Inbreeding depression in two species of Mimulus (Scrophulariaceae) with contrasting mating systems

We examined the effect of self- and cross-pollination on germination success, flowering probability, pollen and ovule production, survivorship, and adult aboveground biomass in two species of Mimulus with contrasting mating systems: the highly seifing M. micranthus and an outcrossing population of M. guttatus. Cross-pollinations were performed both within and between populations in order to examine the scale at which the genetic load is distributed. We found significant inbreeding depression in M. guttatus in four of the six traits, with the highest inbreeding depression observed in biomass (68% and 69% based on within- and between-population crosses, respectively) and lowest in ovule production (21% based on between-population crosses only). M. micranthus displayed significant inbreeding depression in only two of the six traits examined. Again, we observed the highest inbreeding depression in biomass (47–60% based on within- and between-population crosses, respectively), but both traits showing significant differences between self and outcross progeny expressed lower inbreeding depression than in M. guttatus. We detected no significant inbreeding depression for either pollen or ovule production in M. micranthus. An estimate of total inbreeding depression based on the multiplicative effects of all traits was also lower in M. micanthus than∗∗∗ in M. guttatus. Our results are consistent with the expected purging of genetic load in populations with high selfing rates. The absence of inbreeding depression in M. micranthus pollen and ovule production, two traits with strong links to fitness in a selfing annual, further suggests the important role of directional selection in determining the population's genetic load. Comparison of cross-pollinations made within and between populations revealed little evidence of divergence of genetic load among the M. micranthus and M. guttatus populations examined.     

Hummingbird color preference within a natural hybrid population of Mimulus aurantiacus (Phrymaceae)

Hummingbird flowers are typically red in color but the reasons for this are not well understood. Relatively few studies have examined hummingbird flower color preferences under natural conditions in which flower color varies within a species. We recorded hummingbird visitation rates to flowers that vary in color from yellow to red in a natural hybrid population between red‐ and yellow‐flowered Mimulus aurantiacus subspecies. We also examined whether there were any correlations between color and flower size or nectar content. Finally, we reviewed the literature on hummingbird color choice tests using feeders and flowers. There were no correlations in this population between flower color and flower size, nectar volume, or sugar concentration. Nevertheless, hummingbirds undervisited the two most yellow color classes, overvisited orange flowers, and visited the two most red color classes in proportion to their frequency in the population. While Hummingbirds preferred flowers expressing red pigments to those that did not, the flowers with the most red hue were not the most attractive, as has been observed in similar studies with other species of Mimulus. While feeder studies generally fail to show hummingbird preference for red, all studies using flowers, including those that control all floral traits other than color, find consistent preference for red. Experiments are suggested that might help disentangle hypotheses for why hummingbirds exhibit this preference.     

Developmental analysis of the evolutionary origin of vegetative propagules in Mimulus gemmiparus (Scrophulariaceae)

Mimulus gemmiparus (Scrophulariaceae), a rare endemic of Colorado, has a novel life history that depends on an unusual method of vegetative reproduction. The plants are functionally annuals; however, reproduction is asexual via propagules that have been termed gemmae. The morphological identity and the evolutionary antecedent of these propagules are unclear. We approached this problem through comparative developmental analyses of M. gemmiparus and the presumed progenitor species, Mimulus guttatus. In M. gemmiparus there are two meristems initiated in the axil of each leaf primordium. The distal meristem has the potential to produce either a lateral branch or a flower, and the proximal meristem becomes a vegetative propagule (the gemma) that is ultimately surrounded by an expanded, ensheathing petiole. The first leaves of the propagules are thickened and are the site of nutrient storage. Consequently, these propagules can be characterized morphologically as brood bulbils. Mimulus guttatus also has two meristems in each leaf axil; however, the proximal meristem typically remains dormant and serves no function in the life history of this species. Based on architectural and developmental correspondence, we hypothesize that the propagule of M. gemmiparus is homologous to the proximal meristem of M. guttatus. Comparative analysis shows that evolution of the bulbil has involved both the incorporation of features present in shoots of M. guttatus and the acquisition of novel features.     

