not shared among sister ramets may increase genet fitness by reducing the rhizome production of ramets in poor-nutrient microsites. Consequently, the genet would produce relatively fewer ramets in unfertile areas and make better use of heterogeneous nutrient resources.     

The effects of lack of normality and homogeneity of variance on analysis of variance and clonal heritability in <Emphasis Type="Italic">in vitro</Emphasis> clonal propagation traits

We studied the fulfilment of assumptions of normality and homogeneity of error variance, prior to application of analysis of variance (ANOVA), for in vitro clonal propagation data. We assessed the use of data transformations and mean values for situations when the original data did not satisfy the required assumptions. The purpose of the study was to establish whether the use of original, transformed or mean values had any effect on F values, significance levels and clonal heritability values. The F values, significance levels and values of clonal heritability obtained showed analysis of variance to be reliable, despite deviations with respect to normality and homogeneity of variance and despite the fact that samples sizes were unequal. Original data may be used for ANOVA applied to measured variables such as number of shoots per explant, length of tallest shoot, number of 1-cm segments per explant and also derived variables such as the multiplication coefficient. Frequency data can be used for analysis of variance of categorical-type variables such as apical necrosis and percentage of responsive explant. For shoot colour variables, the distributions were very skewed and the variances were very different, but even though the sample sizes were not identical in all cases, lack of homogeneity of variance did not significantly affect F values, significance levels or clonal heritability values, and thus analysis of variance may be applied to the original data. The use of original and frequency data makes interpretation of the results easier than when transformed data are used and also allows us to calculate variance components more accurately than when using mean values, which do not provide as much information. Clonal heritability values from transformed data and mean values showed differences of less than one hundredth compared with those from original data. Box–Cox-transformed data showed slightly lower heritability values than those corresponding to original data, whereas clonal heritability values from both mean data and angular-transformed data were slightly higher than those obtained using original data. In clonal variability studies with single growth medium, nutritional conditions that encouraged highly unequal growth or characteristics among clones gave rise to data that were unlikely to satisfy the conditions of normality or homogeneity of variance.     

ANALYSIS OF GENOTYPIC DIFFERENCES IN DEVELOPMENTAL STABILITY IN ANNONA CHERIMOLA

The genetic basis of developmental stability, measured as asymmetry (fluctuating asymmetry in leaves), was analyzed in leaves and flowers of cherimoya (Annona cherimola Mill) and atemoya (A. cherimola × A. squamosa). The individuals analyzed belonged to a controlled collection of cultivars (clones) that had previously been characterized by means of isozymes. We used a nested design to analyze the differences in asymmetry at several sampling levels: individual leaves and flowers, individual trees, and genotypes. The clonal repeatability of developmental stability was not significantly different from zero, thus suggesting the absence of heritability of the asymmetry for leaves and flowers under these environmental conditions. No relationship between asymmetry and individual heterozygosity was found, but leaf fluctuating asymmetry was significantly related to particular isozymic genes. Petal and leaf size showed a phenotypically plastic response to the exposure zone of the tree (mainly due to light). Leaf fluctuating asymmetry also showed such a plastic response. No significant correlation was found between asymmetry and any pomological characters (some of these being fitness related). Finally, the hybrid species (atemoya) did not show larger developmental instability than did the parental species (cherimoya). All these data show that cherimoya asymmetry reveals the random nature of developmental noise, with developmental stability for leaves being possibly related to specific chromosome regions, but with weak evidence for genotypic differences in developmental stability.     

WATER SHARING AMONG RAMETS IN A DESERT POPULATION OF DISTICHLIS SPICATA (POACEAE)

Clonal growth enables plants to transport resources among separately rooted but connected ramets, a potential advantage in patchy or unpredictable habitats. Nevertheless, clonal plants are relatively scarce in deserts. To test whether clonal integration of water relations can increase plant performance under natural conditions in a desert species, water movement was traced and connection among ramets was manipulated in the rhizomatous grass Distichlis spicata in Death Valley, California. To examine potential costs of clonal growth form, connections were mapped and analyzed for dry mass and nitrogen content. Movement of dye showed potential transport of water among five ramets up to 1.4 m apart. Severance of connecting rhizomes increased mortality and decreased water potential of individual ramets within 36 hr, indicating that water sharing among ramets could be of significant benefit. However, plants had a high investment of mass and nitrogen in underground organs, which might be a cost of clonal growth associated with desert environments.     

