Estimation of Developmental Stability of Small-Leaved Lime on Reserved and Urbanized Territories

The developmental stability of small-leaved lime (Tina cordata Mill.) was estimated in populations on territories of the Kerzhenskii State Nature Reserve, Nizhni Novgorod District, and in an industrial region of Nizhni Novgorod. The developmental stability was estimated according to the fluctuating asymmetry of the leaf measurements. The results obtained suggest that, in the urbanized territory, the homeostatic developmental mechanisms are weakened, as expressed in the increased degree of leaf asymmetry.     

Peculiarities of Leukocytic Formula of Peripheral Blood of Green Frogs under Conditions of Anthropogenic Load

Changes of the total number of leucocytes and the leucocytic formula of peripheral blood were studied and stability of development was evaluated by the indicator of fluctuating asymmetry of populations of green frogs Rana lessonae Cam. and Rana ridibunda Pall. inhabiting city reservoirs of Nizhni Novgorod and the Nizhegorodskaya Province. The revealed changes in the amphibians white blood depending on the degree of anthropogenic effects in the studied reservoirs can be divided into two types. In the first type (the level of stability of development of populations is 2 scores according to Zakharov) there were revealed neutropenia, increased content of eosinophils, and total leucopenia. In the second type, a more intensive anthropogenic effect and deviation of the organism state from normal conditions, estimated by 3 scores (Zakharov), the leucograms of frogs were characterized by more pronounced negative alterations. Leucocytosis accompanied by neutropenia, eosinophilia, basophilia, and a decrease of the total number of lymphocytes has been established. In the absence of deviations of the stability of development from the conditionally normal state, there was revealed the balance of leucogram indicators, close to norm. An increase of the number of lymphocytes and young forms of neutrophils in the amphibian peripheral blood indicates stimulation of the processes providing the organism protective reaction. Cross-correlation between the integral indicator of stability of development, the content of eosinophils and lymphocytes has been established in individuals from populations of green frogs inhabiting the studied reservoirs.     

Developmental stability of a leaf of Pisum sativum L. under the influence of formaldehyde in a wide range of doses

The influence of formaldehyde in a wide range of doses on the stability of development of the third leaf of pea (Pisum sativum L.) was studied. The developmental stability of the leaf was assessed by the change in the value of the directional asymmetry of the right and left leaflets caused by the fluctuating asymmetry of these morphological structures. When subjected to a toxic agent, the studied parameter exhibited a paradoxical effect. In minimum studied concentrations, formaldehyde disturbed stability of leaf development, which was manifested in an increase in the asymmetry of the right and left leaflets. At medium concentrations of the toxicant, the asymmetry was less than the control level, which indicated an increase in the developmental stability of the pea leaf. Maximum studied concentrations of formaldehyde, close to sublethal, again reduced the stability of development of the pea leaf and led to an increase in the asymmetry of its leaflets compared with the controls.     

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.     

The effect of nutrient stress on developmental instability in leaves of Acer platanoides (Aceraceae) and Betula pendula (Betulaceae)

Studies of developmental stability can provide insights into the amount of genetic or environmental stress experienced by individuals or populations. In the present study, we used young plants of Acer platanoides (Norway maple) and Betula pendula (silver birch), two distantly related tree species with widely different leaf morphologies, to compare the expression of developmental instability in two contrasting environments: one with free access to nutrients and the other with a severely limited supply of nutrients. Using the difference in size between the right and left side of each leaf as a measure of developmental instability, we found no effect of nutrient deficiency on leaf asymmetry, despite large sample sizes (370-380 plants per species and treatment) and evidence for stress-related changes in overall leaf size and plant biomass. Moreover, there was no consistent relationship between individual leaf asymmetry and plant biomass within each nutrient treatment. In view of these observations, leaf asymmetry appears to be a poor indicator of nutrient stress in young plants of Acer platanoides and Betula pendula.     

A meta-analysis of the heritability of developmental stability

The existence of additive genetic variance in developmental stability has important implications for our understanding of morphological variation. The heritability of individual fluctuating asymmetry and other measures of developmental stability have frequently been estimated from parent-offspring regressions, sib analyses, or from selection experiments. Here we review by meta-analysis published estimates of the heritability of developmental stability, mainly the degree of individual fluctuating asymmetry in morphological characters. The overall mean effect size of heritabilities of individual fluctuating asymmetry was 0.19 from 34 studies of 17 species differing highly significantly from zero (P < 0.0001). The mean heritability for 14 species was 0.27. This indicates that there is a significant additive genetic component to developmental stability. Effect size was larger for selection experiments than for studies based on parent-offspring regression or sib analyses, implying that genetic estimates were unbiased by maternal or common environment effects. Additive genetic coefficients of variation for individual fluctuating asymmetry were considerably higher than those for character size per se. Developmental stability may be significantly heritable either because of strong directional selection, or fluctuating selection regimes which prevent populations from achieving a high degree of developmental stability to current environmental and genetic conditions.     

