VARIATION IN SEED CHARACTERS IN NEMOPHILA MENZIESII: EVIDENCE OF A GENETIC BASIS FOR MATERNAL EFFECT

A growing body of evidence indicates that phenotypic selection on juvenile traits of both plants and animals may be considerable. Because juvenile traits are typically subject to maternal effects and often have low heritabilities, adaptive responses to natural selection on these traits may seem unlikely. To determine the potential for evolutionary response to selection on juvenile traits of Nemophila menziesii (Hydrophyllaceae), we conducted two quantitative genetic studies. A reciprocal factorial cross, involving 16 parents and 1960 progeny, demonstrated a significant maternal component of variance in seed mass and additive genetic component of variance in germination time. This experiment also suggested that interaction between parents, though small, provides highly significant contributions to the variance of both traits. Such a parental interaction could arise by diverse mechanisms, including dependence of nuclear gene expression on cytoplasmic genotype, but the design of this experiment could not distinguish this from other possible causes, such as effects on progeny phenotype of interaction between the environmental conditions of both parents. The second experiment, spanning three generations with over 11,000 observations, was designed for investigation of the additive genetic variance in maternal effect, assessment of paternal effects, as well as further partitioning of the parental interaction identified in the reciprocal factorial experiment. It yielded no consistent evidence of paternal effects on seed mass, nor of parental interactions. Our inference of such interaction effects from the first experiment was evidently an artifact of failing to account for the substantial variance among fruits within crosses. The maternal effect was found to have a large additive genetic component, accounting for at least 20% of the variation in individual seed mass. This result suggests that there is appreciable potential for response to selection on seed mass through evolution of the maternal effect. We discuss aspects that may nevertheless limit response to individual selection on seed mass, including trade-offs between the size of individual seeds and germination time and between the number of seeds a maternal plant can mature and their mean size.     

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.     

CONSTANCY OF POPULATION PARAMETERS FOR LIFE-HISTORY AND FLORAL TRAITS IN RAPHANUS SATIVUS L. II. EFFECTS OF PLANTING DENSITY ON PHENOTYPE AND HERITABILITY ESTIMATES

To determine the effect of growing conditions on population parameters in wild radish, (Raphanus sativus L.: Brassicaceae), we replicated maternal and paternal half-sib families of seed across three planting densities in an experimental garden. A nested breeding design performed in the greenhouse produced 1,800 F₁ seeds sown in the garden. We recorded survivorship, measured phenotypic correlations among and estimated narrow-sense and broad-sense heritabilities (h²) of: days to germination, days to flowering, petal area, ovule number/flower, pollen production/flower, and modal pollen grain volume. Survivorship declined with increasing density, but the relative abundances of surviving families did not differ significantly among densities. Seeds in high-density plots germinated significantly faster than seeds sown in medium- or low-density plots, but they flowered significantly later. Plants in high-density plots had fewer ovules per flower than those in the other treatments. Petal area and pollen characters did not differ significantly among densities. Densities differed with respect to the number and sign of significant phenotypic correlations. Analyses of variance were conducted to detect additive genetic variance (V_a) of each trait in each density. At low density, there were significant paternal effects on flowering time and modal pollen grain volume; in medium-density plots, germination time, flowering time and ovule number exhibited significant paternal effects; in high-density plots, only pollen grain volume differed among paternal sibships. The ability to detect maternal effects on progeny phenotype also depended on density. Narrow-sense h² estimates differed markedly among density treatments for germination time, flowering time, ovule number and pollen grain volume. Maternal, paternal and error variance components were estimated for each trait and density to examine the sources of variation in narrow-sense h² across densities. Variance components did not change consistently across densities; each trait behaved differently. To provide qualitative estimates of genetic correlations between characters, correlation coefficients were estimated using paternal family means; these correlations also differed among densities. These results demonstrate that: a) planting density influences the magnitude of maternal and paternal effects on progeny phenotype, and of h² estimates, b) traits differ with respect to the density in which heritability is greatest, c) density affects the variance components that comprise heritability, but each trait behaves differently, and d) the response to selection on any target trait should result in different correlated responses of other traits, depending on density.     

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.     