A STATISTICAL CHARACTERIZATION OF GROWTH AMONG CLONES OF SYNURA PETERSENII (SYNUROPHYCEAE)1

Each of four clones from the Synura petersenii complex was grown at different pHs (5.5, 6.5, 7.5, 8.5) in batch culture experiments. Growth response curves and exponential growth rates were compared among clones and pH treatments in order to examine growth trend variation among the clonal groups. The clones were isolated from geographically distant North American localities. The clonal groups represented distinct mating types, an isolate and its subisolate, and S. petersenii- and S. glabra-like scale morphologies. No consistent relationship existed between growth response curve, and culture medium pH. Additionally, the trends across time differed according to clone and pH combination. Pairwise comparisons of linear trends from transformed growth response curves indicated two distinct clonal associations. Although the clonal associations corresponded with the final cell density of the cultures, growth response curves did not correspond with mating type, the parent-isolate and subisolate, or scale morphology. Clones with glabra-like scales had greater growth rates than the clone with petersenii-like scales. The conflicting results generated from growth response curve and growth rate analyses support the concept that S. petersenii and S. glabra form a highly variable, homogeneous grouping.     

Regeneration growth of the invasive clonal forb Rorippa austriaca (Brassicaceae) in relation to fertilization and interspecific competition

A special type of clonal growth, spread by lateral roots, ishypothesized to be a favourable trait of invasive, opportunistic plant speciesof disturbed habitats. We tested this hypothesis for the invasive forbRorippa austriaca (Brassicaceae). Regenerationfrom root fragments, subsequent vegetative spread and allocation patterns inrelation to varied nutrient supply and intensity and pattern of interspecificcompetition were analyzed in container experiments. Regeneration success fromroot fragments was 100% and clonal spread was rapid but vegetativeperformance was strongly reduced under unfertilized conditions and,particularly, when interspecific competition was present. While the ratio ofabove- to belowground bio-mass did not differ considerably betweentreatments, R. austriaca allocated a high amount ofresources to belowground growth resulting in low aboveground but highbelowground biomass at harvest time relative to the matrix vegetation.Differences in shoot number or biomass between simulated gaps and denselyvegetated quadrants in the containers were (relatively) weak.Reproductive effort was less reduced under low resource levels, and the clonesdid not set seed at all, irrespective of the treatment. Our results show thatclonal growth by lateral roots and plasticity in clonal growth patterns inR. austriaca promote both exploitation of gaps andnutrient-rich microsites and resistance to competitors. Such plasticity,combined with its ability to regenerate from widely-dispersed rootfragments, contribute to the ability of the species to invade and persistwithindisturbed and spatially heterogeneous habitats.     

Demographic vulnerability of the clonal and endangered meadow thistle

For effective management of endangered species it is pivotal to understand why a species is endangered and which key life cycle components are involved in its response to environmental changes. Our objective was to investigate the response of rosettes of the redlisted clonal herb Cirsium dissectum to anthropogenic nutrient enrichment, which threatens its populations, and the consequences of these responses for its population dynamics. We constructed matrix population models with demographic data from three populations and four annual transitions and we decomposed the spatiotemporal variation in projected population growth rates into contributions from life cycle components. These patterns were compared with below-ground rosette dynamics in different fields, and with the below- and above-ground rosette dynamics in a garden experiment with nutrient enrichment and competing grasses. The decomposition analysis revealed that increased clonal rosette formation and decreased rosette survival were driving the spatial variation in the population growth rate. Excavating the below-ground rhizome network revealed a higher rosette turn-over in experimentally fertilized garden plots, which not only resulted in increased plot-level extinction, but also in increased spread of the clonal offspring. This supported the observed trend among field populations: rosette formation trades off with rosette survival. Surviving seedlings were only found in areas where the topsoil had been removed. The endangered C. dissectum is vulnerable when its habitat becomes more productive, because this species does not have the necessary capability to build up biomass. Small-scale disturbances such as created by sod-cutting or trampling cattle are essential for seedling establishment and necessary to render the explorative strategy of rhizomatous clonal spread successful.     

