FLUCTUATING ASYMMETRY IN A SALIX HYBRID SYSTEM: THE IMPORTANCE OF GENETIC VERSUS ENVIRONMENTAL CAUSES

To examine the effects of hybridization and environmental stress on developmental instability, we examined fluctuating asymmetry (FA), the variance in random deviations from perfect symmetry in bilaterally symmetrical traits, for leaf symmetry in a Salix hybrid system. An abiotic environmental stress (water stress), an interspecific biotic stress (pathogen attack), and an intraspecific biotic stress (competition) were examined to determine which factors increase developmental instability. None of these three environmental stressors significantly increased FA. However, genetic stress through hybridization was detected; hybrid plants showed significantly higher levels of FA than parental species. In contrast to hybridization providing greater developmental stability through heterozygosity, these results suggest that complex, nonadditive interactions provided developmental stability and that developmental instability increased when coadapted gene complexes were disrupted through hybridization. In addition, plant biomass was significantly, negatively correlated with FA, suggesting that those individuals that were more able to buffer themselves against the disruptive effects of environmental stress may have a selective advantage over those that are less able to buffer themselves against these disruptive effects.     

BILATERAL DIRECTIONAL ASYMMETRY OF THE APPENDICULAR SKELETON OF THE HARBOR PORPOISE (PHOCOENA PHOCOENA)

Directional asymmetry ( DA ) of the lengths, diameters, and masses of the scapula, humerus, radius, and ulna were analyzed on a sample of 213 harbor porpoises from Denmark and West Greenland. The levels of DA were consistent across yearlings and older animals, mature and immature animals, sexes, and populations. All investigated variables showed significant DA favoring the right side. For the pooled sample, DA of lengths ranged from 0.2% SE 0.1 (ulna) to 1.2% SE 0.1 (scapula). DA s of diameters were 1.5% SE 0.2 for the humerus, 0.6% SE 0.1 for the radius, and 1.3% SE 0.2 for the ulna. DA of mass ranged from 2.8% SE 0.5 (humerus) to 4.3% SE 0.7 (ulna). The humerus and ulna had significantly larger mean diameter/ length ratios on the right side than the left, making them more robust. The large DA of scapula length indicates larger muscle mass associated with the right flipper, while the generally more robust right humerii and ulnae may be designed for higher levels of mechanical stress. These DA s and the examples of lateralized behavior recorded in cetaceans, point to the existence of lateralized use of the flippers at the population level in harbor porpoises and possibly other cetacean species.     

ESTIMATING THE HERITABILITY OF FLUCTUATING ASYMMETRY IN FIELD DROSOPHILA

Some studies have found intermediate heritabilities for fluctuating asymmetry (FA) in traits, but almost all of these are flawed and/or based on laboratory experiments. We therefore tested if there was heritable variation for FA in bristle and wing traits in three field collections of Drosophila melanogaster by rearing F₁s from field flies under laboratory conditions. One of the collections was reared to the F₂ generation in the laboratory to compare heritability estimates from the laboratory with those from the field-laboratory comparison. Trait means indicated an increase in size under laboratory rearing. FAs increased in one collection, decreased in another collection, and showed no changes in the third collection under laboratory rearing. FAs from the collections tended to converge under laboratory conditions. Morphological traits were heritable under field conditions. However, FA was not significantly heritable for any of the individual traits or when FA was determined by combining traits. Comparisons of the two laboratory generations showed that FA heritability was low under laboratory conditions, in contrast to the morphological traits themselves. These findings suggest a very low heritability for FA in field and laboratory Drosophila. FA in bristle and wing traits may therefore be a poor indicator of genetic quality in Drosophila.     

