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PubMed1997

Microsatellite variation and evolution in the Mimulus guttatus species complex with contrasting mating systems.

Mutational variability at microsatellite loci is shaped by both population history and the mating system. In turn, alternate mating systems in flowering plants can resolve aspects of microsatellite loci evolution. Five species of yellow monkeyflowers (Mimulus sect. Simiolis) differing for historical rates of inbreeding were surveyed for variation at six microsatellite loci. High levels of diversity at these loci were found in both outcrossing and selfing taxa. In line with allozyme studies, inbreeders showed more partitioning of diversity among populations, and diversity in selfing taxa was lower than expected from reductions in effective population size due to selfing alone, suggesting the presence of either population bottlenecks or background selection in selfers. Evaluation of the stepwise mutation model (a model of DNA replication slippage) suggests that these loci evolve in a stepwise fashion. Inferred coalescent times of microsatellite alleles indicate that past bottlenecks of population size or colonization events were important in reducing diversity in the inbreeding taxon.

Alleles↗
PubMed1993

Variation in the ribosomal internal transcribed spacers (ITS1 and ITS2) among eight taxa of the Mimulus guttatus species complex.

The internal transcribed spacer region (ITS1 and ITS2) of the 18S-25S nuclear ribosomal DNA sequence and the intervening 5.8S region were sequenced from three individuals in each of eight taxa of the Mimulus guttatus species complex. Three discrete variants, or "types," of ITS sequences were found, among which 30%-40% of sites differed, compared with 1%-2% within types. Dot plots indicate that these types were not related by conspicuous rearrangements or inversions. More than one ITS type was often found in the same taxon, and two of three ITS types span species boundaries, indicating their presence prior to speciation. These ITS sequences showed essentially no positional homology with the nearest sequenced relative, tomato. In contrast, the 5.8S region was relatively unvaried, with 8 of 162 sites varied in the sample among all eight taxa. The phylogeny inferred by the most common ITS sequence type, rooted by the two other ITS types, agreed with isozymes in showing the distinctness of M. nudatus, M. laciniatus, and M. tilingii from the other five taxa.

Base Sequence↗
PubMed2001

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.

Animals↗
PubMed2001

Evidence for Dobzhansky-Muller incompatibilites contributing to the sterility of hybrids between Mimulus guttatus and M. nasutus.

Both chromosomal rearrangements and negative interactions among loci (Dobzhansky-Muller incompatibilities) have been advanced as the genetic mechanism underlying the sterility of interspecific hybrids. These alternatives invoke very different evolutionary histories during speciation and also predict different patterns of sterility in artificial hybrids. Chromosomal rearrangements require drift, inbreeding, or other special conditions for initial fixation and, because heterozygosity per se generates any problems with gamete formation, F1 hybrids will be most infertile. In contrast, Dobzhansky-Muller incompatibilities may arise as byproducts of adaptive evolution and often affect the segregating F2 generation most severely. To distinguish the effects of these two mechanisms early in divergence, we investigated the quantitative genetics of hybrid sterility in a line cross between two members of the Mimulus guttatus species complex (M. guttatus and M. nasutus). Hybrids showed partial male and female sterility, and the patterns of infertility were not consistent with the action of chromosomal rearrangements alone. F2 and F1 hybrids exhibited equal decreases in pollen viability (> 40%) relative to the highly fertile parental lines. A large excess of completely pollen-sterile F2 genotypes also pointed to the segregation of Dobzhansky-Muller incompatibility factors affecting male fertility. Female fertility showed a pattern similarly consistent with epistatic interactions: F2 hybrids produced far fewer seeds per flower than F1 hybrids (88.0 +/- 2.8 vs. 162.9 +/- 8.5 SE, respectively) and either parental line, and many F2 genotypes were completely female sterile. Dobzhansky-Muller interactions also resulted in the breakdown of several nonreproductive characters and appear to contribute to correlations between male and female fertility in the F2 generation. These results parallel and contrast with the genetics of postzygotic isolation in model animal systems and are a first step toward understanding the process of speciation in this well-studied group of flowering plants.

Asteraceae↗
PubMed2002

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.

Animals↗
PubMed2001

Deleterious mutations and genetic variation for flower size in Mimulus guttatus.

Mimulus guttatus is a wildflower that exhibits substantial genetic variation in flower size. Here, we test the hypothesis that this variation is caused by deleterious mutations maintained through mutation-selection balance. The deleterious-mutation model predicts that rare, partially recessive alleles will be the primary source of variation. We test this prediction by measuring the change in the mean flower size (deltaM) and the directional dominance of flower size (deltaB) within a selection experiment. If variation is due to rare (partially) recessive alleles, deltaB/deltaM is expected to be positive and exceed one. However, we obtain negative values for deltaB/deltaM from three independent selection lines. This result is statistically inconsistent with the deleterious-mutation model.

Alleles↗
PubMed1999

Stigma behavior in Mimulus aurantiacus (Scrophulariaceae).