Testing population differentiation in plant species – how important are environmental maternal effects

The maternal environment may contribute to population differentiation in offspring traits if growing conditions of mother plants are different. However, the magnitude of such environmental maternal effects compared with genetic differentiation is often not clear. We tested the importance of environmental maternal effects by comparing population differentiation in parental seed directly collected in the field and in F₁ seed grown under homogeneous conditions. The F₁ seeds were obtained by random crosses within populations. We used five populations in each of four plant species to analyse seed mass and growth chamber germination of both generations at the same time. In two species, we additionally tested offspring performance in the field. We found a significant population differentiation in all species and for nearly all measured traits. Population‐by‐generation interactions indicating environmental maternal effects were significant for germination (three species) and for seed mass (two species) but not for growth and reproduction. The significant interaction was partly due to a reduction of among‐population differentiation from the parental to the F₁ generation that can be explained by a decrease of maternal provisioning effects. However, in some species by trait combinations a change in population ranking and not a decrease of variation was responsible for significant population‐by‐generation interactions indicating environmental maternal effects beyond maternal provisioning. Fitting of seed mass as covariate was not successful in reducing environmental maternal effects on population differentiation in germination. We discuss alternative methods to account for environmental maternal effects in studies on genetic differentiation among populations.     

The effect of maternal and paternal environments on seed characters in the herbaceous plant Campanula Americana (Campanulaceae)

Maternal environments typically influence the phenotype of their offspring. However, the effect of the paternal environment or the potential for joint effects of both parental environments on offspring characters is poorly understood. Two populations of Campanula americana, a woodland herb with a variable life history, were used to determine the influence of maternal and paternal light and nutrient environments on offspring seed characters. Families were grown in the greenhouse in three levels of light or three levels of nutrients. Crosses were conducted within each environmental gradient to produce seeds with all combinations of maternal and paternal environments. On average, increasing maternal nutrient and light levels increased seed mass and decreased percentage germination. The paternal environment affected seed mass, germination time, and percentage germination. However, the influence of the paternal environment varied across maternal environments, suggesting that paternal environmental effects should be evaluated in the context of maternal environments. Significant interactions between family and the parental environments for offspring characters suggest that parental environmental effects are genetically variable. In C. americana, the timing of germination determines life history. Therefore parental environmental effects on germination timing, and genetic variation in those parental effects, suggest that parental environments may influence life history evolution in this system.     

Genetic control of seed proteins in wheat

Summary Electrophoretic profiles of crude protein extracts from seed of F₁ hybrids and reciprocal crosses among diploid, tetraploid and hexaploid wheats were compared with those of their respective parental species. The electrophoretic patterns within each of three pairs of reciprocal crosses, T.boeoticum X T.urartu, T.monococcun X T. urartu and T.dicoccum X T. araraticum, were different from one another but were identical with those of their respective maternal parents. Protein bands characteristic of the paternal parents were missing in F₁ hybrid seed suggesting that the major seed proteins in wheat were presumably regulated by genotype of the maternal parent rather than by the seed genotype. However, in another three pairs of reciprocal crosses, T.boeoticum X T. durum, T.dicoccum X T.aestivum and T. zhukovskyi x T. aestivum, protein bands attributable to the paternal parents were present in the F₁ hybrid seeds indicating that the seed proteins were not always exclusively regulated by the maternal genotype. The expression of paternal genomes is presumably determined by dosage and genetic affinity of the maternal and paternal genomes in the hybrid endosperm. The maternal regulation of seed protein content is probably accomplished through the maternal control over seed size. The seed protein quality may, however, depend upon the extent of expression of the paternal genome.     

The influence of light on paternal plants in Campanula americana (Campanulaceae): pollen characteristics and offspring traits

Offspring trait expression is determined by the combination of parental genes and parental environments. Although maternal environmental effects have been widely characterized, few studies have focused on paternal environmental effects. To determine whether light availability influences pollen and offspring traits in the woodland herb Campanula americana, we reared clones of 12 genotypes in two light levels. In the parental generation we measured pollen number and size. Plants grown under high light produced more pollen grains per flower than those grown under low light. However, the response was genotype specific; some individuals responded little to changes in light availability while others substantially reduced pollen production. As a consequence, paternity ratios may vary between light environments if more pollen is associated with greater siring success. We crossed a subset of these plants to produce the offspring generation. The paternal and maternal light environments influenced offspring seed mass, percentage germination, and days to germination, while only maternal light levels influenced later life traits, such as leaf number and size. Maternal and paternal environmental effects had opposite influences on seed mass, percentage germination and days to germination. Finally, there was no direct relationship between light effects on pollen production and offspring trait expression.     

