Life history variation in Campanula americana (Campanulaceae): population differentiation

Genetic variation in life history traits has important consequences for life-history evolution. Here we report the results of a greenhouse experiment investigating the broad sense genetic basis of variation in life history traits within and among five populations of Campanula americana distributed along a latitudinal gradient. The populations exhibit differentiation for a number of morphological traits (seed weight, number of branches, final plant size, number of capsules) and the phenological traits, days to emergence, days to bolting, the onset of flowering, and the duration of flowering. Families within populations differed only in days to emergence and seed weight. These results suggest that the life history differences among populations are genetically based. In addition, two life history types—winter annuals and biennials—have previously been reported from natural populations of Campanula americana. This experiment identified a third type—summer annuals from the Florida population.     

Floral size variation in Campanula rotundifolia (Campanulaceae) along altitudinal gradients: patterns and possible selective mechanisms

We investigated patterns of flower‐size variation along altitudinal gradients in the bee‐pollinated perennial Campanula rotundifolia (Campanulaceae) by examining 22 Norwegian populations at altitudes between 240 and 1100 m a.s.l. We explored potential mechanisms for the underlying pattern by quantifying pollinator–faunal composition, pollinator‐visitation rates and pollen limitation of seed set in subsets of the study populations. Despite a decrease in plant size, several measures of flower size increased with elevation. Bumble bees were the main pollinators at both alpine and lowland sites in the study area. However, species composition of the pollinator fauna differed, and pollinators were larger in higher‐elevation than in lower‐elevation sites. Pollinator visitation rates were lower at higher‐elevations than at lower elevations. Pollen limitation of seed set did not vary significantly with altitude. Our results are consistent with differences in bumble‐bee size and visitation rates as causal mechanisms for the relatively larger flowers at higher elevations, in three non‐mutually exclusive ways: 1) Larger flowers reflect selection for increased attractiveness where pollinators are rare. 2) Larger and fewer flowers represent a risk avoidance strategy where the probability of pollination is low on any given day. 3) Flower size variation reflects selection to improve the fit of pollinators with fertile structures by matching flower size to pollinator size across sites.     

Resolving the evolutionary history of Campanula (Campanulaceae) in western North America

Recent phylogenetic works have begun to address long-standing questions regarding the systematics of Campanula (Campanulaceae). Yet, aspects of the evolutionary history, particularly in northwestern North America, remain unresolved. Thus, our primary goal in this study was to infer the phylogenetic positions of northwestern Campanula species within the greater Campanuloideae tree. We combined new sequence data from 5 markers (atpB, rbcL, matK, and trnL-F regions of the chloroplast and the nuclear ITS) representing 12 species of Campanula with previously published datasets for worldwide campanuloids, allowing us to include approximately 75% of North American Campanuleae in a phylogenetic analysis of the Campanuloideae. Because all but one of North American Campanula species are nested within a single campanuloid subclade (the Rapunculus clade), we conducted a separate set of analyses focused specifically on this group. Our findings show that i) the campanuloids have colonized North America at least 6 times, 4 of which led to radiations, ii) all but one North American campanuloid are nested within the Rapunculus clade, iii) in northwestern North America, a C. piperi-C. lasiocarpa ancestor gave rise to a monophyletic Cordilleran clade that is sister to a clade containing C. rotundifolia, iv) within the Cordilleran clade, C. parryi var. parryi and C. parryi var. idahoensis exhibit a deep, species-level genetic divergence, and v) C. rotundifolia is genetically diverse across its range and polyphyletic. Potential causes of diversification and endemism in northwestern North America are discussed.     

Genetic Diversity in Alpine and Foothill Populations of Campanula rotundifolia (Campanulaceae)

Climatic constraints on insects in alpine environments have important consequences for the biology of their plant mutualists; in particular, reduced insect diversity and activity in alpine plant populations can result in pollinator-limited seed set and, potentially, in low genetic diversity. However, highly effective pollination by bumblebees in alpine populations can compensate for low visitation rates. In this study we hypothesized that, because of highly effective pollination by bumblebees, alpine populations of Campanula rotundifolia would not experience more frequent cycles of pollinator limitation than low-elevation populations and would therefore exhibit comparable levels of genetic variability and inbreeding to those found in foothill populations. Enzyme electrophoresis was used to assess genetic variability at nine putative loci in alpine and foothill populations of C. rotundifolia in Colorado. Genetic variability in C. rotundifolia was found to be comparable to that reported for other long-lived herbaceous perennials. Measures of genetic variability and fixation indices did not differ between high- and low-elevation populations and were consistent with Hardy-Weinberg expectations. Nonsignificant F(ST) values indicated no genetic differentiation among all populations.     