FLUCTUATING ASYMMETRY DOES NOT CONSISTENTLY REFLECT SEVERE DEVELOPMENTAL DISORDERS IN HUMAN FETUSES

Developmental instability (DI), as measured by fluctuating asymmetry (FA), may reflect fitness and facilitate the expression of morphological variation. Insights in the underlying mechanisms and magnitude of DI during early development would increase our understanding of its role in evolutionary biology. We studied associations between FA and congenital abnormalities of different origins and functional systems in deceased human fetuses. Major congenital abnormalities corresponded to severe, often-lethal developmental disorders disrupting normal development from early organogenesis onward, but only moderately increased FA. Lower FA with age also supported the hypothesis that more severe abnormalities, leading to an earlier death, increased DI. Although FA related significantly to measures of fitness or health, we anticipated stronger associations because fetal health problems were detrimental. Furthermore, elevated FA occurred in only 4 of 17 disorders (left–right patterning, limb defects, and problems of bronchopulmonary and urogenital system). Fetuses experiencing major abnormalities other than these four types did not show increased FA. This suggests that the functional importance of symmetry in limbs has resulted in strong selection for symmetry and reduced its sensitivity to stress. Finally, the observed patterns suggest that specific developmental pathways have a stronger effect on DI than others do.     

A SEARCH FOR QUANTITATIVE TRAIT LOCI AFFECTING ASYMMETRY OF MANDIBULAR CHARACTERS IN MICE

An interval mapping procedure was used to search for and describe the effects of any quantitative trait loci (QTLs) for directional asymmetry (DA) and fluctuating asymmetry (FA) of 10 bilateral mandible characters in house mice. It was hypothesized that more QTLs would be found for DA than for FA, but that any discovered for FA should tend to exhibit dominance. All mandible characters were triply measured and 76 microsatellite markers were scored in an average of 471 mice from the F₂ intercross of the Large (LG/J) and Small (SM/J) inbred strains. A total of 16 QTLs significantly affected DA in nine of the 10 mandible characters, and this was more than the 9.5 expected by chance alone. These QTLs were found on seven of the 19 chromosomes, often at or near locations of QTLs affecting the mean of the two sides for various dimensions on the mandible. It was concluded that there is genetical variability for DA in these characters, although its level was low (4.4% of the total variation in this particular F₂ population). Eleven QTLs were detected for FA, suggesting that there is very little genetic variability for FA, at least as seen in the mandible characters in this particular F₂ population. As hypothesized, however, these QTLs did tend to exhibit dominance.     

THE ASSOCIATION BETWEEN FLUCTUATING ASYMMETRY,TRAIT VARIABILITY,TRAIT HERITABILITY,AND STRESS: A MULTIPLY REPLICATED EXPERIMENT ON COMBINED STRESSES IN DROSOPHILA MELANOGASTER

A number of hypotheses have been proposed about the association between developmental stability phenotypic variability, heritability, and environmental stress. Stress is often considered to increase both the asymmetry and phenotypic variability of bilateral traits, although this may depend on trait heritability. Empirical studies of such associations often yield inconsistent results. This may reflect the diversity of traits and conditions used or a low repeatability of any associations. To test for repeatable associations between these variables, multiply replicated experiments were undertaken on Drosophila melanogaster using a combination stress at the egg, larval and adult stages of reduced protein, ethanol in the medium, and a cold shock. Both metric and meristic traits were measured and levels of heritable variation for each trait estimated by maximum likelihood and parent-offspring regression over three generations. Trait means were reduced by stress, whereas among-individual variation increased Fluctuating asymmetry (FA) was increased by stress in some cases, but few comparisons were significant. Only one trait orbital bristle, showed consistent increases in FA. Changes in trait means, trait phenotypic variability, and developmental stability as a result of stress were not correlated. Extreme phenotypes tended to have higher levels of FA but only the results for orbital bristles were significant. All traits had low to intermediate heritabilities except orbital bristle, which showed no heritable variation. Only traits with low heritability and high levels of phenotypic variability may show consistent increases in FA under stress. Overall, the independence of phenotypic variability, plasticity, and the developmental stability of traits extend to changes in these measures under stressful conditions.     