Structure of a geranyl-alpha-pyrone from Mimulus aurantiacus leaf resin

A previously reported but misidentified geranyl-alpha-pyrone, in addition to six known compounds, was isolated from the leaf resin of Mimulus aurantiacus. HMBC NMR analyses of the geranyl-alpha-pyrone resolved uncertainties in the site of attachment of the two side chains and necessitated a revision of the previously reported structure. This compound is shown to be 3-geranyl-4-hydroxy-6-(2-hydroxypropyl)-2-pyrone.     

Environment and pollinator-mediated selection on parapatric floral races of Mimulus aurantiacus

We tested whether selection by pollinators could explain the parapatric distribution of coastal red- and inland yellow-flowered races of Mimulus aurantiacus (Phrymaceae) by examining visitation to natural and experimental populations. As a first step in evaluating whether indirect selection might explain floral divergence, we also tested for local adaptation in early life stages using a reciprocal transplant experiment. Hummingbirds visited flowers of each race at similar rates in natural populations but showed strong (>95%) preference for red flowers in all habitats in experimental arrays. Hawkmoths demonstrated nearly exclusive (>99% of visits) preference for yellow flowers and only visited in inland regions. Strong preferences for alternative floral forms support a direct role for pollinators in floral divergence. Despite these preferences, measures of plant performance across environments showed that red-flowered plants consistently survived better, grew larger and received more overall pollinator visits than yellow-flowered plants. Unmeasured components of fitness may favour the yellow race in inland habitats. Alternatively, we document a marked recent increase in inland hummingbird density that may have caused a change in the selective environment, favouring the eastward advance of red-flowered plants.     

美国凌霄柱头行为--一种促进传粉的机制

引起美国(Campsis radicans(L.)Seem.ex Bureau.)柱头开合行为的原因迄今未知.在本研究中,机械刺激、自花授粉、异花授粉以及不同授粉数量分别给予美国凌霄的柱头,以期找到影响其柱头行为的原因.研究结果表明,单纯的机械剌激可以使柱头闭合,但不能使之水永久闭合;只有柱头表明落置了足够量的花粉(>350)时柱头才会最终闭合.美国凌霄的柱头对换自花或者花粉没有差异,两种花粉授于柱头后花粉管都能成功萌发并生长.研究结果表明:美国凌霄的柱头行为不是一种异交机制,而是一种促进传粉进而增强繁殖成功的机制.     

美国凌霄柱头行为——一种促进传粉的机制(英文)

引起美国凌霄(campsis radicans(L.)Seem.ex Bureau.)柱头开合行为的原因迄今未知。在本研究中,机械刺激、自花授粉、异花授粉以及不同授粉数量的授粉分别给予美国凌霄的柱头,以期找到影响其柱头行为的原因。研究结果表明,单纯的机械刺激可以使柱头闭合,但不能使之永久闭合;只有柱头表面落置了足够量的花粉(>350)时柱头才会最终闭合。美国凌霄的柱头行为对自花或者异花花粉没有差异,两种花粉授于柱头后花粉管都能成功地萌发并生长。研究结果说明:美国凌霄的柱头行为不是一种异交机制,而是一种促进传粉进而增强繁殖成功的机制。     

Mimicry on the QT(L): genetics of speciation in Mimulus

Ecological studies suggest that hummingbird-pollinated plants in North America mimic each other to increase visitation by birds. Published quantitative trait locus (QTL) data for two Mimulus species indicate that floral traits associated with hummingbird versus bee pollination results from a few loci with major effects on morphology, as predicted by classical models for the evolution of mimicry. Thus, the architecture of genetic divergence associated with speciation may depend on the ecological context.     

