NATURAL SELECTION ON FLORAL TRAITS IN TWO SPECIES OF LOBELIA WITH DIFFERENT POLLINATORS

Using both multivariate and univariate regression techniques, I measured selection acting through female reproductive success in two hermaphroditic species with precise pollen placement but different pollinators: hummingbird-pollinated Lobelia cardinalis and bumblebee-pollinated L. siphilitica. Six traits were analyzed in two populations of L. cardinalis and one population of L. siphilitica: flower number, mean number of flowers open per day, inflorescence height, number of days in flower, median-flower date and nectar-stigma distance. In another study it was found that female reproductive success in one population of L. cardinalis was much less pollen limited than in the other two populations, and it was therefore expected that selection of female reproductive traits in this population would be weaker. In the univariate analyses correlations caused nearly all traits to have significant directional selection coefficients. However, in the multivariate analyses no traits in L. siphilitica experienced directional or quadratic selection. Selection acted differently in the two L. cardinalis populations. The less pollen-limited population experienced positive directional selection on flower number and median-flower date, while in the other L. cardinalis population there was positive directional selection on flower number and nectar-stigma distance and both positive directional and positive quadratic selection on height. The functional significance of floral traits in these two species and the probable effect of increased sample sizes are discussed.     

MEASUREMENTS OF NATURAL SELECTION ON FLORAL TRAITS IN WILD RADISH (RAPHANUS RAPHANISTRUM). II. SELECTION THROUGH LIFETIME MALE AND TOTAL FITNESS

It has often been suggested that selection on floral traits in hermaphroditic plants should occur primarily through differences in male fitness. However, measurements of selection on floral traits through differences in lifetime male fitness have been lacking. We measured selection on a variety of wild radish floral traits using lifetime male fitness measures derived from genetic paternity analysis. These male fitness estimates were then combined with estimates of lifetime female fitness of the same plants to produce measurements of selection based on lifetime total fitness. Contrary to the prediction above, there was no strong evidence for selection on floral morphology through male fitness differences in any of the three years of the study, but there was strong selection for increased flower size through female fitness differences in one year. The main determinant of both male and female fitness in all years was flower number; this lead to moderately positive correlations between male and female fitness in all three years.     

STRONG INBREEDING DEPRESSION IN TWO SCANDINAVIAN POPULATIONS OF THE SELF‐INCOMPATIBLE PERENNIAL HERB ARABIDOPSIS LYRATA

Inbreeding depression is a key factor influencing mating system evolution in plants, but current understanding of its relationship with selfing rate is limited by a sampling bias with few estimates for self‐incompatible species. We quantified inbreeding depression (δ) over two growing seasons in two populations of the self‐incompatible perennial herb Arabidopsis lyrata ssp. petraea in Scandinavia. Inbreeding depression was strong and of similar magnitude in both populations. Inbreeding depression for overall fitness across two seasons (the product of number of seeds, offspring viability, and offspring biomass) was 81% and 78% in the two populations. Chlorophyll deficiency accounted for 81% of seedling mortality in the selfing treatment, and was not observed among offspring resulting from outcrossing. The strong reduction in both early viability and late quantitative traits suggests that inbreeding depression is due to deleterious alleles of both large and small effect, and that both populations experience strong selection against the loss of self‐incompatibility. A review of available estimates suggested that inbreeding depression tends to be stronger in self‐incompatible than in self‐compatible highly outcrossing species, implying that undersampling of self‐incompatible taxa may bias estimates of the relationship between mating system and inbreeding depression.     

SELECTION ON VARIANCE IN FLOWERING TIME WITHIN AND AMONG INDIVIDUALS

We model the impact of pollinator visitation rate and behavior on the short‐term evolution of population flowering phenologies determined by the distributions of flowering times within and among individual plants. Evolution of population flowering phenologies depends on the phenotypic variances and heritabilities of the within‐individual mean and variance of flowering time. In the ecological scenarios we investigate selection does not produce a correlation of the mean and variance of individual flowering time. Self‐incompatibility causes weak stabilizing selection on flowering time that acts to reduce the within‐individual variance in flowering time. Disruptive selection due to pollinator limitation acts mostly to increase the among‐individual variance in flowering time. Stabilizing selection due to pollinator attraction, or short reproductive season, acts mostly to decrease the within‐individual variance in flowering time. Temporal autocorrelation of environmental stochasticity in pollinator visitation rate strongly selects to increase the within‐individual variance in flowering time. These predictions can be tested by measuring the causal factors described above, partitioning the variance in population phenology within and among individuals, and estimating the inheritance of, and selection on, within‐individual mean and variance of flowering time.     