The effect of maternal and paternal environments on seed characters in the herbaceous plant Campanula Americana (Campanulaceae)

Maternal environments typically influence the phenotype of their offspring. However, the effect of the paternal environment or the potential for joint effects of both parental environments on offspring characters is poorly understood. Two populations of Campanula americana, a woodland herb with a variable life history, were used to determine the influence of maternal and paternal light and nutrient environments on offspring seed characters. Families were grown in the greenhouse in three levels of light or three levels of nutrients. Crosses were conducted within each environmental gradient to produce seeds with all combinations of maternal and paternal environments. On average, increasing maternal nutrient and light levels increased seed mass and decreased percentage germination. The paternal environment affected seed mass, germination time, and percentage germination. However, the influence of the paternal environment varied across maternal environments, suggesting that paternal environmental effects should be evaluated in the context of maternal environments. Significant interactions between family and the parental environments for offspring characters suggest that parental environmental effects are genetically variable. In C. americana, the timing of germination determines life history. Therefore parental environmental effects on germination timing, and genetic variation in those parental effects, suggest that parental environments may influence life history evolution in this system.     

Seed coat morphology of some Turkish Campanula (Campanulaceae) species and its systematic implications

The seed coat morphology of 10 Campanula species from Turkey, 3 of which are endemic, was studied using scanning electron microscopy. Characteristics of the seeds and their surface morphology were described and compared. Two different types were described based on seed surface features. The Type 1 seed surface was characterized by a reticulate pattern; only C. Olympica belonged to this type. The Type 2 seed surface had a striate testa and was the most common type. It included C. Lyrata subsp. lyrata, C. rapunculoides L. subsp. rapunculoides, C. glomerata L. subsp. hispida, C. involucrata, C. saxonorum, C. persicifolia, C. latiloba subsp. latiloba, C. lactiflora and C. Rapunculus L. var. lambertiana. However, some differences among these species were determined on the basis of their seed surface features at the microstructural level. These features were evaluated as possibly consistent parameters in the delimination of the Campanula taxa studied.     

Effects of inbreeding on male function and self-fertility in the partially self-incompatible herb Campanula rapunculoides (Campanulaceae)

We examined the effect of inbreeding on fitness (through both male and female functions) and changes in self-fertility in the partially self-incompatible species Campanula rapunculoides. Individuals in natural populations of C. rapunculoides varied extensively in their strength of self-incompatibility (SI). We crossed 11 individuals that differed in their strength of SI to generate families with four levels of inbreeding (f = 0.0, 0.25, 0.5, and 0.75). Progeny were scored for three traits related to male fitness and for outcrossed and selfed seed production. Analyses of variance revealed significant inbreeding depression for the three male traits and seed set. Families with strong or weak SI differed in their response to inbreeding. Families with weak SI had lower levels of inbreeding depression for most traits than families with strong SI, but strong SI families had a greater increase in selfed seed set, but not self-fertility, with inbreeding. Finally, we found evidence of a significant linear response to inbreeding for all three male reproductive traits and outcrossed seed, indicating that inbreeding depression was primarily caused by partially or fully recessive deleterious alleles. Variation in genetic load was associated with variation in self-fertility, a finding that suggests an evolutionary role for partial self-fertility in natural populations of C. rapunculoides.     

