POSTPOLLINATION EFFECTS ON SEED PATERNITY: MECHANISMS IN ADDITION TO MICROGAMETOPHYTE COMPETITION OPERATE IN WILD RADISH

After pollen arrives on a stigma, the paternity of seeds may be influenced by microgametophyte competition, maternal choice, genetic complementation between parents, and embryo competition. While microgametophyte competition has been well accepted, the other mechanisms are more difficult to demonstrate, and their effects are often confounded. Here, wild radish plants were pollinated with single and mixed pollen loads, and some plants were stressed such that reproduction was reduced. Effects of pollen donors, maternal families, maternal × paternal interaction, pollen donor number, and stress on fruit abortion, seed number per fruit, seed weight, and total seed weight per fruit were measured. Maternal-plant × pollen-donor interaction effects were found for all variables, indicating that genetic complementation or maternal choice occurred. Values of the components of reproduction were generally higher for multiply sired fruits than for singly sired fruits, indicating that either competition among embryos changed under multiple paternity or maternal choice for multiply sired fruits occurred. Finally, when maternal plants were stressed, the components of reproduction were more strongly affected by seed and fruit paternity. This result indicates that either competitive regimes among embryos were affected by stress or maternal plants become more selective under stress. In both cases where embryo competition might have been an explanation of the results, variation in seed weight within fruits was unaffected, suggesting that competitive regimes were unchanged. Clearly, mechanisms in addition to microgametophyte competition are important in sorting the pollen that arrives on stigmas of wild radish. These data suggest that maternal choice is likely to be important. In addition, these processes are likely to occur in the field, since the effects are stronger in stressed than in control plants.     

PARENTAL EFFECTS ON SEED DEVELOPMENT AND SEED YIELD IN RAPHANUS RAPHANISTRUM: IMPLICATIONS FOR NATURAL AND SEXUAL SELECTION

The possibility that sexual selection operates in angiosperms to effect evolutionary change in polygenic traits affecting male reproductive success requires that there is additive genetic variance for these traits. I applied a half-sib breeding design to individuals of the annual, hermaphroditic angiosperm, wild radish (Raphanus raphanistrum: Brassicaceae), to estimate paternal genetic effects on, or, when possible, the narrow-sense heritability of several quantitative traits influencing male reproductive success. In spite of significant differences among pollen donors with respect to in vitro pollen tube growth rates, I detected no significant additive genetic variance in male performance with respect to the proportion of ovules fertilized, early ovule growth, the number of seeds per fruit, or mean individual seed weight per fruit. In all cases, differences among maternal plants in these traits far exceeded differences among pollen donors. Abortion rates of pollinated flowers and fertilized ovules also differed more among individuals as maternal plants than as pollen donors, suggesting strong maternal control over these processes. Significant maternal phenotypic effects in the absence of paternal genetic or phenotypic effects on reproductive traits may be due to maternal environmental effects, to non-nuclear or non-additive maternal genetic effects, or to additive genetic variance in maternal control over offspring development, independent of offspring genotype. While I could not distinguish among these alternatives, it is clear that, in wild radish, the opportunity for natural or sexual selection to effect change in seed weight or seed number per fruit appears to be greater through differences in female performance than through differences in male performance.     

Effects of parentage,prior fruit set and pollen load on fruit and seed production in Campanula americana L

Summary We conducted a controlled crossing experiment to examine the effects of maternal and paternal parentage, the size of the pollen load, and prior fruit production on the proportion of flowers that set fruit, seed number per fruit and seed weight in a natural population of Campanula americana. Effects due to the maternal parent were large for all measures of fruit and seed production, while the paternal parent had a significant effect only upon mean seed weight. As the number of prior fruits on the maternal plant increased the probability that a flower would produce a mature fruit, the number of seeds per fruit, and total seed weight per fruit all decreased. We found no effect of the size of the pollen loads used in this study on fruit or seed production. These results are consistent with those of other studies that suggest in natural plant populations maternal effects, especially environmental maternal effects, can have an overwhelming effect on fruit and seed production and on seed characteristics.     

