COVARIATION OF SELFING RATES WITH PARENTAL GENE FIXATION INDICES WITHIN POPULATIONS OF MIMULUS GUTTATUS

Wright's gene fixation index F and two single-locus effective selfing rates—the selfing rate at loci with fixed alleles, and the selfing rate at loci without fixed alleles—were estimated in five populations of Mimulus guttatus. These two effective selfing rates describe the inbreeding observed at a single locus when both uniparental and biparental inbreeding are practiced. Estimates were made using progeny arrays assayed for six allozyme loci and two morphological loci exhibiting dominance. The average of the two selfing rates computed for subpopulations (ca. 10 m diameter) ranged from 24% to 59%, with a mean of 37%. When computed for populations (ca. 1 km diameter), average selfing rates were about 10% higher. In four populations, the selfing rate at loci with fixed alleles was higher than the selfing rate at loci without fixed alleles. Thus, the covariance of selfing with parental gene fixation was positive. In one of the populations, estimates for individual plants sampled along a transect gave positive correlations for selfing rates and for gene-fixation indices between adjacent plants. A highly positive correlation between selfing rate and gene fixation of individual plants was also observed. In another population, the covariance of selfing with gene fixation was higher for a locus causing leaf spots than for allozyme loci. This covariance is partially caused by 1) variation in homozygosity among neighborhoods and 2) biparental inbreeding within neighborhoods. The consequences of this covariance are discussed.     

EFFECTS OF DIFFERENT LEVELS OF INBREEDING ON FITNESS COMPONENTS IN MIMULUS GUTTATUS

Recent theoretical work has shown that there can be selection favoring the maintenance of sexual reproduction and the evolution of increased recombination when deleterious mutations at different loci interact synergistically, such that the logarithm of fitness declines at a greater than linear rate with the number of harmful mutations per genome. The purpose of this experimental study was to determine whether synergism exists for genes affecting fitness components in two partially selfing populations of the monkey flower Mimulus guttatus. For each wild population, a large randomly mated base population was constructed and many independent lines, inbred to differing degrees, were extracted from this base population. Lines with expected inbreeding coefficients of 0, 0.25, 0.5, and 0.75 were raised simultaneously in the greenhouse and were scored for germination, flowering, flower production, and pollen viability. All fitness traits except germination success declined with increased inbreeding, but in spite of the substantial inbreeding depression found in this study, relatively little evidence of synergistic epistasis was found. The only trait that showed evidence of synergism was pollen viability. These results indicate that synergism is not strong for the fitness components measured in this study. The evidence for synergism from other published studies is also reviewed.     

DEVELOPMENTAL SELECTION IN RESPONSE TO ENVIRONMENTAL CONDITIONS OF THE MATERNAL PARENT IN MIMULUS GUTTATUS

Copper tolerance is expressed in the diploid sporophyte as well as the microgametophyte of Mimulus guttatus. Previous studies, based on reproductive output, suggested that selection for copper tolerance could occur within the pistil. The objective of this study was to determine if selection within the pistil can increase sporophytic tolerance to copper and to determine whether this selection occurs pre- or postzygotically. Mixtures of pollen from copper tolerant or sensitive sources or from plants heterozygous for tolerance to copper were applied at two intensities to plants cloned and grown in control or copper supplemented solutions. The proportion of copper tolerant progeny showed a small, 7%, but significant increase when pollen recipients were grown with added copper. Comparisons of the numbers of tolerant progeny, as well as various components of reproduction, following light and heavy pollinations suggested that microgametophytic selection was unlikely to account for this increase. However, the 8 to 10% decrease in the seed/zygote ratio, compared to control values, was sufficient to account for the difference in proportion of copper tolerant progeny from control and copper treated plants. Thus, it appeared likely that selection for copper tolerance could occur within the pistil, and that much of this selection occurred postzygotically through the early failure of developing seeds.     

CORRELATED MATINGS IN THE PARTIAL SELFER MIMULUS GUTTATUS

In partially selfing populations, siblings may be correlated for both selling and paternity. A model of the mating system based upon sampling pairs of progeny from a maternal parent is described. The model separates the correlation of selfing from the correlation of outcrossed paternal alleles and is an approach to paternity analysis suited for larger populations with fewer marker loci. Its parameters determine the components of genetic covariance between sibs and provide information about the average number of fathers in a maternal sibship. Electrophoretic markers were used to obtain estimates of correlated matings for two Mimulus guttatus populations. In both populations, about 50% selfing was observed. For two sibs randomly selected from the same capsule, the correlations of selfing between these sibs were 17% and 12% in the two populations, and the correlations of paternity (the proportion of full-sibs among outcrossed sib-pairs) were 37% and 44%. Sibs from different capsules were not correlated for selfing, and the paternity correlation dropped to near 20% in both populations. However, estimates of correlated matings have high variance, lack statistical independence, and can be difficult to obtain. The use of marker loci with many alleles can alleviate these problems.     

ROLE OF COROLLA ABSCISSION IN DELAYED SELF-POLLINATION OF MIMULUS GUTTATUS (SCROPHULARIACEAE)

Corollas of Mimulus guttatus are shed 1–9 days post-anthesis, and autofertility is often high. The possibility that corolla shedding causes self-pollination in the absence of pollinators was examined. In one experimental population 82% of seed produced under conditions of pollinator exclusion were due to corolla abscission. Corolla abscission, in the absence of pollinators, is a mechanism of delayed self-fertilization in this taxon.     

