EFFECTS OF NONRANDOM SEED ABORTION ON PROGENY PERFORMANCE IN PHASEOLUS COCCINEUS L

In this study, we sought to determine if Phaseolus coccineus normally aborts potentially viable seeds, and whether seed abortion is nonrandom with respect to progeny vigor. The ovaries of Phaseolus coccineus typically have six linearly arranged ovules. The three ovule positions at the stylar end are more likely to mature seeds than the three ovules at the basal end of the ovary. When we destroyed the developing ovules at the stylar end of the fruit after fertilization but before seed abortion, there was a significant increase in the probability that the ovules in the three basal positions would produce a mature seed. The probability of seed maturation in control fruits (no ovules destroyed) ranged from 38.3 to 42.7% over the three field seasons, whereas in the experimental fruits it ranged from 64.3 to 79.7% (similar to that of ovules at the stylar end in control fruits). We did not find any significant change in the probability of seed production in the three ovule positions in the stylar end of the ovary (the positions with high probability of seed maturation) when the basal ovules were experimentally destroyed. These results indicate that potentially viable seeds are regularly aborted in P. coccineus, with seed abortion more frequent in the basal ovule positions than at the stylar positions. In two greenhouse studies and one field study, we compared the vigor of progeny produced in the control fruits with the vigor of progeny produced in fruits where three ovules (either stylar or basal end) were destroyed. We found that the performance of the progeny from the three positions at the stylar end of the control fruits did not differ from the performance of the progeny from experimental fruits in which the three basal ovule positions were destroyed. In contrast, the progeny from the basal positions of the control fruits outperformed the progeny from the experimental fruits when the seeds were produced in the three basal ovules (stylar ovules destroyed). Our findings indicate that when the experimental treatment increased the probability of seed maturation, there was a significant decrease in the average vigor of the progeny in the remaining (basal) ovule positions.     

EFFECT OF OVULE POSITION IN THE POD ON PATTERNS OF SEED FORMATION IN TWO SPECIES OF LATHYRUS (LEGUMINOSAE: PAPILIONOIDEAE)

Using a combination of observations of fate of ovules in matured fruits and of fluorescence techniques to study pollen tube growth and fertilization of ovules, we examined patterns of seed formation within pods in natural populations of two species of Lathyrus, L. sylvestris and L. latifolius. We also examined variation in these patterns within and among populations and between two consecutive years. In both species, only a portion of the ovules were fertilized. Fertilization occurs over a period of several days and ovules at the stigmatic end of the fruit are the first to be fertilized. Fertilized ovules farthest from the stigma are closest to the maternal nutrition. The pattern of embryo abortion is interpreted as a balance between the early start and genetic quality of embryos near the stigma, on the one hand, and the nutritional advantage of proximity to maternal nutrients, on the other. Differences between the two species in patterns of seed maturation are postulated to be related to differences in breeding system. In L. sylvestris, a higher frequency of selling leads to less genetic diversity of pollen deposited on the stigma, lower competition among potential sires, and a more nearly random pattern of ovule fertilization and maturation within the pod.     

VARIATION IN SEED WEIGHT AND ITS EFFECTS ON GERMINATION IN PASTINACA SATIVA L. (UMBELLIFERAE)

Pastinaca sativa (wild parsnip) produces seeds on the primary, secondary, and tertiary umbels of the flowering stalk. Within plants, variation in seed weight is about twofold. Secondary and tertiary seed weight is 73% and 50% of primary seed weight, respectively. Maximum variation in seed weight between plants is sixfold when tertiary seeds from a small plant are compared to primary seeds from a large plant. Within an umbel order, variation in seed weight between plants is correlated with plant size. Under autumn germinating conditions in the laboratory, final germination of seeds from different umbel orders does not differ but smaller seeds germinate more rapidly than larger seeds. Under spring germination conditions in the laboratory, significantly more primary and secondary seeds germinate than tertiary seeds and the rate of germination is independent of seed weight. Field germination of seeds from different umbel orders produces similar results except that in the spring both secondary and tertiary seed germination is lower than that of primary seeds. These results suggest that with respect to seed germination characteristics small seeds may have a competitive advantage over large seeds in the autumn because they germinate more quickly, but in the spring small seeds are at a disadvantage because they have lower overall germination. Because most germination in the field occurs in the spring, population recruitment from small seeds is likely to be substanially less than that from large seeds.     