Soybean Seed Water Relations during in Situ and in Vitro Growth and Maturation

Water, osmotic, and pressure potentials of soybean (Glycine max [L.] Merrill) embryos and related maternal tissues were measured during periods of seed growth and maturation to test the involvement of embryo water relations in seed maturation. Seeds were matured in situ or in an in vitro liquid culture medium in detached pods or as isolated seeds. Changes in water relations of embryo tissues were independent of maternal tissues. During seed maturation in situ, water and osmotic potentials in both embryo and maternal tissues declined sharply near the time of maximum dry weight. During in vitro seed culture with and without pods, water and osmotic potentials in axis and cotyledon tissues declined continuously during growth. Water and osmotic potentials of the seed coat, which was present only during in vitro seed culture with pods, changed little during the culture period. Positive turgor in the embryo was maintained beyond maximum dry weight and the loss of green color during in vitro culture but declined to zero at maturity in situ. The osmotic potential in embryo tissues declined from −1.1 megapascals at early pod fill to between −1.65 and −2.2 megapascals at maximum seed dry weight across all maturation environments. It is suggested that the decreasing osmotic potential in the growing soybean embryo reaches a threshold level that is associated with cessation of growth and onset of seed maturation.     

FERTILIZATION DYNAMICS AND PARENTAL EFFECTS UPON FRUIT DEVELOPMENT IN RAPHANUS RAPHANISTRUM: CONSEQUENCES FOR SEED SIZE VARIATION

Seed weight varies significantly within and among fruits of wild radish (Raphanus raphanistrum). To determine sources of this variation, we studied fertilization and seed development following controlled pollinations. Within fruits, central ovules were fertilized prior to distal ovules and attained greater seed size. Ninety-seven percent of the variation in mean seed wt per fruit was explained by an analysis of variance incorporating parental effects, pollination date, and the number of seeds per fruit. We document strong maternal effects on the number of ovules per ovary, the number of fertilized ovules per ovary, the number of seeds per fruit, and mean individual seed wt per fruit. Across females, pollen donor had a slight but significant effect on seed wt; no paternal effects on fertilization rate, zygote number, or seed number per fruit were detected. Within females, with one exception, pollen donor had no significant effect on these components of seed development. Stronger maternal main effects may be due to donor x recipient interactions, cytoplasmic factors, the genetic inequity within triploid endosperm, and/or strict maternal control over resource allocation. The large maternal effects relative to paternal effects may limit the rate at which natural selection acts on paternal traits expressed prior to seed maturation.     

Dynamics of maternal and paternal effects on embryo and seed development in wild radish (Raphanus sativus)

Background and Aims

Variability in embryo development can influence the rate of seed maturation and seed size, which may have an impact on offspring fitness. While it is expected that embryo development will be under maternal control, more controversial hypotheses suggest that the pollen donor and the embryo itself may influence development. These latter possibilities are, however, poorly studied. Characteristics of 10-d-old embryos and seeds of wild radish (Raphanus sativus) were examined to address: (a) the effects of maternal plant and pollen donor on development; (b) the effects of earlier reproductive events (pollen tube growth and fertilization) on embryos and seeds, and the influence of embryo size on mature seed mass; (c) the effect of water stress on embryos and seeds; (d) the effect of stress on correlations of embryo and seed characteristics with earlier and later reproductive events and stages; and (e) changes in maternal and paternal effects on embryo and seed characteristics during development.

Methods

Eight maternal plants (two each from four families) and four pollen donors were crossed and developing gynoecia were collected at 10 d post-pollination. Half of the maternal plants experienced water stress. Characteristics of embryos and seeds were summarized and also compared with earlier and later developmental stages.

Key Results

In addition to the expected effects of the maternal plants, all embryo characters differed among pollen donors. Paternal effects varied over time, suggesting that there are windows of opportunity for pollen donors to influence embryo development. Water-stress treatment altered embryo characteristics; embryos were smaller and less developed. In addition, correlations of embryo characteristics with earlier and later stages changed dramatically with water stress.

Conclusions

The expected maternal effects on embryo development were observed, but there was also evidence for an early paternal role. The relative effects of these controls may change over time. Thus, there may be times in development when selection on the maternal, paternal or embryo contributions to development are more and less likely.     

EFFECTS OF PARENTAGE AND SIZE OF THE POLLEN LOAD ON PROGENY PERFORMANCE IN CAMPANULA AMERICANA

To examine the effects of maternal and paternal parentage and the size of the pollen load on seed size and weight and on progeny performance we conducted a controlled crossing experiment using a natural population of Campanula americana. We found that seed size was positively correlated with early seedling performance for all but one of traits we measured (days to emergence), but was not significantly correlated with any of the later vegetative measures or reproductive output. We detected significant effects due to the maternal parent for the vegetative traits days to emergence, days to first leaf, and final plant height, as well as total seed weight, and mean seed weight per fruit. Significant paternal effects were found for all of the seedling traits except number of leaves after vernalization. The progeny from fruits receiving high pollen loads significantly outperformed the progeny from fruits receiving low pollen loads for the traits days to first and second leaf, numbers of leaves after vernalization, and days to first flower. These results not only demonstrate the importance of parentage and seed weight on progeny performance, but also indicate that variations in the size of the pollen load may be important in seedling establishment in natural populations.     