Parental effects in a partially self-incompatible herb Campanula rapunculoides L. (Campanulaceae): influence of variation in the strength of self-incompatibility on seed set and progeny performance

We employ a full reciprocal diallel design between 10 parental plants that differed in their strength of self-incompatibility (SI; strong, intermediate, and weak) to examine parental effects on seed set and 10 components of fitness of progeny performance in Campanula rapunculoides. We perform ANOVAs to separate the influence of the strength of SI and the identity of the maternal and paternal parent on family performance. We calculate the phenotypic and genetic correlations between traits to determine potentially evolutionary constraints. Finally, we employ maximum likelihood methods to estimate the components of quantitative genetic variance, as defined by Cockerham and Weir in their BioModel c. Our most significant finding is that weak SI plants have high outcrossed seed set as maternal parents. We argue that direct or pleiotropic effects of modifiers of SI probably cause this. Second, we find that extranuclear interactions, as defined by the BioModel, have strong effects on seed set and several vegetative and flowering traits. These findings indicate that some maternal plants selectively provision seeds sired by specific paternal donors and that some of this variation appears to be associated with modifiers of the strength of SI. We find other sources of significant quantitative genetic variation for all of the traits we examine and discuss the possible role these play in the evolution of the reproductive system. Taken together, our findings show that variation in the strength of SI may influence levels of quantitative genetic variation that, in turn, can influence the reproductive success of individuals in C. rapunculoides     

The influence of light on paternal plants in Campanula americana (Campanulaceae): pollen characteristics and offspring traits

Offspring trait expression is determined by the combination of parental genes and parental environments. Although maternal environmental effects have been widely characterized, few studies have focused on paternal environmental effects. To determine whether light availability influences pollen and offspring traits in the woodland herb Campanula americana, we reared clones of 12 genotypes in two light levels. In the parental generation we measured pollen number and size. Plants grown under high light produced more pollen grains per flower than those grown under low light. However, the response was genotype specific; some individuals responded little to changes in light availability while others substantially reduced pollen production. As a consequence, paternity ratios may vary between light environments if more pollen is associated with greater siring success. We crossed a subset of these plants to produce the offspring generation. The paternal and maternal light environments influenced offspring seed mass, percentage germination, and days to germination, while only maternal light levels influenced later life traits, such as leaf number and size. Maternal and paternal environmental effects had opposite influences on seed mass, percentage germination and days to germination. Finally, there was no direct relationship between light effects on pollen production and offspring trait expression.     

箭叶淫羊藿同质园栽培居群非腺毛多样性及其分类学启示

淫羊藿是国际上多年来重点研究的药用植物.箭叶淫羊藿(Epimedium sagittatum)是淫羊藿属分布最广,也是形态变异最大的物种,箭叶淫羊藿复合群种分类中存在诸多疑点和争议.为了重建淫羊藿属的系统发育,我们利用数码显微摄影技术对10省16个箭叶淫羊藿同质园栽培居群的叶背非腺毛性状进行了比较研究.结果表明,箭叶淫羊藿非腺毛性状的主要特征包括:细胞数、非腺毛长度、是否存在伸长细胞、顶细胞形态及由此导致的顶细胞直径和壁厚差异.不同居群的非腺毛形态和特征存在巨大差异和丰富的多样性,其中项细胞形状是最重要的特征,并与全长、有无伸长细胞、项细胞比例、顶细胞直径和壁厚等多个性状密切关联.聚类分析将16个居群划分为两个类型,即梭形组和剑形组.变异式样的分析表明个体间和居群间整体呈现连续变异,丰富的多样性可能是多型种的表现.但形态上的分化往往来源于关键性状特征的强烈选择作用引起的居群间适应性分化,同质园栽培个体和居群间的形态差异暗示着适应性相关性状的遗传分化,居群间的形态差异应为地方适应性的结果,这种差异可能是物种形成的雏形.     

Polymorphism in the Aegean "five-loculed" species of the genus Campanula, Section Quinqueloculares (Campanulaceae)

Population variation in species of Campanula Section Quinqueloculares and related taxa has been studied. 19 species and 48 populations from the Aegean region have been sampled. There are few distinct character states available for analysis so that metric variables have been scored. 11 leaf variables and 19 flower variables have been measured in up to 683 plants. Regressions of variables with overall size have been calculated and where significant allometric variation has been detected residual scores have been used as variables. Hierarchical analysis of variance has indicated that a large part of the total variation is found between populations within species rather than within populations. Between the 10 most closely related species there is a considerable overlap in population means of variables. Multivariate analysis also fails to identify distinct species clusters. In particular principal components analysis provides additional evidence of overlap between species. It is suggested that a detailed study of genetic variation within and between species is required in order to establish the presence of any distinct phylogenetic lineages.     