烟台甜樱桃柱头的可授性、形态特征与坐果率

采用数码照相、电镜扫描、联苯胺-过氧化氢测试及去雄套袋等技术手段, 对烟台甜樱桃(Cerasus avium)花期不同发育阶段柱头的可授性、形态特征和坐果状况进行了观察。结果表明, 烟台甜樱桃在套袋状态下, 柱头可授期从开花前1天开始可持续5–7天。从杯状花期到花瓣平展期, 柱头逐渐有乳突细胞破裂并呈现分泌液, 出现渐强的可授性; 从花瓣平展期到花瓣脱落期, 柱头由暗黄渐变至暗黑, 逐渐萎缩并丧失可授性。去雄套袋及人工授粉实验结果显示, 在大蕾期、杯状花期、花瓣展放期、花瓣平展期和花瓣脱落期进行人工授粉, 烟台甜樱桃的坐果率分别为60.50%、58.33%、62.08%、57.14%和39.13%。在自然条件下烟台甜樱桃的坐果率一般为30%–42%, 传粉成功的最佳期主要发生在杯状花期至花瓣平展期。     

烟台甜樱桃柱头的可授性、形态特征与坐果率

采用数码照相、电镜扫描、联苯胺-过氧化氢测试及去雄套袋等技术手段,对烟台甜樱桃（Cerasus avium）花期不同发育阶段柱头的可授性、形态特征和坐果状况进行了观察。结果表明,烟台甜樱桃在套袋状态下,柱头可授期从开花前1天开始可持续5–7天。从杯状花期到花瓣平展期,柱头逐渐有乳突细胞破裂并呈现分泌液,出现渐强的可授性;从花瓣平展期到花瓣脱落期,柱头由暗黄渐变至暗黑,逐渐萎缩并丧失可授性。去雄套袋及人工授粉实验结果显示,在大蕾期、杯状花期、花瓣展放期、花瓣平展期和花瓣脱落期进行人工授粉,烟台甜樱桃的坐果率分别为60.50%、58.33%、62.08%、57.14%和39.13%。在自然条件下烟台甜樱桃的坐果率一般为30%–42%,传粉成功的最佳期主要发生在杯状花期至花瓣平展期。     

Inbreeding alters resistance to insect herbivory and host plant quality in Mimulus guttatus (Scrophulariaceae)

Previous studies have demonstrated genetic variation for resistance to insect herbivores and host plant quality. The effect of plant mating system, an important determinant of the distribution of genetic variation, on host plant characteristics has received almost no attention. This study used a controlled greenhouse experiment to examine the effect of self- and cross-pollination in Mimulus guttatus (Scrophulariaceae) on resistance to and host plant quality for the xylem-feeding spittlebug Philaenus spumarius (Homoptera: Cercopidae). Spittlebugs were found to have a negative effect on two important fitness components in M. guttatus, flower production and above ground biomass. One of two M. guttatus populations examined showed a significant interaction between the pollination and herbivore treatments. In this case, the detrimental effects of herbivores on biomass and flower production were much more pronounced in inbred (self) plants. The presence of spittlebug nymphs increased inbreeding depression by as much as three times. Pollination treatments also had significant effects on important components of herbivore fitness, but these effects were in opposite directions in our two host plant populations. Spittlebug nymphs maturing on self plants emerged as significantly larger adults in one of our host plant populations, indicating that inbreeding increased host plant quality. In our second host plant population, spittlebug nymphs took significantly longer to develop to adulthood on self plants, indicating that inbreeding decreased host plant quality. Taken together these results suggest that the degree of inbreeding in host plant populations can have important and perhaps complex effects on the dynamics of plant-herbivore interactions and on mating-system evolution in the host.     