MEASUREMENTS OF NATURAL SELECTION ON FLORAL TRAITS IN WILD RADISH (RAPHANUS RAPHANISTRUM). I. SELECTION THROUGH LIFETIME FEMALE FITNESS

Although the role of natural selection in the evolution of floral traits has been of great interest to biologists since Darwin, studies of selection on floral traits through differences in lifetime fitness have been rare. We measured selection acting on flower number, flower size, stigma exsertion, and ovule number per flower using field data on lifetime female fitness (seed production) in wild radish, Raphanus raphanistrum. The patterns of selection were reasonably consistent across three field seasons, with strong directional selection for increased flower production in all three years, weaker selection for increased ovule number per flower in two years, and selection for increased flower size in one year. The causes of the selection were investigated using path analysis combined with multiplicative fitness components. Increased flower production increased fruit production directly, and increased numbers of ovules per flower increased the number of seeds per fruit in all three years; pollinator visitation did not influence either of these fitness components. Increased flower size was associated with increases in both the number of fruit and the number of seeds per fruit in one year, with the latter relationship being stronger. Total lifetime seed production was affected more strongly by differences in fruit production than by differences in either the number of seeds per fruit or the proportion of fertilized seeds that were viable, but all three fitness components were positively correlated with total seed production.     

RESPONSES OF FLORAL TRAITS TO SELECTION ON PRIMARY SEXUAL INVESTMENT IN SPERGULARIA MARINA: THE BATTLE BETWEEN THE SEXES

Two widespread assumptions underlie theoretical models of the evolution of sex allocation in hermaphroditic species: (1) resource allocations to male and female function are heritable; and (2) there is an intrinsic, genetically based negative correlation between male and female reproductive function. These assumptions have not been adequately tested in wild species, although a few studies have detected either genetic variation in pollen and ovule production per flower or evidence of trade-offs between male and female investment at the whole plant level. It may also be argued, however, that in highly autogamous, perfect-flowered plant taxa that exhibit genetic variation in gamete production, strong stabilizing selection for an efficient pollen:ovule ratio should result in a positive correlation among genotypes with respect to mean ovule and mean pollen production per flower. Here we report the results of a three-generation artificial selection experiment conducted on a greenhouse population of the autogamous annual plant Spergularia marina. Starting with a base population of 1200 individuals, we conducted intense mass selection for two generations, creating four selected lines (high and low ovule production per flower; high and low anther production per flower) and a control line. By examining the direct and correlated responses of several floral traits to selection on gamete production per flower, we evaluated the expectations that primary sexual investment would exhibit heritable variation and that resource-sharing, variation in resource-garnering ability, or developmental constraints mold the genetic correlations expressed among floral organs. The observed direct and correlated responses to selection on male and female gamete production revealed significant heritabilities of both ovule and anther production per flower and a significant negative genetic correlation between them. When plants were selected for increased ovules per flower over two generations, ovule production increased and anther production declined relative to the control line. Among plants selected for decreased anthers per flower, we observed a decline in anther production and an increase in ovule production relative to the control line. In contrast, the lines selected for low ovules per flower and for high anthers per flower exhibited no evidence for significant genetic correlations between male and female primary investment. Correlated responses to selection also indicate a genetically based negative correlation between the production of normal versus developmentally abnormal anthers (staminoid organs); a positive correlation between the production of ovules versus staminoid organs; and a positive correlation between the production of anthers and petals. The negative relationship between male versus female primary investment supports classical sex allocation theory, although the asymmetrical correlated responses to selection indicate that this relationship is not always expressed.     

NATURAL SELECTION ON BODY SIZE AND LAYING DATE IN THE TREE SWALLOW

I measured natural selection on body size and laying date in a population of tree swallows (Tachycineta bicolor) from 1986 to 1988. There was little evidence of selection on body size associated with overwinter survival. Disruptive selection on tarsus length, associated with female reproductive success, was detected in one of three years. Both repeatability and mother-daughter regression suggested that laying date was heritable. I found weak evidence of selection on laying date, associated with both overwinter survival and reproduction in females. The ecological implications of both tarsus length and laying date variation in this population could not be identified. Consequently, although I was able to identify the targets of natural selection, the ecological link between trait variation and selection remains unknown.     