EMBRYO GROWTH AND SEED SIZE IN RAPHANUS SATIVUS: MATERNAL AND PATERNAL EFFECTS IN VIVO AND IN VITRO

Theories on the evolution of the angiosperm seed disagree as to the effects of different plant tissues on embryo growth. To examine the relative contributions of maternal and paternal genes on embryo growth, we conducted controlled crosses in the greenhouse with wild radish plants (Raphanus sativus), looked for maternal, paternal, and interaction effects on embryo development, and compared the performance of embryos within fruits and in embryo culture. Maternal plant identity affected fruit set, seeds per fruit, embryo developmental stage, and mean seed weight. In embryo culture, maternal effects were found for cotyledon size and embryo weight. Paternal effects were fewer or smaller in magnitude than maternal effects. The identity of the pollen donor affected embryo developmental stage and mean seed weight. In culture, paternal effects were detected for cotyledon size and embryo weight. Our results demonstrate that both maternal and paternal elements affect embryo growth. The fact that maternal effects are greater than paternal effects on embryo development in culture may result from cytoplasmic elements or maternal nuclear genes. Embryo performance in vivo compared to that in vitro varied among maternal plants. The interaction between an embryo and its endosperm and maternal tissues may be either positive or negative, depending upon the maternal plant and the embryo's developmental stage.     

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.     

CROSSING-DISTANCE EFFECTS IN DELPHINIUM NELSONII: OUTBREEDING AND INBREEDING DEPRESSION IN PROGENY FITNESS

Depending on its genetic causes, outbreeding depression in quantitative characters may occur first in the free-living F₁ generation produced by a wide cross. In 1981–1985, we generated F₁ progenies by hand-pollinating larkspurs (Delphinium nelsonii) with pollen from 1-m, 3-m, 10-m, or 30-m distances. From the spatial genetic structure indicated by previous electrophoretic and reciprocal transplantation studies, we estimate that these crosses range from being inbred (f ≈ 0.06) to outbred. We planted 594 seeds from 66 maternal sibships under natural conditions. As of 1992, there was strong evidence for both inbreeding depression and outbreeding depression. Progeny from intermediate crossing distances grew approximately twice as large as more inbred or outbred progeny in the first 5 yr after planting (P = 0.013, repeated measures ANOVA), and survived almost 1 yr longer on average (contrast of 3-m and 10-m treatments versus 1 m and 30 m; P = 0.028, ANOVA). Twenty maternal sibships produced flowering individuals; only four and two of these represented 1-m and 30-m crossing distances, respectively (P = 0.021, G-test). The cumulative fitness of intermediate distance sibships averaged about twice that of 1-m sibships, and five to eight times that of 30-m sibships (P = 0.017, ANOVA). Thus, even though progeny of 1-m crosses were inbred to a degree only about one-eighth that of selling, inbreeding depression approximated 50%, and outbreeding depression equaled or exceeded 50% for all fitness components.     

Seed mass,genotype, and density effects on growth and yield of Oenothera biennis L.

Summary Seed mass and genotypic effects on the growth and reproduction of Oenothera biennis L. over a gradient of intraspecific density were examined in a greenhouse experiment. By using genetically identical seeds from five parental genotypes we were able to examine independently the effects of seed mass and genotype on seedling and adult performance. Seedling size was significantly correlated with seed mass for the first five weeks but had no effect on adult size or reproductive output. In contrast, genotype differences became increasingly apparent with time. In particular, there were striking differences in reproductive output among genotypes. Plants grown from two of the genotypes consistently produced more, but lighter, seeds and a greater proportion flowered at high density than the other three genotypes. In all five genotypes, seed number was much more variable than seed mass across the density gradient. Initial seed mass accounted for a significant proportion of the variation in progeny seed mass, and mean seed mass produced in the greenhouse was positively correlated with mean seed mass of the parent (in the field). This result, together with the observed constancy of seed mass within a genotype across the density gradient, indicates the differences in reproductive output among these genotypes are genetically determined.     

Parental effects in a partially self-incompatible herb Campanula rapunculoides L. (Campanulaceae): influence of variation in the strength of self-incompatibility on seed set and progeny performance