FLUCTUATING ASYMMETRY IN CENTRAL AND MARGINAL POPULATIONS OF LYCHNIS VISCARIA IN RELATION TO GENETIC AND ENVIRONMENTAL FACTORS

Developmental instability in the form of increased fluctuating asymmetry can be caused by either genetic or environmental stress. Because extinctions can be attributed broadly to these factors, fluctuating asymmetry may provide a sensitive tool for detecting such stresses. We studied the level of fluctuating asymmetry of flowers of a perennial outcrossing plant species, Lychnis viscaria, both in natural and common-garden populations. The degree of flower asymmetry was higher in small, isolated, and marginal populations of the species range. These marginal populations also were the most homozygous. In the core area of the species' range, flowers were more symmetrical The level of asymmetry was correlated with both population size and heterozygosity. However, a partial correlation analysis revealed that when the impact of population size was controlled for, there was a negative relationship between fluctuating asymmetry and heterozygosity, whereas when controlling for heterozygosity, no relationship between population size and fluctuating asymmetry was found. This indicates that genetic consequences of small population size probably underlie the relationship between the level of asymmetry and population size. Results from a transplantation experiment showed that individuals subjected to a higher environmental stress had an increased level of asymmetry compared to control plants. In the common-garden conditions the level of fluctuating asymmetry did not differ between the central and marginal populations. This suggests that presumably both genetic and environmental factors affected to the higher level of asymmetry among marginal populations compared to central ones. In all we conclude that even though fluctuating asymmetry seems to be a sensitive tool for detecting stresses, results from studies focusing on only one factor should be interpreted with caution.     

GENETICS OF FLUCTUATING ASYMMETRY: A DEVELOPMENTAL MODEL OF DEVELOPMENTAL INSTABILITY

Although numerous studies have found that fluctuating asymmetry (FA) can have a heritable component, the genetic and developmental basis of FA is poorly understood. We used a developmental model of a trait, according to a diffusion-threshold process, whose parameters are under genetic control. We added a small amount of random variation to the parameter values of this model to simulate developmental noise. As a result of the nonlinearity of the model, different genotypes differed in their sensitivity to developmental noise, even though the noise is completely random and independent of the genotype. The heritable component of FA can thus be understood as genetically modulated expression of variation that is itself entirely nongenetic. The loci responsible for this genetic variation of FA are the same that affect the left/right mean of the trait, showing that genetic variation for FA does not require genes that specifically control FA. Furthermore, the model offers alternative explanations for phenomena widely discussed in the literature on FA, for instance, the correlations between FA and heterozygosity and between FA and trait size. The model underscores the importance of dominance and epistasis, and therefore unites the study of FA with the classical theory of quantitative genetics.     

THE GENETIC BASIS OF DEVELOPMENTAL STABILITY IN APIS MELLIFERA: HETEROZYGOSITY VERSUS GENIC BALANCE

The genetic basis for developmental stability in the haplo-diploid honeybee Apis mellifera was determined by comparing the level of asymmetry between diploid females and haploid males both among and within inbreeding levels. There was no significant relationship between the level of inbreeding and the level of fluctuating asymmetry for both females and males. It is therefore argued that the general level of genomic heterozygosity is not an important factor for the determination and maintenance of developmental stability in this system, but rather that the balance of genes within chromosomes plays the major role. The observation that males were generally more asymmetric than females suggests that developmental stability in females may also be influenced by additional factors such as gene dosage, sex-limited genes or cytoplasmic elements.     

西北干旱区10种荒漠植物地理分布与大气候的关系及其可能潜在分布区的估测

 选择面积广阔、地理位置特殊、气候干旱、地形地貌多样、生态系统脆弱以及人类活动长期干扰,而且目前面临着西部大开发和环境保护双重矛盾的西北干旱区作为研究区域,通过收集西北干旱区多种优势种和常见种的地理分布资料,选取10 种分布范围相对明确、资料相对齐备的荒漠植物种：短叶假木贼（Anabasis brevifolia）、木蓼（Atraphaxis frutescens）、沙拐枣（Calligonum mongolicum）、膜果麻黄（Ephedra przewalskii）、裸果木（Gymnocarpos przewalskii）、梭梭柴（Haloxylon ammodendron）、白梭梭（Haloxylon persicum）、尖叶盐爪爪（Kalidium cuspidatum）、松叶猪毛菜（Salsola laricifolia）和合头草（Sympegma regelii）,定量分析其地理分布与气候因子的关系,并据此估测其潜在中心分布区和潜在最大分布范围,与实际分布范围进行比较。结果表明,10种荒漠植物的Holdridge生物温度、降水量和可能蒸散率的平均值、标准差、最大值和最小值比较准确地反映了我国荒漠地区的典型气候特征,说明植物种分布与气候的定量对应关系较好。其可能潜在分布的估测中,中心分布估测图与植物种的实际分布范围有较好的对应性,而在最大可能分布的估测图中,与实际有一定误差及存在一些不合理的地方,但种与种之间有差异。究其原因,首先在于目前所拥有的植物种分布和气象资料的限制,而且植物种的实际分布范围受到人为活动的影响较大,所以无法完全把握植物种与气候的准确定量关系;其次,气候极值所预测的只是植物种分布的最大可能性,而不能表现出植物种现实分布的主要地区,气候均值与标准差所反映的中心分布区,其准确性更可靠;再者,模拟预测仅考虑了植物种与大气候的关系,而没有考虑决定植物分布的其它环境因素,如土壤、基质、地形等地理因素。     