Patterns of leaf asymmetry changes in Plantago major L. (ssp. major) natural populations exposed to different environmental conditions

In the present study, developmental stability of leaf traits was examined in three natural populations of Plantago major L. (ssp. major), representing two polluted environments (Karaburma and Zemun) and an unpolluted area (Crni Lug). Developmental stability was assessed as fluctuating asymmetry (FA). The magnitude of FA is believed to reflect differences in the ability of individuals to buffer their development in natural populations. We hypothesized that there are differences within characters and among characters in response to environmental conditions. Significant patterns of asymmetry correlations and asymmetry changes were detected both within characters and between characters. The manova results revealed a significant effect of individual and a significant individual × environment interaction on actual asymmetry (logL_i ? logR_i) and on the amount of asymmetry |(logL_i ? logR_i)| for leaf width and vein distances within a leaf. Over time, statistically significant and positive correlations of the FA values were detected for each trait separately per sample (population). For both leaf traits, there were differences for (logL_i ? logR_i) and |(logL_i ? logR_i)| asymmetry values among individuals within samples in response to yearly variations. Statistically significant and negative correlations for (logL_i ? logR_i) versus |(logL_i ? logR_i)| asymmetries were detected for both leaf traits. In summary, our results highlight the importance of differences in the ability of individuals to buffer their development under different environmental conditions and point to the concept that developmental stability is character specific.     

Leaf developmental stability of Platanus acerifolia under urban environmental stress and its implication as an environmental indicator

The developmental stability indices,leaf width based fluctuating asymmetry (FA1),and lateral vein length based directional asymmetry (RDA1) of Platanus acerifolia were studied.All the leaves were sampled from 14 sites that were categorized based on different urban environmental stress levels (UESL) in Shanghai metropolitan,China.Besides,foliar stomatal density and stomatal length were also studied as the subsidiary indices to test the availability of developmental stability indices as the indicator under a stressful environment.Results showed seasonal variation of FA1 and RDA1 existed among the 14 sites,but the data showed significant negative correlation between FA1 and UESL (FA1=0.029-0.000 9UESL+0.000 3UESL2,r=0.766 5,P=0.001 4).However,a similar trend was not found between RDA1 and UESL.Furthermore,the significant correlation among FA1 and leaf stomatal length and stomatal density implied they could be used as indicators of urban stress levels on a small scale.It seemed that RDA1 was possibly a normal parameter during leaf development but it was unavailable for use as an indicator of urban stresses.     

The repeatability of fluctuating asymmetry: a revision and extension

Fluctuating asymmetry (FA) is an imperfect measure of the developmental stability of an individual. Estimates of the heritability of FA will underestimate the heritability of developmental stability; the extent of this bias can be estimated and corrected by estimating the repeatability of developmental stability. This note corrects an error in a previous derivation of this repeatability using the absolute value of the difference between sides as the measure of FA, and derives the repeatability of developmental stability for another common measure of FA, the variance among sides within an individual.     

DEVELOPMENTAL STABILITY IN LEAVES OF CLARKIA TEMBLORIENSIS (ONAGRACEAE) AS RELATED TO POPULATION OUTCROSSING RATES AND HETEROZYGOSITY

Four natural populations of Clarkia tembloriensis, whose levels of heterozygosity and rates of outcrossing were previously found to be correlated, are examined for developmental instability in their leaves. From the northern end of the species range, we compare a predominantly selfing population (t? = 0.26) with a more outcrossed population (t? = 0.84), which is genetically similar. From the southern end of the range, we compare a highly selfing population (t? = 0.03) with a more outcrossed population (t? = 0.58). We measured developmental stability in the populations using two measures of within-plant variation in leaf length as well as calculations of fluctuating asymmetry (FA) for several leaf traits. Growth-chamber experiments show that selfing populations are significantly more variable in leaf length than more outcrossed populations. Developmental instability can contribute to this difference in population-level variance. Plants from more homozygous populations tend to have greater within-plant variance over developmentally comparable nodes than plants from more heterozygous populations, but the difference is not significant. At the upper nodes of the plant, mature leaf length declines steadily with plant age, allowing for a regression of leaf length on node. On average, the plants from more homozygous populations showed higher variance about the regression (MSE) and lower R² values, suggesting that the decline in leaf length with plant age is less stable in plants from selfing populations than in plants from outcrossing populations. Fluctuating asymmetry (FA) was calculated for four traits within single leaves at up to five nodes per plant. At the early nodes of the plant where leaf arrangement is opposite, FA was also calculated for the same traits between opposite leaves at a node. Fluctuating asymmetry is significantly greater in the southern selfing population than in the neighboring outcrossed population. Northern populations do not differ in FA. Fluctuating asymmetry can vary significantly between nodes. The FA values of different leaf traits were not correlated. We show that developmental stability can be measured in plants using FA and within-plant variance. Our data suggest that large differences in breeding system are associated with differences in stability, with more inbred populations being the least stable.     