We employ a full reciprocal diallel design between 10 parental plants that differed in their strength of self-incompatibility (SI; strong, intermediate, and weak) to examine parental effects on seed set and 10 components of fitness of progeny performance in Campanula rapunculoides. We perform ANOVAs to separate the influence of the strength of SI and the identity of the maternal and paternal parent on family performance. We calculate the phenotypic and genetic correlations between traits to determine potentially evolutionary constraints. Finally, we employ maximum likelihood methods to estimate the components of quantitative genetic variance, as defined by Cockerham and Weir in their BioModel c. Our most significant finding is that weak SI plants have high outcrossed seed set as maternal parents. We argue that direct or pleiotropic effects of modifiers of SI probably cause this. Second, we find that extranuclear interactions, as defined by the BioModel, have strong effects on seed set and several vegetative and flowering traits. These findings indicate that some maternal plants selectively provision seeds sired by specific paternal donors and that some of this variation appears to be associated with modifiers of the strength of SI. We find other sources of significant quantitative genetic variation for all of the traits we examine and discuss the possible role these play in the evolution of the reproductive system. Taken together, our findings show that variation in the strength of SI may influence levels of quantitative genetic variation that, in turn, can influence the reproductive success of individuals in C. rapunculoides     

SEED SET AND SEED MASS IN IPOMOPSIS AGGREGATA: VARIANCE PARTITIONING AND INFERENCES ABOUT POSTPOLLINATION SELECTION

Events that follow pollination, such as pollen-tube growth and seed maturation, comprise an important phase of angiosperm reproduction. Differential success during this “postpollination” phase may represent phenotypic selection, including sexual selection, or interaction between parents caused, for example, by their genetic similarity. By providing a detailed partitioning of variance in success, diallel crossing designs offer great potential to determine which processes are occurring and their relative magnitudes. We performed three partial diallels with the montane herb Ipomopsis aggregata, using a large sample of parental plants (69 total). Embedded in the designs were crossing-distance treatments of 1 m, 10 m, and 100 m, reflecting a range of parental genetic similarity. We partitioned phenotypic variance in seed set per fruit into six components using restricted maximum-likelihood (REML) analysis. For one diallel, we also partitioned variance in seed mass into five components, and estimated two components of covariance between seed set and mass. Variance caused by maternal effects (V_mat) comprised 12%–35% of total variance in seed set and 62% of variance in seed mass, and there was a significant negative environmental covariance between seed set and seed mass. Parental interaction made no detectable contribution to phenotypic variance in either of our measures of postpollination success, although crossing distance did contribute slightly but significantly to fit of the model in some cases. Finally, there was no detectable paternal variance (V_pat) in seed set or seed mass. These results are in keeping with reports from other studies of natural plant populations. The finding of little or no paternal variance in particular suggests little scope for postpollination sexual selection through the male function of cosexual plants such as I. aggregata.     

Maternal effects on early juvenile traits inIris pumila (Iridaceae)

The variation in the maternal effects on a number of the earlyIris pumila seedling traits expressed under high and low light levels in a growthroom was studied. A reciprocal crossing design was used for the establishment of 20 full-sib families. Each family consisted of two maternal groups differing only in genotype used as a seed parent. Maternal variation was estimated from the variance between maternal groups within sibships. Two parameters for maternal variation were computed: the maternal effects coefficient m² and the coefficient of maternal variation CV_M. Maternal effects significantly accounted for the phenotypic variation in seed mass and most of the early seedling traits. Light conditions prevailing during the seedling development markedly influenced overall structure of maternal correlations. Implications of maternal correlations on the integration of juvenile phenotype have also been discussed.     

Maternal and paternal contributions to the fitness of hybrids between red and white mulberry (Morus, Moraceae)

The fitness of hybrids depends on the genetic disparity between parental taxa and the magnitude of their nuclear and non-nuclear contributions. To estimate the role of non-nuclear effects, we crossed red (R), white (W) and hybrid (H) mulberry in all combinations and compared the magnitude of maternal and paternal effects on offspring fitness (seed set, germination, survival and aboveground biomass) in a greenhouse environment. Variation in offspring fitness was determined largely by the identity of the maternal parent; specifically, progeny with white mothers had the highest cumulative fitness. As fathers, red, white, and hybrid mulberry had no effect on fitness, and maternal × paternal interactions were significant only for survival. Individual cross-types differed significantly for all fitness components except seed set. Offspring from hybrid crosses (W × R, H × R, H × W) often differed from at least one of the within-parent crosses (W × W, R × R) as well as from other hybrid crosses, although their fitness values never exceeded the most fit parent. Reciprocal crosses differed in only two of 15 possible parental combinations: W × H (cumulative fitness) and W × R (aboveground biomass). Overall, the strong asymmetry in magnitude of maternal and paternal effects suggests that fitness of hybrid mulberry is governed largely by non-nuclear, parental effects.     