HANDEDNESS AND ASYMMETRY IN SCALE‐EATING CICHLIDS: ANTISYMMETRIES OF DIFFERENT STRENGTH

Individual symmetry is believed to be advantageous and reflecting developmental stability, but frequency‐dependent selection can also maintain polymorphisms of asymmetric phenotypes. There are many examples of so‐called antisymmetry, where mirror image morphs occur at equal frequencies. With very few exceptions, these are caused by nongenetic variation. One notable exception is handedness and mouth bending variation in the scale‐eating cichlid Perissodus microlepis, which has been suggested to be an example of antisymmetry determined by a single genetic locus of large effect. Here, we report that this handedness and mouth bending asymmetry are not jointly and exclusively determined by a single major locus. We found no evidence of a major locus for asymmetry and some support for a major handedness locus. Also, asymmetry is plastic in this species: it can change in adults. We suggest that behavioral handedness in this system precedes and guides morphological asymmetry.     

SIZE AND FLUCTUATING ASYMMETRY OF MORPHOMETRIC CHARACTERS IN MICE: THEIR ASSOCIATIONS WITH INBREEDING AND t-HAPLOTYPE

Abstract Fluctuating asymmetry (FA), a ubiquitous type of asymmetry of bilateral characters, often has been used as a measure of developmental instability in populations. FA is expected to increase in populations subjected to genetic stressors such as inbreeding or environmental stressors such as toxins or parasites, although results have not always been consistent. We tested whether FA in four skeletal size characters and mandible shape was greater in a population of wild‐derived mice reared in the laboratory and subjected to one generation of inbreeding (F = 0.25) versus that in an outbred group (F= 0.00). FA did not significantly differ between the inbred and outbred groups, despite the fact that these two groups differed dramatically in fitness under seminatural population conditions. As far as we know, this is the first study to evaluate the relationship between FA and inbreeding in wild house mice, and our general conclusion is opposite that of earlier work on laboratory inbred strains of mice and their hybrids. Size for two of the characters was significantly less in inbreds than in outbreds, however, and there was a significant difference between inbreds and outbreds in the signed differences of right and left sides in one character (humerus length). Some of the mice in both groups also were heterozygous or homozygous carriers of the t‐complex. Because mice carrying this chromosome 17 variant are known to have reduced fitness, we also tested whether they had greater FA than mice carrying non‐t‐haplotypes. The overall level of a composite FA index calculated from all four characters was in fact significantly higher in the t‐bearing mice. These combined results suggest that FA is not a generally sensitive proxy measure for fitness, but can be associated with fitness reductions for certain genetic stressors.     

THE EVOLUTION OF ASYMMETRY IN SEXUAL ISOLATION: A MODEL AND A TEST CASE

We constructed a model for the evolution of sexual isolation by extending Lande's (1981) model of sexual selection. The model predicts that asymmetric sexual isolation is a transient phenomenon, characteristic of intermediate stages of divergence in sexually selected traits. Unlike the Kaneshiro (1976, 1980) proposal, our model does not depend upon drift and the loss of courtship elements to produce asymmetries in sexual isolation. According to our model, the direction of evolution cannot be predicted from asymmetry in sexual isolation. We tested some features of the model using data from an experimental study of sexual isolation in the salamander Desmognathus ochrophaeus. We tested for sexual isolation between 12 allopatric populations and found significant asymmetry in sexual isolation in about a quarter of the test cases. The highest degrees of asymmetry were associated with intermediate levels of divergence. A curvilinear relationship between isolation asymmetry and divergence was predicted by our model and was supported by statistical analysis of the salamander data.     