Reproduction inCampanula afra (Campanulaceae): Mating system and the role of the pollen collecting hairs

The aim of the present investigation is to compare two geographically well separated populations ofC. afra with different mating systems with emphasis on (1) factors controlling the length of the male phase with special attention paid on the pollen collecting hairs, (2) crossing ability among populations with different mating systems; shouldC. afra be regarded as one or several species?, and (3) correlations between pollen germinability and mating system; how does pollen germinability vary with anthesis and stigma development? From the results it can be concluded that the length of the male phase and the development of the stigma depend on the stimulation of the pollen collecting hairs rather than on the amount of pollen removed. Furthermore, the length of the male phase was found to be correlated with the mating system, i.e., in the population with a mixed mating system, the differences between treated and untreated flowers were less pronounced than in the population being obligately outcrossing. I also found correlations between mating system and pollen germinability, where the population with a mixed mating system had a high germinability at stigma receptivity, whereas for the xenogamous population, the germinability approached zero at stigma development. No crossing incompatibility was found between the two populations ofCampanula afra in any of the combinations.     

Random Amplified Polymorphic DNA (RAPD) Variation among Populations of the Insular Endemic Plant Campanula microdonta(Campanulaceae)

RAPD variation was examined in nine populations of Campanulamicrodonta Koidz., endemic to the Izu Islands, Japan. Ninety-eightbands were obtained for all populations, 94% of which were polymorphicat least within a population. Shannon's H values were calculated;these have frequently been used in RAPD studies to estimategenetic diversity. The values within populations did not correlatewith the allozyme gene diversity estimated by a previous studyor with distance from the Japanese mainland. The possible reasonsfor this discrepancy are different selection regimes betweenthe two markers, higher RAPD mutation rates, and each marker'sdifferent coverage of genomes. Cluster analysis of genetic similaritiessuggested that colonization of each island probably occurredonce, except for Miyake Island, where immigration has occurredat least twice. Copyright 2001 Annals of Botany Company AMOVA, Campanula microdonta Koidz., insular endemic plant, genetic diversity, population genetic structure, RAPD     

Floral longevity in Campanula americana (Campanulaceae): a comparison of morphological and functional gender phases

Plastic responses to pollination and/or pollen removal may shift a flower's realized longevity closer to an optimal longevity that maximizes reproductive output per unit resource input. In particular, conditional responses to pollen removal and pollen deposition are expected in flowers of protandrous species in which the lengths of the male and female phases may be adjusted independently. We investigated plasticity in floral longevity in Campanula americana, a protandrous, insect-pollinated herb. In greenhouse studies, we found that the longevity of the morphological male phase was shortened by pollen removal and that the longevity of the morphological female phase was shortened by pollen deposition. In a natural population, male and female sexual functions saturated within a few hours of morphological gender phase onset. In contrast to theory, morphological gender phases did not terminate immediately upon saturation of sexual function. These findings are discussed in the context of current floral longevity theory.     

Morphological, anatomical and palynological studies on Asyneuma michauxioides (Campanulaceae)

The morphological, anatomical and palynological features of Asyneuma michauxioides (Boiss.) Damboldt (Campanulaceae) were investigated. This species is an East Mediterranean Floristic Element and an endemic in Turkey. In anatomical studies, cross-sections of the root and stem were examined and demonstrated by photographs. It has conspicuous endodermis in which the Casparian bands are indistinct in the stem. According to the result of this study, it was found that pollen grain of this taxon was 3-porate sometimes 4-porate and oblate-spheroidal. Its structure was tectate and sculpture was small granules and pits with densely spaced spinules. The seeds of the taxon were eliptic to ovate in shape, laterally compressed, brown to light brown in colour, glabrous and shiny.     