QTL for yield in bioenergy Populus: identifying G×E interactions from growth at three contrasting sites

Populus is a genus of fast growing trees that may be suitable as a bioenergy crop grown in short rotation, but understanding the genetic nature of yield and genotype interactions with the environment is critical in developing new high-yield genotypes for wide-scale planting. In the present study, 210 genotypes from an F₂ population (Family 331; POP1) derived from a cross between Populus trichocarpa 93-968 and P. deltoides ILL-129 were grown in southern UK, central France and northern Italy. The performance of POP1, based upon first- and second-year main stem traits and biomass production, improved from northern to southern Europe. Trees at the Italian site produced the highest mean biomass ranging from 0.77 to 18.06 oven-dried tonnes (ODT) ha⁻¹ year⁻¹, and the UK site produced the lowest mean biomass ranging from 0.18 to 10.31 ODT ha⁻¹ year⁻¹. Significant genotype × environment interactions were seen despite heritability values across sites being moderate to high. Using a pseudo-testcross analysis, 37 quantitative trait loci (QTL) were identified for the maternal parent and 45 for the paternal parent for eight stem and biomass traits across the three sites. High genetic correlations between traits suggested that collocating QTL could be inferred as a single pleiotropic QTL, reducing the number of unique QTL to 23 and 24 for the maternal and paternal parent, respectively. Additive genetic effects were seen to differ significantly for eight QTL on the maternal map and 20 on the paternal map across sites. An additive main effects and multiplicative interaction analysis was carried out to obtain stability parameters for each trait. These parameters were mapped as QTL, and collocation to trait QTL was accessed. Two of the eight stability QTL collocate to trait QTL on the maternal map, and 8 of the 20 stability QTL collocate to trait QTL on the paternal map, suggesting that a regulatory gene model is prevalent over an allele sensitivity model for stem trait stability across these environments.     

PSA: software for parental structure analysis of seed or seedling patches

Parental structure analysis (PSA) is a computer program to analyse separate contributions of paternal and maternal parents to postdispersal plant offspring. The program provides joint estimates of maternal, paternal and cross‐parental correlations within and among a set of predefined groups of seeds or seedlings, as well as derivative estimates of effective parental numbers. PSA utilizes data sets that distinguish between maternal and paternal contributions to the genotype of each offspring in the sample, but does not require parental samples per se. The approach requires assay of codominant diploid markers from both seed coat (maternally inherited) and seedling/embryo (biparentally inherited) tissues for each offspring. A simulation analysis of PSA's performance shows that it provides fairly accurate parental correlation estimates from affordable sampling effort. PSA should be of interest to plant biologists studying the interplay between dispersal, demography and genetics, as well as plant–animal interactions.     

Quantitative genetics of seed size variation inPhlox

Summary Because seed size is often associated with survival and reproduction in plant populations, genetic variation for seed size may be reduced or eliminated by natural selection. To test this hypothesis we assessed genetic sources of variation in seed size in a population ofPhlox drummondii to determine whether genetic differences among seeds influence the size they attain. A diallel cross among 12 plants from a population at Bastrop, Texas, USA allowed us to partition variance in the mass of seeds among several genetic and parental effects. We found no evidence of additive genetic variance or dominance genetic variance for seed mass in the contribution of plants to their offspring. Extranuclear maternal effects accounted for 56% of the variance in seed mass. A small interaction was observed between seed genotype and maternal plant. Results of this study support theory that predicts little genetic variation for traits associated with fitness.     

ENVIRONMENTAL AND GENETIC MATERNAL EFFECTS ON SEED CHARACTERS IN NEMOPHILA MENZIESII

Nuclear genetic, maternal genetic and maternal environmental effects on seed characters were estimated in the California native annual plant Nemophila menziesii using two greenhouse crosses. In one cross, according to a nested mating design, the narrow sense heritability of seed weight was small (3.9%). A subset of full-sib progenies produced in this cross was grown singly and in competition with the introduced grass Bromus diandrus. In a second cross, these plants were used as mothers (dams) and were each mated to the same three sires. Seeds produced by mothers competing with B. diandrus showed a significant reduction in weight, increase in time to germination, and increase in the incidence of dormancy, when compared to seeds from mothers grown singly. Significant sire components were found for time to germination and incidence of dormancy. Maternal genetic variation for seed weight was largely expressed as maternal genotype by maternal environment interaction, and showed no significant maternal genetic main effect. Time to germination and dormant fraction showed a relatively large maternal genetic effect. Evolution of seed characters in N. menziesii is more likely to occur via indirect response to selection among maternal plants than among the seeds themselves. Maternal genotype by maternal environment interaction could potentially contribute to the maintenance of maternal genetic variation in seed weight, but this does not appear likely for dormancy.