EFFECTS OF POLLEN DONOR IDENTITY ON OFFSPRING QUALITY IN WILD RADISH,RAPHANUS SATIVUS

If pollen donor performance during mating correlates with differences in offspring growth and fitness, processes that sort among potential mates may directly improve offspring fitness. Here seeds sired by three pollen donors on ten maternal plants were grown for eight weeks in the greenhouse. The performance of the pollen donors during pollination and fertilization was known from a previous experiment. There were significant effects of paternity on two measures of early growth: leaf number and plant height. Paternal effects on three measures more closely related to fitness; final plant weight, day of first flower production, and total flower number were also significant. Under the conditions of this experiment, final plant weight was probably the best predictor of fitness. The pollen donor that sired the largest seeds in the previous experiment sired offspring that were largest after 8 weeks of growth. Half of the plants were grown under low-water conditions. Paternal effects on growth were not masked by the environmental effects. In fact, some paternal effects became stronger under stress. This suggests that paternal effects could also be important in the field. Plants sired by donor A bolted very early when water was limited and would probably have an advantage in a season that was very short due to an early and severe drought. During fertilization and seed filling, seeds sired by this donor were more frequent on water-stressed maternal plants than on control maternal plants (Marshall, 1988). The data from this experiment indicate a connection between pollen donor performance during mating and offspring growth. These results suggest that the processes that sort among potential fathers during pollination, fertilization, and seed filling, may improve offspring quality.     

Imprinting of the Polycomb Group Gene MEDEA Serves as a Ploidy Sensor in Arabidopsis

Balanced maternal and paternal genome contributions are a requirement for successful seed development. Unbalanced contributions often cause seed abortion, a phenomenon that has been termed “triploid block.” Misregulation of imprinted regulatory genes has been proposed to be the underlying cause for abnormalities in growth and structure of the endosperm in seeds with deviating parental contributions. We identified a mutant forming unreduced pollen that enabled us to investigate direct effects of unbalanced parental genome contributions on seed development and to reveal the underlying molecular mechanism of dosage sensitivity. We provide evidence that parent-of-origin–specific expression of the Polycomb group (PcG) gene MEDEA is causally responsible for seed developmental aberrations in Arabidopsis seeds with increased paternal genome contributions. We propose that imprinted expression of PcG genes is an evolutionary conserved mechanism to balance parental genome contributions in embryo nourishing tissues.     

Paternal transmission of a seed size reduction gene varies with age of a primary transformant and seed set is also influenced by gene expression in maternal tissues

We have constructed chimaeric genes that are expressed in embryo and endosperm compartments of the seed, induce dominant seed lethality and have potential to reduce seed size in 75% of seeds within a fruit such as Citrus [7]. The genes are not entirely seed-specific as a proportion of primary test tobacco transformants containing their gene were fully male-sterile [7]. Here we investigated why a proportion of apparently male-fertile transgenic plants showed segregation distortion from the 75% seed lethality expected for a single dominant gene. Reciprocal crosses were conducted between pollen fertile, primary tobacco transformants containing various copies of the CG1-400-RNase gene [7] and wild-type tobacco plants to examine the transmission of the gene through maternal and paternal gametes and also the effects of gene dosage in embryo and endosperm compartments on seed viability and phenotype. Pollen viability, seed set and seed phenotype were examined over a 16 month period to assess stability of gene expression in primary transformants because woody, fruit crops containing these genes will be vegetatively propagated from primary transformants. In male-fertile transformants, the gene was observed to be expressed to varying degrees post-meiotically in pollen over the time period examined resulting in lethality of transgenic pollen and reduced paternal transmission. A variable, low-level maternal expression component was also detected that resulted in seed lethality and influenced morphological variation in the seed lethal phenotype. The maternal and paternal expression components caused seed lethality to range from 50 to 75%. This study indicates the need to select for transformants with stable pollen transmission and high seed expression and raises questions in relation to possible environmental and epistatic effects on gene expression in primary, hemizygous transformants over long growth periods.     