城市胁迫生境下二球悬铃木叶片的发育稳定度及其环境指示功能

对采自上海市区14个样点的二球悬铃木叶片发育稳定度、叶片气孔密度和气孔长径进行研究。结果表明,在3个采样时期,根据叶片宽度确定的非稳态不对称指数(FA1)和根据叶片侧脉长度确定的偏向性不对称指数(RDA1)在不同环境胁迫程度(UESL)下均表现出显著的差异,但进一步分析表明,各样点FA1与环境胁迫程度UESL之间呈显著的非线性回归关系,而RDA1与环境胁迫程度之间却未显示类似关系。此外,FA1与叶片气孔密度和气孔长径之间的显著相关关系表明,它们可共同作为指示小尺度上环境胁迫的有效指标,而RDA1是叶片的一种正常发育状态参数,并不适于反映环境胁迫状态。     

Gene pool structure of Russian populations from the European part of Russia inferred from the data on Y chromosome haplogroups distribution

Population structure of Russian population from the European part of Russia was investigated by analyzing the distribution of 23 SNP makers of Y chromosome in Russian populations from Kaluga, Yaroslavl’, Vladimir, Nizhni Novgorod, Pskov, Tula, Belgorod, and Novgorod oblasts. In the populations studied a total of 14 Y-chromosome haplogroups (E, F*, I, J, K*, N3a, N2, P*, R1*, R1a1, C3, G, H, and A) were discovered, of which haplogroups R1a1, I, and N3a were the prevailing. Analysis of Φ statistics in the populations grouped in accordance to the dialect subdivision of the Russian language, showed the absence of statistically significant differences between Russian population groups. Analysis of the Y-chromosome markers distribution patterns among Russian population (10 population groups) in comparison with the population of Germany (11 population groups) and Poland (8 population groups) revealed statistically significant differences between the gene pools of Slavs (Russians and Poles) and Teutons (Germans).     

Effects of extreme temperatures on phenotypic variation and developmental stability in Drosophila melanogaster and Drosophila buzzatii

The effect of stressful and nonstressful rearing temperatures on phenotypic variation of four quantitative characters (thorax length, wing length, number of sternopleural chaetae, number of arista branches) and on developmental stability (fluctuating asymmetry) of the three latter characters was estimated in two Drosophila species: Drosophila melanogaster and Drosophila buzzatii . In both species, a general trend for increasing of phenotypic variation and fluctuating asymmetry at stress temperatures was observed; in fluctuating asymmetry, this effect was more pronounced. An increase of phenotypic variation under stress was shown for all characters examined except sternopleural chaeta number in D. buzzatii . Comparison of species responses suggests that the increase of variation in D. melanogaster was somewhat higher than in D. buzzatii .     

Changes in resilience to fire disturbance in lowland pine forest ecosystems

The effect of major disturbances on the structure of pine forests in the interfluve of the Chernaya and Pugai rivers has been retrospectively analyzed back to the late 19th century. The results show that intensive timber harvesting and fires produced a synergistic effect so that the natural fire regime in these forests has changed; in particular, the severity of fires and the frequency of stand-replacement fires have increased. The ecosystem of lowland pine forests, adapted to periodic fires, largely loses its resilience to fire disturbance if the age structure of pine populations becomes relatively homogeneous and communities at early succession stages gain significant prevalence in the vegetation. The main cause of the largest fire that occurred in pine forests of the Nizhni Novgorod Transvolga region (Nizhegorodskoe Zavolzhie) in 1972 was human activity rather than natural and climatic features of this territory.     

Use of a geometric morphometric method to determine the developmental stability of <Emphasis Type="Italic">Betula pendula</Emphasis> Roth

A geometric morphometric method has been proposed to test the fluctuating asymmetry and the developmental stability of Betula pendula Roth populations. The main factors affecting the developmental stability were the industrial emission level, the relief altitude, and the interaction of both these factors. A strong correlation between the fluctuating asymmetry indices obtained by the normalizing difference method and geometric morphometric method has been revealed. The fluctuating asymmetry determined by the geometric morphometric method is sensitive to the presence of directional asymmetry, which makes it possible to use this method for precise bioindication mapping of the developmental stability.     