DEVELOPMENTAL INSTABILITY IN WILD CHROMOSOMAL HYBRIDS OF THE HOUSE MOUSE

In wild populations of the house mouse from Tunisia, fluctuating asymmetry and character size of tooth traits were compared between chromosomal races (2n = 40, all acrocentric standard karyotype, and 2n = 22, with nine fixed Robertsonian fusions) and their natural hybrids. Developmental stability was impaired in hybrids compared to both parental groups. Because genetic divergence measured by allozyme markers was low, genomic incompatibilities were not expected between the chromosomal races. This suggests that differentiation of gene systems specifically involved in development may have occurred between the chromosomal races. Support for the latter was found in the study of character size which showed that the 2n = 22 mice had smaller teeth than either the hybrid or the standard mice. The study of Tunisian chromosomal races thus shows that chromosomal evolution may lead to important changes in coadapted gene systems without involving extensive genic differentiation.     

川西农区小型兽类分布研究

经3个监测点三年定位布夹监测数据分析发现,川西农区田间小兽的密度分布有明显的院落依托性,即离院落越近的农田中密度越高。分布模式呈Logistic曲线。家栖鼠(如褐家鼠、小家鼠)比野栖鼠(如大足鼠和黑线姬鼠)更集中于院落周围。不同季节的密度分布有显著差异,春、夏季多集中在离院落50—70米范围。秋、冬季别离院落越近,其密度越高。     

ENVIRONMENTAL FACTORS INFLUENCING THE DISTRIBUTION OF SOUTHERN RIGHT WHALES (EUBALAENA AUSTRALIS) ON THE SOUTH COAST OF SOUTH AFRICA II: WITHIN BAY DISTRIBUTION

Environmental factors are thought to strongly influence the distribution and predictability of the coastal distribution of southern right whales ( Eubalaena australis ) off South Africa. Preferred habitat had generally shallow sloping sedimentary floors and was characteristically protected from open ocean swell and prevalent seasonal winds. This study investigated whether habitat choices at smaller scales (within bays) were similar. Fine scale distribution patterns (GPS) from three years' surveys (1997, 1999, 2000) were analyzed separately within the three main concentration areas St Sebastian Bay, De Hoop, and Walker Bay (containing ∼73% of cow-calf pairs and ∼49% of unaccompanied adults in the whole survey region). Whale density at this scale of within particular bays did not correlate well with predicted variables, but Chi-squared analysis strongly supported results at broader scales, in all bays. Post-hoc "choice" tests between similar areas differing in only one variable revealed that cow-calves preferred (presumed) sandy substrates and especially protection from swell. The strength and predictability of preferences shown at fine scale (where individual movement and weather variability could have great influence) provide strong support for findings at larger scales and emphasize the importance of environmental factors in the habitat choice of wintering right whales.     

广西杜鹃花属的修订及其地理分布

本文在简述广西杜鹃花属分类研究历史的基础上,对其作了全面修订,着重讨论了其中１４种（变种）的分类存在问题,并对该属在广西境内的地理分布规律也作了初步的探讨。     

使用分子遗传学方法对版纳鱼螈分布、起源及扩散的初步研究

版纳鱼螈Ichthyophis bannanicus是蚓螈目Gymnophiona在我国分布的唯一物种.由于长期以来缺乏对版纳鱼螈的分布范围及演化历史等信息的了解,本研究通过分子遗传学的方法,就其分布、起源及扩散等内容进行了探讨.结果发现,我国境内的版纳鱼螈均属同一物种,尚未出现种的分化.研究中明确证实了版纳鱼螈在泰国和越南的分布,提示了版纳鱼螈在中南半岛广为分布,甚至可能达到马来西亚境内.结果还提示水系对版纳鱼螈的分布具有较大的影响,尤其在湄公河水系中表现的最为明显.现有的信息提示版纳鱼螈有可能起源于中南半岛,并沿着澜沧江-湄公河水系、红河-珠江水系、克拉地峡-马来半岛3个方向扩散.我国境内的版纳鱼螈主要分布在云南南部地区和两广丘陵以南,来自这两大分布区的个体之间的遗传差异明显,应属于不同的进化显著单元.