REPRODUCTIVE ASSURANCE MECHANISMS IN THREE TAXA OF THE MIMULUS GUTTATUS COMPLEX (SCROPHULARIACEAE)

The evolution of inbreeding in plants has often been attributed to selection for the ability to set seed in the absence of mates or pollinators. Mechanisms of reproductive assurance in five populations of mixed mating Mimulus guttatus, three populations of inbreeding M. platycalyx, and two populations of inbreeding M. nasutus were examined in a pollinator-free greenhouse. Reproductive assurance was manifested in all populations by autofertility, vegetative reproduction, or both. The inbreeding taxa had significantly greater levels of autofertility and less vegetative reproduction. Three modes of autofertility were identified: 1) due to corolla abscission only, occurring in three M. guttatus populations; 2) due to both corolla abscission and direct anther-stigma contact by curling of the lower stigmatic lobe into the anthers, occurring in two M. guttatus populations; and 3) direct stigma-anther contact by stigma curling alone prior to corolla abscission, found in each M. platycalyx and M. nasutus population. Stigma-anther distance and its interaction with stigma curling contributed to differences in autofertility among populations. Significant levels of intrapopulation quantitative genetic variation were found for seven of ten traits examined; average levels were similar between inbreeding and mixed mating populations. Genetic variation within populations for autofertility per se was not detected, but significant levels controlling stigma-anther distance were found in two M. guttatus populations. These results show that evolution of inbreeding by natural selection for reproductive assurance is possible in Mimulus, and illustrate the complex changes in floral dynamics and morphology it may involve.     

DIFFERENTIAL SEED PRODUCTION IN MIMULUS GUTTATUS IN RESPONSE TO INCREASING CONCENTRATIONS OF COPPER IN THE PISTIL BY POLLEN FROM COPPER TOLERANT AND SENSITIVE SOURCES

Selection can occur in the pistil, during a series of stages that include both pre-zygotic (pollen germination, pollen tube growth, and fertilization) and post-zygotic events. This study explores the extent to which selection, at this level, could be due to the environmental conditions under which the maternal parent is growing. Five plants of Mimulus guttatus, tolerant to copper, were vegetatively cloned and each clone was grown in control and in solutions to which copper was added. The maternal plants received pollen from plants either tolerant or sensitive to copper. Seeds and ovules were counted to estimate the number of seeds/capsule, the seed/ovule ratio, the percent fertilization, and the proportion of zygotes aborting for each clone, treatment and pollen source combination. There were large differences among the pollen recipients for each of the measurements. However, there was a consistent pattern to seed production depending on the pollen source and copper treatment. The seed/ovule ratio was unaffected if pollen came from tolerant sources but was reduced by an average of 24% for both copper supplemented treatments if pollen came from copper sensitive sources. Thus, the data indicated that selection due to environmental factors could occur within the pistil. Differences in percent fertilization were not statistically significant, but the seed/zygote ratio showed a pattern that was similar to seed production suggesting that abortion of immature seeds was responsible for most of the difference in seed production.     

THE STRUCTURE OF SOME CYTOPLASMIC COMPONENTS OF PLANT CELLS IN RELATION TO THE BIOCHEMICAL PROPERTIES OF ISOLATED PARTICLES

1. Electron micrographs of thin sections of material fixed with buffered osmium tetroxide have been used for comparison of the fine structure of isolated cytoplasmic particles from silver beet petioles and roots of germinating wheat with that of the cytoplasm of the intact cells. 2. Mitochondria of wheat roots have an external double membrane and poorly oriented internal double membranes. As compared with the structures seen in situ, the isolated mitochondria showed evidence of some disorganisation of the fine internal structure, probably due to osmotic effects. The possible influence of such changes on the enzymic properties of the isolated mitochondria is discussed. 3. The isolated plant microsomes are mainly spherical vesicular structures consisting of (a) an outer membrane enclosing (b) either an homogeneous slightly dense material (wheat root microsomes) or some granular dense material (silver beet microsomes) and (c) small dense particles, mostly associated with the vesicle membranes. 4. The cytoplasm of the wheat root cells does not contain any structures similar to the isolated microsomes but has a very dense reticular network, consisting of membranes with associated small dense particles, here called the endoplasmic reticulum. The observations indicate that the isolated microsomes arise mainly by rupture and transformation of the membranes of this structure. The effects of such extensive changes in the lipoprotein membranes on the enzymic activities of the endoplasmic reticulum, as studied in isolated microsomes, is discussed. 5. Meristematic wheat root cells contain structures which consist of smooth membranes with associated vacuoles and are similar to the Golgi zones of animal cells. The membranes of these zones probably contribute to the microsomal fraction under the conditions of preparation used for the enzymic and chemical studies previously reported.     

INFERENCES ABOUT QUANTITATIVE INHERITANCE BASED ON NATURAL POPULATION STRUCTURE IN THE YELLOW MONKEYFLOWER,MIMULUS GUTTATUS

We used a nonmanipulative, marker-based method to study quantitative genetic inheritance in two habitats of a common monkeyflower population. The method involved regressing quantitative trait similarity on marker-estimated relatedness between individuals sampled in the field. We sampled 300 adult plants from each of two transects, one along a stream habitat and another through a meadow habitat. For each plant we measured 10 quantitative characters and assayed 10 polymorphic isozyme loci. In the meadow habitat, relatedness of plants within 1 m was moderate (r = 0.125, corresponding to half-sibs) as was actual variance of relatedness (V_r = 0.044). Significant heritabilities of 50–70% were found for corolla width and the fitness characters of flower number and plant weight. Genetic correlations were strongly positive, but sharing of environmental effects within 1 m was weak. In the stream habitat, levels of relatedness were lower and similar heritabilities were indicated. To detect dominance variance and the correlation of phenotypes due to shared inbreeding, we also estimated higher-order coefficients of relationship and inbreeding, but these did not significantly differ from zero. Laboratory-based estimates of heritability in the field were lower than the marker-based estimates, indicating that natural heritabilities and genetic correlations may be stronger than indicated by controlled studies.