Growth of clonal modules on <i>Agropyron michnoi</i> in the Songnen Plain of Northeast China

Spatial expansion of clonal plants and growth of their modules are of concern in the field of plant ecology. After measuring a large number of samples, we analyzed the module components and the growth patterns of vegetatively propagated Agropyron michnoi clones in the Songnen Plain on Northeast China. The results showed that the plasticity of clonal growth was large; the coefficients of variation of both extensive areas and the quantitative characters of modules were more than 20%. The numbers of ramets, seedlings, and buds and the cumulative length of the rhizomes showed exponentially and linearly increasing patterns with increases of the area and the total number of modules. The biomass of each module, total number of modules and total biomass showed an allometric growth pattern, which was best described by power functions. For A. michnoi, there was a relatively stable investment to sexual reproduction; it showed a priority for allocating biomass to reproductive ramets, and also to rhizomes and buds formation.     

ACID EFFLUX PATTERNS OF PRIMARY AND LATERAL ROOTS OF PHASEOLUS VULGARIS (FABACEAE)

Graviresponding primary roots of Phaseolus vulgaris were characterized by more acid efflux on the upper (i.e., rapidly growing) side of the root than on the lower side of the root. Acid efflux patterns of the upper and lower sides of horizontally-oriented lateral roots were symmetrical. Addition of sodium orthovanadate (an inhibitor of auxin-induced H⁺ efflux) to the growth medium abolished gravicurvature and development of acid efflux asymmetry in horizontally-oriented roots. These results 1) support the suggestion that auxin redistribution may cause the asymmetry of acid efflux that mediates gravitropism, and 2) indicate that the lack of an auxin-induced asymmetry of acid efflux may be involved in explaining the minimal graviresponsiveness of lateral roots.     

Shoot Growth and Mortality Patterns of Urtica dioica, a Clonal Forb

The growth and mortality patterns of the clonal forb Urticadioica were investigated at the level of the individual shootin two growing seasons, 1991 and 1992, in a natural stand. Shootheight and diameter at ground level of each shoot tagged inspring were measured repeatedly five times during the growingseason. Dry weights of these repeatedly measured shoots wereestimated using an allometric relationship between dry weight,height and diameter of harvested shoots. A large decrease inshoot density occurred with stand development from the beginningof the growing season in both the years: (1) shoot survivalrate was about 30% at the end of the growing season; (2) shootmortality rate per 10 x 10 cm subplot between censuses was positivelydependent on shoot density per subplot; (3) the mortality rateof individual shoots was negatively dependent on shoot size(height, diameter and weight) at each growing stage, suggestingone-sided competition between living and dying shoots; (4) shootsize (height, diameter and weight) variability in terms of thecoefficient of variation and skewness decreased in accordancewith shoot mortality. Symmetric competition between living shootswas detected by regression analysis based on a model for individualshoot growth considering the degree of competitive asymmetry.However, the competitive effect on individual shoot growth wasvery small (nearly absent). The mortality pattern of Urticadioica indicates that shoot self-thinning occurred from theearly growing stage as in non-clonal crowded monospecific stands,and contrasts with many clonal plants where shoot self-thinningrarely occurs or, if any, is confined only to a short periodof the later growing stage. The pattern of growth and competitionbetween living shoots of Urtica dioica contrasts with non-clonalcrowded plants undergoing intense competition (usually asymmetric)between individuals, but is a common feature of many clonalplants where shoot competition is supposed to be reduced by'physiological integration' between shoots. These form a newpattern not reported yet for clonal plants. It is pointed outthat clonal plants show a wider spectrum of the growth, competitionand mortality patterns of shoots than non-clonals. Some possiblemechanisms for the pattern of Urtica dioica are discussed.Copyright1995, 1999 Academic Press Shoot competition, diffusion model, individual shoot growth, shoot self-thinning, shoot size variability, Urtica dioica L     

GENETIC AND MATERNAL EFFECTS ON SEEDLING CHARACTERS OF SOLIDAGO ALTISSIMA (COMPOSITAE)

