PROGENY SEX RATIOS IN DIOECIOUS SILENE LATIFOLIA (CARYOPHYLLACEAE)

Most sex ratios reported for Silene latifolia are female biased. As a result of experiments performed by Correns in the early 1900s, pollen tube competition has generally been accepted as the primary cause of these skewed ratios. We did four sets of hand pollinations in which we varied the size of pollen loads and placement of pollen along the filamentous stigma. The effect of pollen load size on progeny sex ratios was not statistically significant. Of 32 maternal families, 17 contained more females than males (one ratio deviated statistically from 1:1), and 13 contained more males than females. Paternal families exhibited a greater range of sex ratios, including three with a significant female bias and one with a significant male bias. Within experiments, neither the maternal parent nor where pollen was placed had a statistically significant effect on progeny sex ratios; the paternal effect was significant in one experiment. We suggest that sex ratios in Silene latifolia are not necessarily affected by the level of pollen competition. Other factors, including variation among males and sex-linked mortality, may help explain the skewed sex ratios that characterize populations of this species. Further, Correns' observations of excess females may have resulted from his use of interspecific hybrids.     

THE EFFECT OF SERPENTINE ON THE POPULATION STRUCTURE OF SILENE DIOICA (CARYOPHYLLACEAE)

Serpentine soils are rich in heavy metals and have a distinctive flora. Silene dioica is a member of the Scandinavian serpentine plant community but is also widespread outside serpentine soils. To study the population genetic consequences of serpentine stress and the origin and evolution of serpentine populations we analyzed the isozyme genetic structure of S. dioica. Seventeen populations located in the mountains of Västerbotten and Jämtland, central Sweden, were investigated by starch gel enzyme electrophoresis. About one half of the populations grow in serpentine soils and the rest on adjacent non-serpentine sites. Analyses of allele frequencies show that both serpentine and non-serpentine populations in the northern part of the studied area (Västerbotten) are genetically similar. Evidently serpentine does not exert strong selection acting upon isozyme loci. In the south (Jämtland), however, the serpentine populations exhibit genetic differentiation. This allozyme divergence is probably not due to direct selection but rather represents the effects of isolation and genetic drift. The results suggest that S. dioica has colonized serpentine repeatedly and that the tolerant populations have a multiple origin.     

EFFECT OF DENSITY ON SECONDARY SEX CHARACTERISTICS AND SEX RATIO IN SILENE ALBA (CARYOPHYLLACEAE)

A field survey of plant and flower sex ratio and secondary sex characteristics was made in Silene alba. Female-biased plant sex ratios were found, as seems typical for the species. Sex ratio distribution correlated with a gradient of soil moisture (with the more moist area having a more female-biased ratio) and with changes in the density of Silene (intermediate and higher density areas having greater female bias). The floral sex ratio was significantly female-biased only at the site that was most female-biased in terms of plant sex ratio. Otherwise the population of flowers was significantly male-biased. Male and female plants harvested from the field differed in secondary sexual characteristics. Males had more flowers and invested proportionately more biomass in leaf, but less in root, stem and reproductive tissue than did females. Although both males and females were larger in terms of total dry weight at the moist site, males produced more flowers at the driest (high density) site. Here the female bias in plant sex ratio was intermediate, but the floral sex ratio was significantly male-biased. A glasshouse experiment was performed in which plants were grown at four densities. Density significantly influenced plant survivorship and the probability of flowering, and increased female bias in the pots, but it did not affect patterns of biomass allocation in flowering plants. Patterns of male and female biomass allocation did not differ in the experiment, except in terms of reproductive allocation (greater in females) and allocation to leaf, greater in males, but only at the lowest density. This work urges caution in interpreting differences between males and females in the field as secondary sex characteristics, since we find such properties to be overlapping under experimental conditions. It supports the idea that males and females of a species may sustain different reproductive output under differing conditions.     