The bilobed stigma of many species in the order Scrophulariales closes in response to touch by an animal pollinator. In hummingbird-pollinated bush monkey flower, Mimulus aurantiacus (Scrophulariaceae), closure is rapid, occurring within seconds of tactile stimulus. We investigated the proximate causes of stigma closure and subsequent reopening in M. aurantiacus, as well as potential costs and benefits of stigma closure for female fitness. Stigma closure is elicited by both touch and pollen, but closure in response to pollen is much slower, requiring 0.5-1.5 h. Stigmata reopen within 2.5-4.5 h if touch, but no pollen, is applied. Upon receipt of pollen, most stigmata remain closed for the remaining lifetime of the flower, even if less pollen is received than is needed for full seed set. Those stigmata that do reopen after pollination generally require between 20 and 28 h to do so, much longer than for unpollinated stigmata. Reopening after pollination appears to be a response to low seed set rather than to low pollen load. Natural pollination of stigmata manipulated to prevent closure shows that closure does not increase capture of pollen or seed set. In fact, closure reduces the average pollen load received by flowers. Despite this, there is no evidence that stigma closure has any negative effect on female fitness in terms of seed set or germinability. Hypotheses for the adaptive significance of stigma closure are discussed. Understanding proximate causes of stigma closure and reopening is essential in the evaluation of these hypotheses.

Journal Article↗
PubMed2006

Mating system and the evolution of quantitative traits: an experimental study of Mimulus guttatus.

The mating system of a population profoundly influences its evolution. Inbreeding alters the balance of evolutionary forces that determine the amount of genetic variation within a population. It redistributes that variation among individuals, altering heritabilities and genetic correlations. Inbreeding even changes the basic relationships between these genetic statistics and response to selection. If populations differing only in mating system are exposed to the same selection pressures, will they respond in qualitatively different ways? Here, we address this question by imposing selection on an index of two negatively correlated traits (flower size and development rate) within experimental populations that reproduce entirely by outcrossing, entirely by self-fertilizing, or by a mixture of outcrossing and selfing. Entirely selfing populations responded mainly by evolving larger flowers whereas outcrossing populations also evolved more rapid development. Divergence occurred despite an equivalent selection regime and no direct effect of mating system on fitness. The study provides an experimental demonstration of how the interaction of selection, genetic drift, and mating system can produce dramatic short-term changes in trait means, variances, and covariances.

Crosses, Genetic↗
PubMed2007

Compost-based growing media: influence on growth and nutrient use of bedding plants.

The agronomic performance and the mineral composition and trace element content in Begonia semperflorens "Bellavista F1", Mimulus "Magic x hybridus", Salvia splendens "maestro", and Tagete patula xerecta "Zenith Lemon Yellow", were tested by growing the plants on substrates of white peat and 25-50-75-100% green waste and sewage sludge (80%+20%v/v) compost (CP). A commercial peat medium of black and white peat (2:1v/v) was used as control. At flowering, the agronomic parameters were compared by ANOVA and plant nutritional status was compared by vector analysis. Substrate-species interactions (P<0.001) were evident for all measured parameters. In the 25% CP medium all the species showed an increase or preservation of the studied agronomic parameters. Begonia grown in 25% CP, showed the highest dry weight (DW) and number of flowers. Other treatments were comparable to the control. Mimulus and Salvia showed the highest DW in the 25-50% CP. Mimulus, after a DW increase up to 50% CP, showed the steepest reduction as the CP increased further. Tagete showed no differences in DW up to 50% CP, or in flower number up to 25% CP, compared to the control. The additional increases of CP in the medium showed a DW decrease similar to that of Salvia. Vector analysis showed the use of compost mainly induced a decrease of P concentration in tissues, except for Begonia which remained unchanged. Plant tissues showed a general P reduction due to a dilution effect in the low compost mixtures (25-50%) and a deficiency in the higher CP mixtures. In contrast, an increase of Mg in the aboveground tissues of all species was detectable as compost usage increased, with the exception of Salvia which suffered a Mg deficiency. Vector analysis also highlighted a Ni and partial Fe deficiency in Tagete and Salvia.

Begoniaceae↗
PubMed2006

Growth and leaf physiology of monkeyflowers with different altitude ranges.

Every species is limited both geographically and ecologically to a subset of available habitats, yet for many species the causes of distribution limits are unknown. Temperature is thought to be one of the primary determinants of species distributions along latitudinal and altitudinal gradients. This study examined leaf physiology and plant performance under contrasting temperature regimes of sister species of monkeyflower, Mimulus cardinalis and Mimulus lewisii (Phrymaceae), that differ in altitude distribution to test the hypothesis that temperature is the primary determinant of differences in fitness versus altitude. Each species attained greatest aboveground biomass, net photosynthetic rate, and effective quantum yield of photosystem II when grown under temperatures characteristic of the altitudinal range center. Although both species exhibited greater stem length, stomatal conductance, and intercellular CO2 concentration in hot than in cold temperatures, these traits showed much greater reductions under cold temperature for M. cardinalis than for M. lewisii. Survival of M. lewisii was also sensitive to temperature, showing a striking decrease in hot temperatures. Within each temperature regime, the species native to that temperature displayed greatest growth and leaf physiological capacity. Populations from the altitude range center and range margin of each species were used to examine population differentiation, but central and marginal populations did not differ in most growth or leaf physiological responses to temperature. This study provides evidence that M. cardinalis and M. lewisii differ in survival, growth, and leaf physiology under temperature regimes characterizing their contrasting low and high altitude range centers, and suggests that the species' altitude range limits may arise, in part, due to metabolic limitations on growth that ultimately decrease survival and limit reproduction.