The relative contributions of genetic and maternal factors to phenotypic variation were studied in seedlings of the clonal herb Solidago altissima L. Pairwise crosses were made to establish 19 full-sibships. Because crosses were reciprocal, each parent served as seed parent to half its offspring and as pollen parent to the other half. For 18 characters, genetic variation was estimated from the between sibship variance and maternal variation from the variance between maternal groups within sibships. Early in the growing season, maternal effects exerted strong influence over variation in leaf and stem dimensions, while genetic influences were weak or absent. At the end of the season genetic influences on leaf and stem dimensions, relative biomass allocation between stem and leaves, and the propensity to flower were significant but maternal effects were not. However, both genetic and maternal influences were found on the number and lengths of new rhizomes. Phenotypic, genetic, and maternal correlation matrices were examined through principal component analysis for trade-offs among vegetative growth, clonal expansion and floral reproduction which could constrain the evolution of life history patterns. At the phenotypic level, seedlings with strong vegetative growth produced more and longer rhizomes, and were more likely to flower. Genetically, correlations among these functions were weak or absent. Strong maternal contributions to seedling vegetative growth correlated positively with the number and length of rhizomes. Thus, no trade-offs acting during the seedling stage were found. However, there were indications of a genetically influenced trade-off between the number of rhizomes produced in the seedling year with the size of the ramets produced in the second year.     

Initial genetic diversity enhances population establishment and alters genetic structuring of a newly established Daphnia metapopulation

When newly created habitats are initially colonized by genotypes with rapid population growth rates, later arriving colonists may be prevented from establishing. Although these priority effects have been documented in multiple systems, their duration may be influenced by the diversity of the founding population. We conducted a large‐scale field manipulation to investigate how initial clonal diversity influences temporal and landscape patterns of genetic structure in a developing metapopulation. Six genotypes of obligately asexual Daphnia pulex were stocked alone (no clonal diversity) or in combination (‘high’ clonal diversity) into newly created experimental woodland ponds. We also measured the population growth rate of all clones in the laboratory when raised on higher‐quality and lower‐quality resources. Our predictions were that in the 3 years following stocking, clonally diverse populations would be more likely to persist than nonclonally diverse populations and exhibit evidence for persistent founder effects. We expected that faster growing clones would be found in more pools and comprise a greater proportion of individuals genotyped from the landscape. Genetic composition, both locally and regionally, changed significantly following stocking. Six of 27 populations exhibited evidence for persistent founder effects, and populations stocked with ‘high’ clonal diversity were more likely to exhibit these effects than nonclonally diverse populations. Performance in the laboratory was not predictive of clonal persistence or overall dominance in the field. Hence, we conclude that although laboratory estimates of fitness did not fully explain metapopulation genetic structure, initial clonal diversity did enhance D. pulex population establishment and persistence in this system.     

Using DNA Methylation Patterns to Infer Tumor Ancestry

Background

Exactly how human tumors grow is uncertain because serial observations are impractical. One approach to reconstruct the histories of individual human cancers is to analyze the current genomic variation between its cells. The greater the variations, on average, the greater the time since the last clonal evolution cycle (“a molecular clock hypothesis”). Here we analyze passenger DNA methylation patterns from opposite sides of 12 primary human colorectal cancers (CRCs) to evaluate whether the variation (pairwise distances between epialleles) is consistent with a single clonal expansion after transformation.

Methodology/Principal Findings

Data from 12 primary CRCs are compared to epigenomic data simulated under a single clonal expansion for a variety of possible growth scenarios. We find that for many different growth rates, a single clonal expansion can explain the population variation in 11 out of 12 CRCs. In eight CRCs, the cells from different glands are all equally distantly related, and cells sampled from the same tumor half appear no more closely related than cells sampled from opposite tumor halves. In these tumors, growth appears consistent with a single “symmetric” clonal expansion. In three CRCs, the variation in epigenetic distances was different between sides, but this asymmetry could be explained by a single clonal expansion with one region of a tumor having undergone more cell division than the other. The variation in one CRC was complex and inconsistent with a simple single clonal expansion.

Conclusions

Rather than a series of clonal expansion after transformation, these results suggest that the epigenetic variation of present-day cancer cells in primary CRCs can almost always be explained by a single clonal expansion.     