Chemical Constituents of Pedicularis tricolor (Scrophulariaceae)

To search for new bioactive constituents from Pedicularis tricolor, a new xanthone named pedicutricone A (1)and a new iridoid named pedicutricoside A (2), along with fifteen known compounds, were isolated from the ethanol extracts of the whole plants of P. tricolor for the first time. Their structures were elucidated based on spectroscopic methods, including two-dimensional nuclear magnetic resonance analysis.     

High evolutionary divergence of the 5.8S ribosomal DNA in Mimulus glaucescens (Scrophulariaceae)

Ribosomal DNA sequences for the ITS 1, 5.8S, ITS 2 and adjoining regions of the 18S and 25S were obtained from Mimulus glaucescens (Scrophulariaceae) via cloned PCR products. The spacer sequences were completely unrelated to other plant taxa, although spacer lengths were approximately the same. Interestingly, the Mimulus 5.8S sequence was much more divergent than other higher-plant rDNA sequences. Consideration of the secondary structure of the 5.8S rRNA shows that most of the changes in Mimulus are compensatory and preserve the basic secondary structure of the mature RNA molecule.     

Contrasting patterns of floral and molecular variation across a cline in Mimulus aurantiacus

Abstract Steep clines in ecologically important traits may be caused by divergent natural selection. However, processes that do not necessarily invoke ongoing selection, such as secondary contact or restricted gene flow, can also cause patterns of phenotypic differentiation over short spatial scales. Distinguishing among all possible scenarios is difficult, but an attainable goal is to establish whether scenarios that imply selection need to be invoked. We compared the extent of morphological and genetic differentiation between geographically structured red and yellow floral races of Mimulus aurantiacus (bush monkeyflower; Phrymacea). Flower color was assessed in a common garden as well as in the field to determine whether variation was genetic and to quantify the extent of geographical differentiation. Population genetic differentiation at marker loci was measured for both chloroplast and nuclear genomes, and the degree of population structure within and among the floral races was evaluated. Flower color shows both a strong genetic basis and a sharp geographic transition, with pure red-flowered populations in western San Diego County and pure yellow-flowered populations to the east. In the zone of contact, both pure and intermediate phenotypes occur. Patterns of genetic differentiation at marker loci are far less pronounced, as little of the variation is partitioned according to the differences in flower color. Phenotypic differentiation (Q_ST) between populations with different flower colors is much greater than neutral genetic differentiation (F_ST). When comparisons are made between populations of the same flower color, the opposite trend is evident. Limited neutral genetic structure between the floral races, combined with sharp differentiation in flower color, is consistent with the hypothesis that current or recent natural selection maintains the cline in flower color.     

Stigma development and receptivity in almond (Prunus dulcis)

BACKGROUND AND AIMS: Fertilization is essential in almond production, and pollination can be limiting in production areas. This study investigated stigma receptivity under defined developmental stages to clarify the relationship between stigma morphology, pollen germination, tube growth and fruit set. METHODS: Light and scanning electron microscopy were employed to examine stigma development at seven stages of flower development ranging from buds that were swollen to flowers in which petals were abscising. Flowers at different stages were hand pollinated and pollen germination and tube growth assessed. Artificial pollinations in the field were conducted to determine the effect of flower age on fruit set. KEY RESULTS: Later stages of flower development exhibited greater stigma receptivity, i.e. higher percentages of pollen germination and more extensive tube growth occurred in older (those opened to the flat petal stage or exhibiting petal fall) than younger flowers. Enhanced stigma receptivity was associated with elongation of stigmatic papillae and increased amounts of stigmatic exudate that inundated papillae at later developmental stages. Field pollinations indicated that the stigma was still receptive and nut set was maintained in older flowers. CONCLUSIONS: Stigma receptivity in almond does not become optimal until flowers are past the fully open stage. The stigma is still receptive and fruit set is maintained in flowers even at the stage when petals are abscising. Strategies to enhance pollination and crop yield, including the timing and placement of honey bees, should consider the effectiveness of developmentally advanced flowers.     