Asymmetry of the a-b ridge count in low-back pain sufferers

Fluctuating asymmetry of paired morphological structures is regarded as a measure of developmental stability. To test whether poorly canalized individuals are highly sensitive to postnatal environmental influence, we accepted the exogenous disease low-back pain as an example of such sensitivity. Asymmetry of the palmar a-b ridge count was examined in 217 males suffering from low-back pain against 300 healthy controls. Low-back pain patients showed significantly higher values of asymmetry indices indicating lower developmental stability. The results suggest that dermatoglyphic asymmetry can mark phenotypes weakly adapted to postnatal stress.     

Tolerance assessment of Cistus ladanifer to serpentine soils by developmental stability analysis

Developmental instability is the result of random environmental perturbations during development. Its absence (developmental stability) depends on an organism's ability to buffer environmental disturbances. Both genotype and environment influence the phenotypic expression of developmental instability and it is susceptible to selection pressure. We studied developmental instability (as indicated by increased within-individual asymmetry of repeated traits) in vegetative and reproductive structures of three populations of Cistus ladanifer L. living in different soil substrates (serpentine, siliceous and contact zone) to detect tolerance to serpentine soils. Serpentine soils, characterized by high concentrations of heavy metals (Ni, Cr, and Co), low levels of Ca/Mg ratio and high water deficit, can adversely affect plant performance. In this study we demonstrated that asymmetry and within-plant variance were higher in the contact zone population than either the silica or serpentine populations, proving the adaptation of C. ladanifer to serpentine soils. Within-population estimates of developmental instability were concordant for both vegetative and reproductive traits. There was little or no within-individual correlation among estimates of developmental instability based on different structures, i.e., plants that had highly asymmetric leaves always had high developmental instability in translational symmetry. Radial asymmetry of petals was negatively correlated with petal size, especially in silica soil plants, providing evidence of selection for symmetric and large petals. While leaf size was positively correlated with absolute fluctuating asymmetry, suggesting selection for small or intermediate size leaves. Serpentine soils presented the largest foliar and floral traits, as well as shoot elongation, while silica soil plants had the smallest scores. On the contrary, aboveground plant biomass was larger in silica soil plants, while the contact zone plants had the lowest biomass.     

Fluctuating dental asymmetry: A measure of developmental instability in Down syndrome

Subjects with Down syndrome provide a useful model for investigating the effect of chromosomal aneuploidy on developmental pathways. Studies suggest that a major effect of trisomy is a decrease in developmental stability. The present study examines fluctuating dental asymmetry in Down syndrome. Mesiodistal crown diameters were measured from dental casts of 114 Down syndrome subjects. Correlation coefficients for antimeric permanent teeth served as an index of dental asymmetry. These values were compared with normal values obtained from the literature. Fluctuating dental asymmetry is thought to reflect the relative success of developmental homeostasis in countering developmental disturbances. Down syndrome subjects have significantly increased dental asymmetry. In addition, they show a disproportionate increase in dental asymmetry for those teeth reported to have the least developmental stability. These results support the contention that the chromosomal imbalance in Down syndrome results in amplified developmental instability.     

Variational and genetic properties of developmental stability in Dalechampia scandens

Because low developmental stability may compromise the precision with which adaptations can be reached, the variability and genetic basis of developmental stability are important evolutionary parameters. Developmental stability is also an important clue to understanding how traits are regulated to achieve their phenotypic target value. However, developmental stability must be studied indirectly through proxy variables, such as fluctuating asymmetry, that are suggested to have noisy and often nonlinear relationships to the underlying variable of interest. In this paper we first show that mean-standardized measures of variance and covariance in fluctuating asymmetry, unlike heritabilities, repeatabilities, and correlations, are linearly related to corresponding measures of variation in underlying developmental stability. We then examine the variational properties of developmental stability in a population of the Neotropical vine, Dalechampia scandens (Euphorbiaceae). By studying fluctuating asymmetry in a large number of floral characters in both selfed and outcrossed individuals in a diallel design, we assemble strong evidence that both additive genetic and individual variation and covariation in developmental stability are virtually absent in this population.     

Parasitic infection and developmental stability: fluctuating asymmetry in Gammarus pulex infected with two acanthocephalan species

Several studies have reported a negative association between developmental stability and parasitic infection. However, the host-parasite associations examined so far consist only of a limited number of parasite taxa, and developmental stability was appraised on definitive hosts. The present study examines the association between infection by 2 acanthocephalan parasites. Pomphorhynchus laevis and Polymorphus minutus, and the developmental stability of their common intermediate host Gammarus pulex. Developmental stability was estimated from the fluctuating asymmetry (FA) levels of 6 morphological traits. A positive association was found between FA and infection. Infected gammarids tended to be more asymmetrical than the noninfected ones for an index generated by combining FA scores from 2 characters out of the 6 studied, even though no significant relationships were found between FA levels and parasitic loads. The simultaneous presence of both acanthocephalan species in the same host seems to be associated with increased FA levels of gammarids, but this trend was not statistically significant. For the same characters, males exhibited higher levels of FA than females.