SPATIAL DISTRIBUTION OF GENETIC INDIVIDUALS IN THICKETS OF ALNUS INCANA SSP. RUGOSA,A CLONAL SHRUB

Thickets of speckled alder (Alnus incana ssp. rugosa (Du Roi) Clausen) consist of numerous discrete clumps of stems. Presumably all stems in a single clump are part of a single genetic individual, but a genet could comprise more than one clump. Starch-gel electrophoresis was used to identify genetic individuals in four alder populations in central New York. A single genetic marker, a tetramer with three alleles, could discriminate five genotypes. Nearest neighbor analysis revealed that genotypes were distributed randomly. That is, the pattern of genotypes was statistically indistinguishable from a model where each clump is considered a unique individual and where clump genotypes are randomly distributed. Calculation of Morisita's index of dispersion confirmed that clumps of a single genotype were not aggregated. Although alder is capable of forming root suckers and offsets, lateral expansion of genets is apparently ineffective. Apparently, spatial distribution of genetic individuals within alder thickets is not influenced by clonal growth or by other factors acting to cause patterns in the genetic structure of plant populations.     

Patterns of Solidago altissima ramet growth and mortality: the role of below-ground ramet connections

Summary For the rhizomatous perennial, Solidago altissima, I identified clonal fragments in the field, mapped ramet spatial locations, and documented patterns of ramet recruitment, growth, and mortality. Parent ramet size influenced the size and number of daughter ramets produced, and small ramets had lower survivorship and fecundity than large ramets. Similarly, small rhizomes tended to develop into small ramets, and ramets that survived to produce daughter ramets had longer parent-daughter rhizome connections than ramets that did not survive. In addition, most ramets that died during the growing season were connected to (genetically identical) ramets that persisted. There were large size inequalities among rhizomes, ramets, and clonal fragments. Inequalities in the size of ramets increased during the early part of the growing season, then decreased at the end of the season; similar patterns were observed for the growth of clonal fragments. In both instances, the decrease in size inequality could be attributed to the mortality of small individuals (ramets or clonal fragments). I found little evidence that ramet size hierarchies were structured by intraspecific competition. For example, path analyses and randomization tests indicated that size variation among S. altissima ramets was influenced little by the size of their near neighbors (but was influenced by parent size and rhizome size). In addition, within-season variation for the relative size and growth rate of individual ramets led to poor correlations between early and final ramet size; this result suggests that there was no stable hierarchy of dominant and suppressed ramets. I discuss implications of my results for contrasting interpretations of clonal plant population dynamics.     

Tree growth and competition in a post-logging Quercus mongolica forest on Mt. Sobaek, South Korea

Secondary woodlands in South Korea cover most mountains from low to middle elevations. While general patterns of forest succession are well understood, little is known about mechanisms of stand recovery after disturbance. We examined the spatio-temporal variations in establishment, growth, size inequality, and mode of competition among trees in a 50-year-old post-logging Quercus mongolica-dominated stand. We further compared the growth and stem allometry of single trees, presumably of seed origin, with multi-stemmed trees resprouting from stumps. Q. mongolica formed the upper canopy 16–22 m tall, 88.3% of total stand basal area, and 36.2% of total stem density, with most trees established during the first post-logging decade (51.2% were resprouts). During the first three decades, the Q. mongolica recruits grew exponentially, and disproportionately more in diameter than few older reserved trees left after the last cutting. This substantially decreased size inequality. The reverse trend was observed from 1994 to 2004: larger trees grow more, indicating an increasing asymmetry of competition for light. Neighborhood analysis revealed that when target trees had more or larger neighbors, their exponential phase of growth was reduced and maximum size was decreased. After the 50 years of stand development, more than 70% of Q. mongolica showed growth decline as a result of competitive stress, and mortality was about 30%, concentrated in smaller size classes. Compared to single stems, resprouts within clones do not seem to compete less asymmetric as might be expected based on studies of clonal herbaceous plants and physiological integration within genets. As Q. mongolica was also negatively affected by competition from woody species currently prevailing in the lower tree stratum (Tilia amurensis, Acer mono, Fraxinus rhynchophylla, Acer pseudosieboldianum), we predict the stand will become increasingly dominated by these more shade-tolerant trees.