Floral volatile alleles can contribute to pollinator‐mediated reproductive isolation in monkeyflowers (Mimulus)

Pollinator‐mediated reproductive isolation is a major factor in driving the diversification of flowering plants. Studies of floral traits involved in reproductive isolation have focused nearly exclusively on visual signals, such as flower color. The role of less obvious signals, such as floral scent, has been studied only recently. In particular, the genetics of floral volatiles involved in mediating differential pollinator visitation remains unknown. The bumblebee‐pollinated Mimulus lewisii and hummingbird‐pollinated Mimulus cardinalis are a model system for studying reproductive isolation via pollinator preference. We have shown that these two species differ in three floral terpenoid volatiles – d ‐limonene, β‐myrcene, and E‐β‐ocimene – that are attractive to bumblebee pollinators. By genetic mapping and in vitro analysis of enzyme activity we demonstrate that these interspecific differences are consistent with allelic variation at two loci, LIMONENE‐MYRCENE SYNTHASE (LMS) and OCIMENE SYNTHASE (OS). Mimulus lewisii LMS (MlLMS) and OS (MlOS) are expressed most strongly in floral tissue in the last stages of floral development. Mimulus cardinalis LMS (McLMS) is weakly expressed and has a nonsense mutation in exon 3. Mimulus cardinalis OS (McOS) is expressed similarly to MlOS, but the encoded McOS enzyme produces no E‐β‐ocimene. Recapitulating the M. cardinalis phenotype by reducing the expression of MlLMS by RNA interference in transgenic M. lewisii produces no behavioral difference in pollinating bumblebees; however, reducing MlOS expression produces a 6% decrease in visitation. Allelic variation at the OCIMENE SYNTHASE locus is likely to contribute to differential pollinator visitation, and thus promote reproductive isolation between M. lewisii and M. cardinalis. OCIMENE SYNTHASE joins a growing list of ‘speciation genes’ (‘barrier genes’) in flowering plants.     

Stigma factors regulating self-compatible pollination

Pollination is one of the most important steps during fertilization and sexual reproduction in plants, and numerous cell-cell interaction events occur between the pistil and the pollen grain/tube during this process. The pollen-stigma interaction is a highly selective process which leads to compatible or incompatible pollination. Previous studies in Solanaceae, Papaveraceae, and Brassicaceae provided some important insights into pollen-stigma recognition in self-incompatible systems. In recent years, considerable data have been available regarding pollen-stigma interaction during self-compatible pollination. In this review, we focus on discussing current knowledge on stigma factors that regulate pollen-stigma interaction in self-compatible systems in comparison with self-incompatible systems.     

Genetics of sex allocation in Mimulus (Scrophulariaceae)

Theoretical models of the evolution of resource allocation patterns to male and female function make the assumption that there are inherent trade-offs between the two. Here we use a quantitative genetic approach to quantify trade-offs between male and female function and to determine whether plant populations could readily respond to natural selection by quantifying the amount of genetic variation for pollen and ovule production. Both intra- and interspecific crossing designs were applied to two populations of the predominantly outcrossing Mimulus guttatus and two populations of the highly selfing congener, M. micranthus. The only significant correlations observed among pollen number, pollen size and ovule number were positive. Positive genetic correlations among the traits were sometimes reduced after removing the effect of flower size but still no significant negative correlations were detected. These results suggest that positive correlations between pollen and ovule production may be due to the joint positive correlation of these characters with the resource pool available for pollen and ovule production, as reflected by flower size. Heritabilities were moderate to high for ovule production but low for pollen number and pollen size and suggest that responses to selection would differ between the two traits. Crosses between the species revealed that there are additional genetic factors contributing to differences between the two species for corolla width, vs. pollen:ovule ratio. This is consistent with the hypothesis that genetic variation for resource acquisition may in part be responsible for the overall lack of a negative correlation between pollen and ovule production and provides a genetic explanation for little evidence of trade-offs between sexual functions in Mimulus.