DNA methylation causes predominant maternal controls of plant embryo growth

The parental conflict hypothesis predicts that the mother inhibits embryo growth counteracting growth enhancement by the father. In plants the DNA methyltransferase MET1 is a central regulator of parentally imprinted genes that affect seed growth. However the relation between the role of MET1 in imprinting and its control of seed size has remained unclear. Here we combine cytological, genetic and statistical analyses to study the effect of MET1 on seed growth. We show that the loss of MET1 during male gametogenesis causes a reduction of seed size, presumably linked to silencing of the paternal allele of growth enhancers in the endosperm, which nurtures the embryo. However, we find no evidence for a similar role of MET1 during female gametogenesis. Rather, the reduction of MET1 dosage in the maternal somatic tissues causes seed size increase. MET1 inhibits seed growth by restricting cell division and elongation in the maternal integuments that surround the seed. Our data demonstrate new controls of seed growth linked to the mode of reproduction typical of flowering plants. We conclude that the regulation of embryo growth by MET1 results from a combination of predominant maternal controls, and that DNA methylation maintained by MET1 does not orchestrate a parental conflict.     

SEED ABORTION AND SEED SIZE VARIATION WITHIN FRUITS OF PHASEOLUS VULGARIS: POLLEN DONOR AND RESOURCE LIMITATION EFFECTS

A new plant cohort starts with the formation of zygotes at ovule fertilization. This paper presents an ecological study of the effects of resource limitation and pollen donor on the survivorship and size of plant embryos in maturing fruits of the common bean, Phaseolus vulgaris L. For naturally selfed flowers of domesticated and wild beans, embryos in basal ovular positions were more likely to abort or, if they survived, become lighter seeds than stylar embryos. Embryo survivorship was higher for the cultivar when maternal plants grew in larger pots. Early seed abortion in the cultivar was random with respect to ovular position. In late abortion basal embryos had a higher mortality than stylar ones. The basal embryos lacked gross genetic defects since they developed adult organs in embryo culture. In more outbred crosses with the cultivar the position effect in embryo survivorship and size disappeared. This difference among paternal parents implicates a genetic component to the observed position effects. The variation in mature seed size may affect adult characteristics since large seeds produced larger juvenile plants than small seeds.     

DEVELOPMENTAL AND GENETIC SOURCES OF SEED WEIGHT VARIATION IN RAPHANUS RAPHANISTRUM L. (BRASSICACEAE)

Seed weight is known to have a marked impact on emergence and post-emergence productivity in wild radish (Raphanus raphanistrum). In this paper, I describe several levels of seed weight variation in plants taken from a natural population in Hamden, Connecticut. Six maternal plants from the 1981 season were analyzed in detail: the weights and positions of all seeds within a fruit were recorded, and some of these seeds were used the following summer for competition studies and progeny analysis. Within a plant, average seed weight decreased as the number of seeds within a fruit increased, suggesting that developing embryos compete for maternal resources. Seed weight also varied significantly among the six maternal plants used in the study. Comparison of the average weights of seeds produced by offspring of those six plants with the average weights of seeds borne by the maternal plant revealed a significant genetic component to seed weight variation. Seed weight varied up to six-fold within single fruits of R. raphanistrum; large seeds tend to occur near the pedicel or in the middle positions. Seed size variation seen within single fruits is of sufficient magnitude to result in differential reproductive output among closely related seeds under competitive field conditions.     

PLANTING DENSITY INFLUENCES THE EXPRESSION OF GENETIC VARIATION IN SEED MASS IN WILD RADISH (RAPHANUS SATIVUS L.: BRASSICACEAE)

To determine the effects of density, genotype, and their interaction on individual seed mass in Raphanus sativus L., we replicated maternal and paternal families of seed across two planting densities in an experimental garden. Seeds were produced by a nested breeding design performed in the greenhouse. Among garden-raised plants, density had a strong negative effect on the mass of seeds produced. At low density, the identity of the greenhouse-grown maternal plants had a strong effect on F₂ seed mass, while in high-density plots, there were no significant parental effects on mean seed mass. Significant parental genotype density interactions contributed to variation in F₂ seed mass. Norms of reaction for each of the 15 paternal sibships illustrate paternal family density interactions. Three sibships exhibited significant declines in mean seed mass with increasing density; 12 sibships showed no change. Maternal family density interaction effects on seed mass were also detected; among maternal sibships, mean seed mass at low density was negatively correlated with mean seed mass at high density. These results demonstrate: a) planting density has a strong effect on mean individual seed mass produced by adults; b) density influences the magnitude of maternal effects on progeny phenotype; and c) genotype density interactions influence seed mass, potentially contributing to the maintenance of maternal genetic variation in seed mass in natural populations of wild radish.     