沙枣花蜜腺的发育解剖学研究

沙枣的花蜜腺位于花柱基部的筒状花盘上,属花盘蜜腺,其蜜腺位于花盘外方,由分履表皮和产蜜组织组成。分泌表皮具有角质层和变态的气孔器。产蜜组织在发育过程中,其液泡和淀粉粒都随着蜜腺的发育呈现一定的消长规律,最后形成的蜜汁由盘状蜜腺表面的气孔泌出。     

SEXUAL SELECTION AND THE EVOLUTION OF SEXUAL SIZE DIMORPHISM IN THE WATER STRIDER,AQUARIUS REMIGIS

Sexual size dimorphism (SSD) is often attributed to sexual selection, particularly when males are the larger sex. However, sexual selection favoring large males is common even in taxa where females are the larger sex, and is therefore not a sufficient explanation of patterns of SSD. As part of a more extensive study of the evolution of SSD in water striders (Heteroptera, Gerridae), we examine patterns of sexual selection and SSD in 12 populations of Aquarius remigis. We calculate univariate and multivariate selection gradients from samples of mating and single males, for two sexually dimorphic traits (total length and profemoral width) and two sexually monomorphic traits (mesofemoral length and wing form). The multivariate analyses reveal strong selection favoring larger males, in spite of the female-biased SSD for this trait, and weaker selection favoring aptery and reduced mesofemoral length. Selection is weakest on the most dimorphic trait, profemoral width, and is stabilizing rather than directional. The pattern of sexual selection on morphological traits is therefore not concordant with the pattern of SSD. The univariate selection gradients reveal little net selection (direct + indirect) on any of the traits, and suggest that evolution away from the plesiomorphic pattern of SSD is constrained by antagonistic patterns of selection acting on this suite of positively correlated morphological traits. We hypothesize that SSD in A. remigis is not in equilibrium, a hypothesis that is consistent with both theoretical models of the evolution of SSD and our previous studies of allometry for SSD. A negative interpopulation correlation between the intensity of sexual selection and the operational sex ratio supports the hypothesis that, as in several other water strider species, sexual selection in A. remigis occurs through generalized female reluctance rather than active female choice. The implications of this for patterns of sexual selection are discussed.     

群心菜花蜜腺的发育解剖学研究

群心菜（Ｃａｒｄａｒｉａｄｒａｂａ（Ｌ．）Ｄｅｓｖ）花蜜腺６枚,包括４枚侧蜜腺的和２枚中蜜腺,属十字花科侧中蜜腺类型中的侧分离中间亚型,侧中蜜腺结构相同,都由分泌表皮,产蜜组织组成,分泌表皮顶部分布的有变态气孔器,产蜜组织中无维管束分布,属较原始的十字花科花蜜腺亚型类型,在花的各部分基本分化完成后,由花托表层细胞恢复分裂能力形成蜜腺原基,蜜腺原基经分裂,分化和形态建成,发育形成成熟蜜腺,侧中蜜腺发     

Fruit set in a deceptive orchid: The effect of flowering phenology,display size,and local floral abundance

We investigated the effect of flowering time, display size, and local floral density on fruit set in Tolumnia variegata, a pollination-limited orchid that offers no reward to its pollinator(s). During 1990, natural variation in flowering time, display size, and fruit set were monitored in 508 plants at one locality in Puerto Rico. The following season, orchid floral abundance per host tree (Randia aculeata) was manipulated to investigate its effect on fruit set. Four floral abundance treatments were established (700, 500, 300, and 100), each replicated four times. Flowering time was the most important trait affecting fruit set. The proportion of plants setting at least one fruit was significantly high early and late in the season, but low during the flowering peak. Thus, strong disruptive selection differential on flowering phenology was found. Display size had little effect on fruit set. A weak, but significant disruptive selection differential on display size was found. Orchid floral abundance per host tree had a significant effect on fruit set. Early in the season, T. variegata flowers with intermediate number of conspecific flowers exhibited a greater probability of setting fruit than those in host trees with fewer or more flowers. Our results show that flowering phenology may be evolutionarily unstable, possibly a consequence of the deception pollination system. Furthermore, a deception strategy would be relatively unsuccessful in populations where plants are found in either very dense or sparse patches.     