Altitude↗
PubMed2026

Climate Gradients and Habitat Discontinuity Structure Genetic Variation in a Spring-Specialist Plant.

BACKGROUND AND AIMS: Groundwater-dependent ecosystems support disproportionate biodiversity in arid regions, yet the population genetics of spring-specialist plants remains poorly understood. Here, we present the first species-wide genetic dataset for crimson monkeyflower (Mimulus verbenaceus, Phrymaceae), a spring-specialist plant distributed in seeps, springs, and associated riparian areas across desert regions of North America. We aim to relate landscape features and climate gradients to the spatial genetic structuring within this system. METHODS: Using genome-wide reduced representation sequencing data consisting of 10,760 SNPs from 175 individuals across 17 populations, we characterized the patterns of genetic differentiation and diversity. Population structure was assessed using ADMIXTURE and Principal Component Analysis. We examined the contributions of climate to range-wide genetic variation in crimson monkeyflower using redundancy analysis. KEY RESULTS: Patterns of genetic differentiation were more consistent with those of spring-specialist animal taxa than those of upland plants or generalist riparian plants. We found strong population structure at both broad regional scales and at fine local scales. While geographic and spatial structuring was a primary driver of genetic structure across all scales, riparian connectivity influenced local patterns of diversity, and adaptation to local climatic variation was more influential at regional scales, with temperature, relative humidity, and a monsoon-driven climate gradient contributing to genetic differentiation. CONCLUSIONS: Our findings highlight the distinctive association with isolated perennial groundwater sources, as well as climate gradients, with genetic variation in this spring-specialist plant. These findings suggest that spring-specialist plants deserve special consideration in ecological theory, management, and conservation.

Mimulus↗
PubMed2003

Allele substitution at a flower colour locus produces a pollinator shift in monkeyflowers.

The role of major mutations in adaptive evolution has been debated for more than a century. The classical view is that adaptive mutations are nearly infinite in number with infinitesimally small phenotypic effect, but recent theory suggests otherwise. To provide empirical estimates of the magnitude of adaptive mutations in wild plants, we conducted field studies to determine the adaptive value of alternative alleles at a single locus, YELLOW UPPER (YUP). YUP controls the presence or absence of yellow carotenoid pigments in the petals of pink-flowered Mimulus lewisii, which is pollinated by bumblebees, and its red-flowered sister species M. cardinalis, which is pollinated by hummingbirds. We bred near-isogenic lines (NILs) in which the YUP allele from each species was substituted into the other. M. cardinalis NILs with the M. lewisii YUP allele had dark pink flowers and received 74-fold more bee visits than the wild type, whereas M. lewisii NILs with the M. cardinalis yup allele had yellow-orange flowers and received 68-fold more hummingbird visits than the wild type. These results indicate that an adaptive shift in pollinator preference may be initiated by a single major mutation.

Adaptation, Physiological↗
PubMed2004

Predicting evolution of floral traits associated with mating system in a natural plant population.

Evolution of floral traits requires that they are heritable, that they affect fitness, and that they are not constrained by genetic correlations. These prerequisites have only rarely been examined in natural populations. For Mimulus guttatus, we found by using the Riska-method that corolla width, anther length, ovary length and number of red dots on the corolla were heritable in a natural population. Seed production (maternal fitness) was directly positively affected by corolla width and anther size, and indirectly so by ovary length and number of red dots on the corolla. The siring success (paternal fitness), as estimated from allozyme data, was directly negatively affected by anther-stigma separation, and indirectly so by the corolla length-width ratio. Genetic correlations, estimated with the Lynch-method, were positive between floral size measures. We predict that larger flowers with larger reproductive organs, which generally favour outcrossing, will evolve in this natural population of M. guttatus.

Biological Evolution↗
PubMed2005

Epistasis in monkeyflowers.

Epistasis contributes significantly to intrapopulation variation in floral morphology, development time, and male fitness components of Mimulus guttatus. This is demonstrated with a replicated line-cross experiment involving slightly over 7000 plants. The line-cross methodology is based on estimates for means. It thus has greater power than the variance partitioning approaches historically used to estimate epistasis within populations. The replication of the breeding design across many pairs of randomly extracted, inbred lines is necessary given the diversity of multilocus genotypes residing within an outbred deme. Male fitness is shown to exhibit synergistic epistasis, an accelerating decline in fitness with inbreeding. Synergism is a necessary, but not sufficient, condition for a mutational deterministic hypothesis for the evolutionary maintenance of sexual reproduction. Unlike male fitness measures, flower morphology and development time yield positive evidence of epistasis but not of synergism. The results for these traits suggest that epistatic effects are variable across genetic backgrounds or sets of interacting loci.

Crosses, Genetic↗