POLLEN AND OVULE SOURCES AFFECT SEED PRODUCTION OF LOBELIA CARDINALIS (LOBELIACE AE)

To examine the breeding system and components of male and female reproductive success in the hermaphroditic plant Lobelia cardinalis, we performed three crossing experiments with plants taken from natural populations. The experiments were designed to determine if the crossing success of plants as pollen and ovule parents was affected by the distance among mates, including self-pollinations and pollinations between populations; to determine if plants differed in their abilities to sire or mature seed; and to determine if there was a correlation between a plant's success at siring and maturing seed. Selfpollinations resulted in significantly fewer seeds per fruit and significantly smaller seeds. There were no significant differences in germinability between selfed and outcrossed seeds. Distance among parents within a population did not affect any of the traits. Outcrosses within and between population produced similar numbers of seeds per fruit, similar seed weights, and similar germination success. There were highly significant differences among maternal plants in all three experiments in the number of seeds they matured, mean seed weight, and seed germinability. The maternal parent was the most important factor determining seed production, but there were also significant differences among paternal plants in the number of seeds they sired (all three experiments), in the germinability of the seeds they sired (two experiments), and in the sizes of seeds they sired (one experiment). Our results indicate that differences in success of Lobelia plants as male parents cannot be due solely to their relatedness to the female parent.     

Developmental Genetic Analysis of Seed Size in Soybean (Glycine max)

Seed size is one of the important factors of soybean [Glycine max (L.) Merrill] yield. There have been lots of reports about genetic effects and physiology—ecological researches on seed size, but the genetic behaviors of genes during seeds development were rarely discussed. Analysis of main genetic effects for fresh seed size (FSS) and dry seed size (DSS) of soybean was conducted with diallel cross data by using a seed genetic model. Analyses of unconditional and conditional variances and correlations were used to evaluate the developmental behavior of soybean. The phenotypic means of FSS and DSS in soybean at eight stages among three generations reached the highest value at 9/6 and 9/13, respectively. The means of FSS decreased dramatically after 9/6, but the means of DSS maintained relatively stable tendency at corresponding periods. The unconditional variance analysis showed that FSS and DSS were controlled by embryo, cytoplasmic and maternal effects in the whole growth period. Genetic effects due to cytoplasmic and maternal effects were relatively important for FSS and DSS at most of the growth periods. Conditional variance analysis showed that genes from different genetic system expressed discontinuously in the whole growth period. The net genetic effects due to cytoplasmic and maternal plant on FSS and DSS were larger than those of embryo effects at most of the growth periods. Different genetic system can affect the relationship of various stages to mature solely or simultaneously. Embryo additive effects at 8/16, embryo dominance effects at 8/9 and 8/16, maternal plant dominance effects at 8/2 and 8/16 could ultimately affect the performance of FSS at maturing stage. Embryo additive effects at 8/2 and 9/13, cytoplasm effects at 8/9, maternal plant dominance effects at 8/2 could ultimately affect the performance of DSS.     

大豆籽粒大小的发育遗传分析

籽粒大小是大豆产量的一个重要因素。有关大豆籽粒的遗传学和生理生态学研究已有一些研究,而对于籽粒发育过程中的遗传效应却报道很少。文章采用种子广义遗传模型,分析了大豆双列杂交组合3个世代遗传材料8个时期的鲜籽粒大小和干籽粒大小的数据,应用非条件和条件遗传方差及相关方法分析了发育遗传规律。8个时期的亲本、F1、F2的鲜籽粒大小和干籽粒大小的平均数分别在9／6和9／13达到最高值,鲜籽粒大小在9／6后迅速下降,干籽粒大小在9／13后区于稳定。非条件方差分析表明在整个生育期中,胚遗传效应、细胞质遗传效应和母体植株遗传效应对大豆鲜籽粒大小和干籽粒大小有影响。在多数生育阶段中,细胞质和母体植株的遗传效应对鲜籽粒大小和干籽粒大小影响较大。条件方差分析表明,在大豆生育期中,各遗传体系的基因间断性表达。在多数生育阶段中,细胞质和母体植株的净遗传效应高于胚净遗传效应。不同时期的各遗传体系的基因效应可以单独或同时影响鲜籽粒大小和干籽粒大小的最终表现。8／16的胚加性效应、8／9和8／16的胚显性效应、8／2和8／16的母体植株显性效应影响到鲜籽粒大小的最终表现。8／2和9／13的胚加性效应、8／9的细胞质效应、8／2的母体植株显性效应对干籽粒大小的最终表现有影响。     