准噶尔无叶豆的开花物候与生殖特征

 为了研究沙漠稀有植物准噶尔无叶豆(Eremosparton songoricum)的开花物候特征及其对生殖成功的影响, 2005和2006年连续两年对其自然种群的开花物候和开花过程中的花部表型变化进行了观察, 并运用相对开花振幅、开花强度和开花同步性等开花物候指数研究了开花物候特征。结果表明: 准噶尔无叶豆在5月下旬至6月中下旬开花, 其种群、个体、花序和单花的花期分别历时26~29 d、8~10 d、5~7 d和2~3 d。单花开花进程依其形态和散粉特征可分为散粉前期、散粉初期、散粉盛期和凋谢期4个时期。其个体水平的开花物候进程(开花振幅曲线)呈渐进式单峰曲线, 具有很高的开花同步指数, 表现出一种集中开花的模式。开花物候指数与座果数之间的相关分析结果表明, 始花日期与花期持续时间存在负相关关系, 而与开花数和座果数存在正相关关系; 花期持续时间与开花数和座果数存在显著正相关关系。准噶尔无叶豆个体开花物候在很大程度上是由其遗传因子决定的, 而开花物候在年度间的变异, 可能是由于荒漠气候的差异(主要是水分和温度的差异)所引起的。作为沙漠窄域分布特有种, 准噶尔无叶豆在环境和人为干扰的双重选择压力下, 为了吸引更多的传粉者访问而达到生殖成功, 形成了大量集中开花的模式。     

SEXUAL SELECTION AND SURVIVAL SELECTION ON WING COLORATION AND BODY SIZE IN THE RUBYSPOT DAMSELFLY HETAERINA AMERICANA

I review methodological problems that can lead to false evidence for selection on secondary sexual characters and present a study of selection in rubyspot damselflies (Hetaerina americana) that avoids these pitfalls. Male rubyspots have a large red spot on each wing that grows to a terminal size after sexual maturity. Selection gradient analyses revealed evidence for positive sexual and survival selection on both terminal wing spot size and body size. Phenotype manipulations confirmed that wing spot size was subject to direct sexual selection, but showed that the positive slope of survival on wing spot size was an indirect effect of selection on unmeasured traits. This study provides the strongest evidence yet for sexual selection on coloration in Odonata, but also provides clear examples of why phenotypic selection statistics must be calculated and interpreted cautiously.     

荆条花蜜腺发育解剖学研究

荆条（Ｖｉｔｅｘ ｃｈｉｎｅｎｓｉｓ Ｍｉｌｌ．）花蜜腺属于淀粉型子房蜜腺,呈圆筒状环绕于子房的基部。蜜腺外观上无特殊结构,表面有。由分泌表皮和泌蜜组织组成,包括分泌表皮、气孔器、泌蜜薄壁组织和维管束。密腺和子房壁起源相同。花蕾膨大期,泌蜜组织细胞中产生大液泡;露冠期,泌蜜组织中形成维管束;花蕾初放期,分泌表皮细胞分化形成气孔器,无气孔下室,淀粉粒的积累在此期达到高峰;盛花期,蜜腺中已无淀粉粒,密     

GENETIC CONSTRAINTS ON LIFE-HISTORY EVOLUTION: QUANTITATIVE-TRAIT LOCI INFLUENCING GROWTH AND FLOWERING IN ARABIDOPSIS THALIANA

We have mapped genes causing life-history trade-offs, and they behave as predicted by ecological theory. Energetic and quantitative-genetic models suggest a trade-off between age and size at first reproduction. Natural selection favored plants that flower early and attain large size at first reproduction. Response to selection was opposed by a genetic trade-off between these two components of fitness. Two quantitative-trait loci (QTLs) influencing flowering time were mapped in a recombinant inbred population of Arabidopsis. These QTLs also influenced size at first reproduction, but did not affect growth rate (resource acquisition). Substitutions of small chromosomal segments, which may represent allelic differences at flowering time loci, caused genetic trade-offs between life-history components. One QTL explained 22% of the genetic variation in flowering time. It is within a few centiMorgans (cM) of the gigantea (GI) locus, and may be allelic with GI. Sixteen percent of the genetic variation was explained by another QTL, FDR1, near 18 cM on chromosome II, which does not correspond to any previously identified flowering-time locus. These life-history genes regulate patterns of resource allocation and life-history trade-offs in this population.     