SEX POLYMORPHISM IN SILENE ACAULIS (CARYOPHYLLACEAE) AND THE POSSIBLE ROLE OF SEXUAL SELECTION IN MAINTAINING FEMALES

Pollen accumulation rates and their relationship to stigma morphology and pollinator visitation behavior were compared between female and hermaphrodite sex morphs of the alpine plant Silene acaulis var. subacaulescens (Caryophyllaceae). Although stigmas of female and perfect flowers collected comparable pollen loads, stigmas of female flowers became receptive earlier in anthesis and therefore recruited a larger number of pollen tubes. Together with early receptivity young female flowers had a larger stigmatic area with longer papillae than did perfect flowers. Pollinator behavior also differed between morphs, with bumble bees spending more time probing female than perfect flowers. Differences in stigma receptivity schedules of female and perfect flowers have consequences for different opportunities for sexual selection in the two sex morphs. Female flowers, by providing a more effective gametophytic screen, have the potential to produce higher quality offspring. This is proposed as a further compensatory advantage maintaining females, with a single fitness function, in populations containing hermaphrodites which have both male and female fitness functions.     

THE QUANTITATIVE GENETICS OF SEXUAL DIMORPHISM IN SILENE LATIFOLIA (CARYOPHYLLACEAE). II. RESPONSE TO SEX-SPECIFIC SELECTION

A well-established theoretical relationship exists between genetic correlations between the sexes and the dynamics of response to sex-specific selection. The present study investigates the response to sex-specific selection for two sexually dimorphic traits that have been documented to be genetically variable, calyx diameter and flower number, in Silene latifolia. Following the establishment of a base generation with a known genetic background, selection lines were established and two generations of sex-specific selection were imposed. Calyx diameter responded directly to sex-specific selection, and the positive genetic correlation between the sexes was reflected in correlated responses in the sex that was not the basis for selection within a particular line. Flower number showed a more erratic response to sex-specific selection in that selection in some lines was initially in the wrong direction, that is, selection for a decrease in flower number resulted in an increase. These erratic responses were attributable to genotype-environment interaction as reflected in significant heteroscedasticity in variance among families. Correlated responses to selection in the sex that was not the immediate basis for selection indicated the possible existence of a negative genetic correlation between the sexes for this trait. These results test for the first time the impact of genetic correlations between the sexes on the evolutionary dynamics of sexually dimorphic traits in a plant species.     

POLLEN TUBE COMPETITION AND SELECTION FOR METAL TOLERANCE IN SILENE DIOICA (CARYOPHYLLACEAE) AND MIMULUS GUTTATUS (SCROPHULARIACEAE)

An experiment was conducted to test whether metal tolerance expressed in pollen would provide a competitive advantage during pollen tube growth and fertilization. Copper or zinc was introduced into the pistil by growing metal tolerant plants of Silene dioica or Mimulus guttatus in a nutrient solution to which metals were added. Flowers from treated and control plants were pollinated with either metal tolerant or metal sensitive pollen. The rate of pollen tube growth, the number of seeds set, and the number of viable seeds produced were measured. In general, no effects were found on the rates of pollen tube growth. However, the number of fertilizations and viable seeds were affected. When toxic metals were present, the relative success of pollen from the nontolerant parent was reduced.     

LEAF DEVELOPMENT OF PHASEOLUS VULGARIS L. IN LIGHT AND IN DARKNESS

Development of the primary bean leaf in the dark and under continuous white light was studied during 14 days after sowing. The increase in surface area of the blade is the result of a number of sequential processes. Both in the darkness and under illumination, leaf growth is characterized by an initial cell enlargement followed by intensive cell division. Cell division in etiolated leaves continues for one day longer than in illuminated ones, but it proceeds at a slower rate. Mature leaves grown under white light undergo a phase of cell enlargement after cell division has stopped. This increases their surface area up to 800 times when compared with the blade area of the embryo. This enlargement phase is almost absent in dark-grown seedlings. Consequently the blade area of etiolated leaves is only 50 times that of the embryonic state. Thus light appears to have a dual effect on leaf development: it activates cell division and induces cell expansion.     