TEMPORAL VARIATION OF FLOWER AND FRUIT SIZE IN RELATION TO SEED YIELD IN CELANDINE POPPY (CHELIDONIUM MAJUS; PAPAVERACEAE)

Ecological and evolutionary studies typically consider variation in single reproductive characters in isolation, without considering how they might be correlated with other reproductive and vegetative characters. In our study, we examined temporal patterns of variation and correlation in flower diameter and fruit length during a reproductive phase in two Massachusetts populations of the herb, Chelidonium majus. We also examined the relationships of such variation to measurements of seed yield components (mean seed weight and number per fruit) and aspects of plant vegetative size. Most of the variation in the sizes of reproductive characters occurred within individual plants, instead of among plants or between populations. Flower and fruit sizes as well as seed number per fruit declined significantly during the season in both populations. Only mean seed size per fruit was relatively stable for individual plants in both populations. Conserving resources by a gradual reduction in the size of reproductive characters over the season may be a strategy for maternal plants to continue seed production. The strong, persistent patterns of correlation between certain characters, such as flower and fruit size, in spite of extensive phenotypic plasticity, was interpreted as indirect evidence for developmental correlation. Furthermore, vegetatively larger plants produced not only more flowers and fruits, but also consistently larger flowers and fruits. The results emphasize that variation in fitness characters, such as seed size and number, should not be viewed in isolation from vegetative characters, flower, and fruit sizes in ecological and evolutionary studies, if the goal is to understand the mechanisms of natural selection in wild populations.     

EFFECTS OF MATERNAL AND PATERNAL ENVIRONMENT AND GENOTYPE ON OFFSPRING PHENOTYPE IN SOLIDAGO ALTISSIMA L.

To predict the possible evolutionary response of a plant species to a new environment, it is necessary to separate genetic from environmental sources of phenotypic variation. In a case study of the invader Solidago altissima, the influences of several kinds of parental effects and of direct inheritance and environment on offspring phenotype were separated. Fifteen genotypes were crossed in three 5 × 5 diallels excluding selfs. Clonal replicates of the parental genotypes were grown in two environments such that each diallel could be made with maternal/paternal plants from sand/sand, sand/soil, soil/sand, and soil/soil. In a first experiment (1989) offspring were raised in the experimental garden and in a second experiment (1990) in the glasshouse. Parent plants growing in sand invested less biomass in inflorescences but produced larger seeds than parent plants growing in soil. In the garden experiment, phenotypic variation among offspring was greatly influenced by environmental heterogeneity. Direct genetic variation (within diallels) was found only for leaf characters and total leaf mass. Germination probability and early seedling mass were significantly affected by phenotypic differences among maternal plants because of genotype ( genetic maternal effects ) and soil environment ( general environmental maternal effects ). Seeds from maternal plants in sand germinated better and produced bigger seedlings than seeds from maternal plants in soil. They also grew taller with time, probably because competition accentuated the initial differences. Height growth and stem mass at harvest (an integrated account of individual growth history) of offspring varied significantly among crosses within parental combinations ( specific environmental maternal effects ). In the glasshouse experiment, the influence of environmental heterogeneity and competition could be kept low. Except for early characters, the influence of direct genetic variation was large but again leaf characters (= basic module morphology) seemed to be under stricter genetic control than did size characters. Genetic maternal effects, general environmental maternal effects, and specific environmental maternal effects dominated in early characters. The maternal effects were exerted both via seed mass and directly on characters of young offspring. Persistent effects of the general paternal environment ( general environmental paternal effects ) were found for leaf length and stem and leaf mass at harvest. They were opposite in direction to the general environmental maternal effects, that is the same genotypes produced “better mothers” in sand but “better fathers” in soil. The general environmental paternal effects must have been due to differences in pollen quality, resulting from pollen selection within the male parent or leading to pre- or postzygotic selection within the female parent. The ranking of crosses according to mean offspring phenotypes was different in the two experiments, suggesting strong interaction of the observed effects with the environment. The correlation structure among characters changed less between experiments than did the pattern of variation of single characters, but under the competitive conditions in the garden plant height seemed to be more directly related to fitness than in the glasshouse. Reduced competition could also explain why maternal effects were less persistent in the glasshouse than in the garden experiment. Evolution via selection of maternal effects would be possible in the study population because these effects are in part due to genetic differences among parents.