SEXUAL SELECTION ON MATURATION TIME AND BODY SIZE IN SPHENARIUM PURPURASCENS (ORTHOPTERA: PYRGOMORPHIDAE): CORRELATED RESPONSE TO SELECTION

We measured in the field the intensity and mode (i.e., directional, stabilizing) of sexual selection acting jointly on body size and time of sexual maturity in the univoltine, polygamous grasshopper Sphenarium purpurascens. Statistical analyses indicated that selection favored large and protandrous males in terms of a higher mating success. At the same time, evidence of correlational selection acting simultaneously on body size and time to sexual maturity was found. Thus, selection should strengthen the relationship between body size and the time of sexual maturity. Theoretical work suggests the existence of a trade-off between reaching a large size and early sexual maturation in insects. The present study does not support the existence of this kind of trade-off. Recent theoretical and empirical work like the one reported here suggests that such a trade-off may not be necessarily expected if growth rates (which are often assumed to be invariable) are affected by environmental and genetic factors.     

POPULATION STRUCTURE ALONG A STEEP ENVIRONMENTAL GRADIENT: CONSEQUENCES OF FLOWERING TIME AND HABITAT VARIATION IN THE SNOW BUTTERCUP,RANUNCULUS ADONEUS

Few studies have determined how gene flow and selection interact to generate population genetic structure in heterogeneous environments. One way to identify the potential role played by natural selection is to compare patterns of spatial genetic structure between different life cycle stages and among microenvironments. We examined patterns of spatial structure in a population of the snow buttercup (Ranunculus adoneus), using both adult plants and newly emerged seedlings. The study population spans a steep environmental gradient caused by gradual melting of snow within a permanent snowbed. Early-melting sites are characterized by denser vegetation, more fertile soils, and a longer growing season than late-melting sites tens of meters away. The flowering time of R. adoneus is controlled entirely by time of snowmelt, so the contiguous population is phenologically substructured into a series of successively flowering cohorts, reducing the opportunity for direct pollen transfer between early- and late-melting sites. For four highly polymorphic enzyme loci in this tetraploid species, there was subtle, but statistically significant, genetic differentiation between early, middle, and late-melting cohorts; adults usually showed greater differentiation among snowmelt zones than did seedlings. At two loci in adults and one locus in seedlings, homozygotes were more common than predicted at Hardy-Weinberg equilibrium, even when assuming maximum levels of double reduction during meiosis. This pattern suggests the occurrence of self-fertilization and/or population substructure. To determine how spatial isolation and phenological separation each contribute to genetic substructure, we used bivariate regression models to predict the numbers of allele differences between randomly paired individuals as a function of meters separation in space and days separation in flowering time. For newly emerged seedlings, we found that spatial separation was positively associated with genetic difference, but that the additional contribution of phenological separation to genetic difference was not significant. This implies that seeds and/or pollen move effectively across the snowmelt gradient, despite differences in flowering time. As was true for seedlings, spatial separation between paired adults contributed to greater genetic difference, but for a given spatial separation, the genetic difference between adult plants was reduced by phenological separation. This result implies that postemergence selection is favoring at least some seeds that migrate across the snowmelt gradient. Directional gene flow across the snowmelt gradient probably results from a genetic source-sink interaction, that is, the colonization of ecologically marginal late-melting sites by high quality seeds produced by the larger subpopulation in early-melting sites. Effective gene flow from high to low quality microenvironments is likely to impede adaptation to late-melting locations.     

Pollen limitation and its influence on natural selection through seed set

Stronger pollen limitation should increase competition among plants, leading to stronger selection on traits important for pollen receipt. The few explicit tests of this hypothesis, however, have provided conflicting support. Using the arithmetic relationship between these two quantities, we show that increased pollen limitation will automatically result in stronger selection (all else equal) although other factors can alter selection independently of pollen limitation. We then tested the hypothesis using two approaches. First, we analysed the published studies containing information on both pollen limitation and selection. Second, we explored how natural selection measured in one Ontario population of Lobelia cardinalis over 3 years and two Michigan populations in 1 year relates to pollen limitation. For the Ontario population, we also explored whether pollinator‐mediated selection is related to pollen limitation. Consistent with the hypothesis, we found an overall positive relationship between selection strength and pollen limitation both among species and within L. cardinalis. Unexpectedly, this relationship was found even for vegetative traits among species, and was not found in L. cardinalis for pollinator‐mediated selection on nearly all trait types.