菜豆胚乳游离核中的线粒体(简报)

在植物细胞中,细胞核以及具DNA的细胞器——质体和线粒体,在结构上通常各具自身的被膜而独立存在。关于核与这两种细胞器以及两种细胞器之间通过外膜的连接在一些植物中已有报道。最近,Yu和Russell在烟草中发现了细胞核中存在线粒体的现象。本     

矮生菜豆胚乳细胞形成时的超微结构研究

应用透射电镜对矮生菜豆的游离核胚乳、胚乳细胞及珠被绒毡层的超微结构进行了观察,揭示了胚乳细胞化过程中自由生长壁的形成方式。实验表明,在游离核胚乳中存在大量有被小泡,胚乳游离核的外层核膜发生明显外突并以出芽的方式产生有被小泡。初始胚乳细胞壁为自由生长壁,大量有被小泡融合于自由生长壁末端使之不断延伸并产生分枝。与珠被绒毡层相邻的胚乳细胞产生发达的壁内突,具有传递细胞特征;在这些细胞中出现大量的核糖体及粗糙内质网,表现现高度的合成活性。     

DIFFERENCES IN BIOMASS ALLOCATION TO REPRODUCTIVE AND VEGETATIVE STRUCTURES OF MALE AND FEMALE PLANTS OF A DIOECIOUS,PERENNIAL HERB,SILENE ALBA (MILLER) KRAUSE

The pattern of biomass allocation of males and females and the sex ratio and growth characteristics of plants from three seed-size classes in Silene alba were investigated in a greenhouse study. Seed size significantly affected adult plant size and flower production of both male and female plants, but there was no significant difference in the proportion of males and females emerging in three seed-size categories. Male and female plants differed in the proportion of total biomass allocated to vegetative and reproductive structures and these differences were consistent across all seed-size categories. Males allocated a greater proportion of their biomass to flowers than did females. Female reproductive effort was dependent upon the percentage of flowers producing mature capsules. Only females with greater than 20% fruit set have a higher reproductive expenditure than males. Consequently, female expenditure is potentially greater than males, but is spread out over a longer portion of the growing season. This difference in the timing of reproductive expenditures by males and females allows females to allocate more biomass to growth during the early flowering period and may therefore account for the common pattern in herbaceous perennial dioecious species in which adult females are larger than adult males.     

DISTANCE-DEPENDENT REGULATION OF STAMEN NUMBER IN CROSSES OF SCLERANTHUS ANNUUS (CARYOPHYLLACEAE) FROM A DISCONTINUOUS POPULATION

Scleranthus annuus is a highly inbreeding annual that has varying numbers of fertile stamens per flower. Two stamen-positions always have fully fertile stamens, whereas the other eight carry staminoids or stamens to varying degrees. I measured male expression in progeny produced by crossing individuals growing in a discontinuous population. Four types of progeny were analyzed: from self-pollinations, from cross-pollinations within a patch, from cross-pollinations between patches, and from cross-pollinations between populations. Selfed progeny showed the lowest total male fertility (25.8), followed by between-population crosses (26.7), between-patch crosses (27.4), and within-patch crosses (27.8). The effect of crossing, as measured by the relative increase in frequency of fully fertile stamens compared to selfed progeny, is highest for within-patch crosses and declines with increasing spatial separation between parents. The increase was strongest for one of the antipetalous stamen positions in progeny produced by between-patch crosses (490%). The response to crossing measured as an increase in stamen fertility was not the same for all ten stamen positions. A strong increase of fertile stamens is noted in all types of crossed progeny for the five stamen positions in the outer whorl (antipetalous stamens), positions that in selfed progeny carry staminoids. The three positions in the inner whorl that are not occupied by fully fertile stamens show varying responses to crossing.     

EFFECTS OF SIMULATED ACID RAIN ON PHASEOLUS VULGARIS L. (FABACEAE)

Experiments were performed to determine the effects of simulated acid rain on Phaseolus vulgaris L. At pH values below 3, plants exhibited a failure to attain normal height, had necrotic and wrinkled leaves, excessive and adventitious budding, and premature abscission of primary leaves. Histologically, leaves had smaller cells, less intercellular space, and smaller starch granules within the chloroplasts. Respiration rates of the treated plants increased only slightly at low pH values. Apparent rates of photosynthesis, however, increased dramatically. Both carbohydrate production and root biomass were reduced by low pH treatments, and application of Congo red indicator to the acid-treated leaf tissue showed that the cell contents were acidified to a pH of below 4.0.