Hereditary variation and genetic recombination in Koji-molds (Aspergillus oryzae and Asp. sojae). I. Natural variation

Natural variation in monospore lines of Koji-molds (Asp. oryzae and Asp. sojae), isolated from commercial Koji material or soil and from laboratory stock cultures, has been observed. We can divide the 58 strains of Koji-molds investigated into two groups; one group consists of inconstant strains which are very liable to produce natural variants, and the other consists of strains which remain constant through successive single spore culture. The inconstant strains develop colonies bearing various proportions of conidia and aerial mycelium (X-type). They generally form large conidia (Asp. oryzae var. magnasporus) but sometimes medium sized conidia (Asp. oryzae s. str.), which produce large conidia occasionally. The colonies of the constant strains show abundant conidial formation and smooth surfaces (C-type). The conidia are mostly small (Asp. oryzae var. microsporus) but sometimes medium in size (Asp. oryzae s. str.). The colony types of the variants are as follows: C (Conidial type, whole colony covered with conidia), M (mycelial type), R (restricted in growth rate), St (sterile type, little sporulation on all media tested), Nit (requiring reduced nitrate, very faint growth on Czapek's agar), and LS (semi lethal, growth cease immediately after germination). Pedigree cultures of the 8 inconstant strains have been made, but no definite segregation ratios for each variant type have been recognized through successive generations. The LS and N types commonly occur spontaneously from the M-type.     

Variation in floral longevity between populations of Campanula rotundifolia (Campanulaceae) in response to fitness accrual rate manipulation

Floral longevity, the time between corolla expansion and senescence, contributes directly and indirectly to a plant's overall fitness. Though mating opportunities for insect-pollinated species often differ among populations, few empirical studies have addressed whether floral longevity varies in a manner consistent with these differences. I conducted experiments at thermally distinct sites to examine whether the prevailing floral longevity model predicted such variation between a montane and an alpine population of Campanula rotundifolia. Staminate phase duration was significantly shorter for montane vs. alpine C. rotundifolia flowers in the presence of pollinators, but significantly longer when pollinators were excluded. Montane flowers had a significantly higher female fitness accrual rate, significantly shorter total longevities, and, unlike alpine flowers, were not pollen-limited. Delaying pollinator access to pistillate phase flowers significantly increased total longevity in alpine flowers only. Significant differences in total longevity between populations resulted from an extended pistillate phase in alpine flowers. Overall, the prevailing model accurately predicted the total floral longevity trends found here. However, I provide novel evidence for geographic and gender-specific plasticity in the floral longevity response to fitness accrual rate variation, suggesting C. rotundifolia populations in this study may be attuned to local schedules of pollinator activity.     

Character transformation and relationships in Corallorhiza (Orchidaceae: Epidendroideae). II. Morphological variation and phylogenetic analysis

Morphological and anatomical study of Corallorhiza, a genus of primarily New World leafless mycoparasitic orchids, was undertaken in order to produce a hypothesis of relationships among the species and to gain some understanding of character transformations. Cladistic analysis of the resulting data set gave two most parsimonious trees. Analysis of combined plastid DNA and morphological data yielded a single topology, identical to one of the two from the analysis of morphological data alone. Molecular data do not conflict with the morphological data set, and provide more resolution within the C. maculata complex. The combined data indicate that C. striata is the sister group to the remainder of the genus; the circumboreal C. trifida also occupies a basal position. Corallorhiza wisteriana and C. odontorhiza comprise the sister group to the C. maculata + C. mertensiana + C. bulbosa clade. Only two synapomorphies, presence of the coralloid rhizome and loss of leaves, unite the species of Corallorhiza. The coralloid rhizome appears to be a paedomorphic development, due to its similarity to a protocorm; if so, it too is a loss character and may be considered only weak support for monophyly of the genus. Predominant autogamy, seen in C. trifida and cleistogamous C. odontorhiza, has probably arisen independently in these taxa.     

Morphological changes in the testis and epididymis of camels (Camelus dromedarius). Part I

The normal anatomy and morphological changes in the testis and epididymis, due to age and seasonal variations, were studied in camels (Camelus dromedarius) between 4 and 20 years of age. The testes had a dark gray colour and were situated in the perineum. The weight ranged from 32 to 225 g. The length and maximum cross-diameter were 6--13 and 3--6.5 cm, respectively. The maximum weight of the left testis in 12-year-old camels during the winter months (December to March) was 225 g, and the minimum weight in the camels of the same age group during the summer months (April to July) was 181 g. The seasonal influence on the weight of the testis was very profound in the camels of 9--14 years. The weight of the epidiymis ranged from 10 to 40 g; their weight increased in all age groups during the winter months.