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 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.     

EFFECT OF MALE POLLEN DONOR AND FEMALE SEED PARENT ON ALLOCATION OF RESOURCES TO DEVELOPING SEEDS AND FRUIT IN CHAMAECRISTA FASCICULATA (LEGUMINOSAE)

In Chamaecrista fasciculata, fruit abortion levels are high and seed mass is highly variable, necessary preconditions for differential resource allocation of the female to seed and fruit sired by different males. This study investigated the relative role of pollen donor and seed parent on the allocation of resouces to developing seed and fruit, and assessed the role of genetic relatedness in contributing to any observed paternal effect in C. fasciculata. In addition, pollen donor effects were contrasted to within-seed parent sources of variation in resource allocation due to pollination date and ovule position in the pod. Plants collected from the field were brought to a greenhouse where single-donor crosses were conducted controlling for pollen donor source and interplant distance, a measure of genetic relatedness. Seed mass, number of seed/fruit, fruit maturation time, and fruit abortion rate were measured as indicators of resource allocation to developing seed and fruit. Variation in resource allocation was largely determined by the seed parent. Pollen donor effects were limited to differences between self vs. non-self pollinations, suggesting that inbreeding depression following mating events between related individuals is the source of any variation among pollen donors on differential resource allocation to developing seed and fruit. Once the effect of inbreeding was removed, however, pollination date and ovule position played larger roles than pollen source. Since there was no detectable variation among male pollen donors in their ability to accrue resources from the female seed parent apart from inbreeding effects, it is concluded that the opportunity for postzygotic mate choice is limited in C. fasciculata.     

PRODUCTION AND EFFECT OF SEED SIZE IN A GRASSLAND ANNUAL (ERODIUM BRACHYCARPUM,GERANIACEAE)

Over its fruit maturation period in the field, Erodium brachycarpum (Geraniaceae) exhibited a striking decrease in average seed mass; seeds produced in the 1st week of fruiting were twice the size of those produced 6 weeks later at the time of drought-induced plant senescence. Individuals with earliest fruit maturation produced the most seeds with the widest range of sizes and over a longer period than the others. In a greenhouse experiment, a similar decline in mean seed weight was generated by inducing drought conditions. Large seeds remained dormant longer and, once the seed coat was scarified, took longer to germinate than small seeds. This study indicates that small E. brachycarpum seeds (36% of the seed pool) produced late in the fruiting period are likely to germinate in the year of seed production, approximately 4–8 months after seed maturation. In contrast, the large seeds (23% of the seed pool), most of which are produced early in the fruiting period, are more likely to germinate in subsequent years (i.e., 15 months or more after seed maturation).     

DEMOGRAPHIC ASPECTS OF FLOWER AND FRUIT PRODUCTION IN BEAN PLANTS,PHASEOLUS VULGARIS L.

In this paper we assess the relative contributions to total pod yield of cohorts of flowers of known age. Bean plant (Phaseolus vulgaris L.) growth was monitored and 'births' of individual flowers were recorded and their fates followed. For every cohort of flowers an array of yield measures was calculated, including the mean and total cohort dry weight of fruits, and the numbers of viable and aborted seeds and completely undeveloped ovules. When mature, pods were removed from half of the plants. Plants whose mature pods were removed produced significantly greater numbers of flowers, ovules and viable seeds and a greater total weight of pods than did untreated plants. However, the proportion of ovules giving rise to viable seeds and to aborted seeds and undeveloped ovules was the same in plants of both groups. Approximately 50% of ovules of marketable pods, in all plants, yielded viable seeds. The 12 cohorts of flowers contributed markedly different amounts to plant yield. Early and later cohorts contributed much less than did middle cohorts in terms of marketable pod dry weight and numbers of fully developed seeds, aborted seeds, and undeveloped ovules. Peak productivity was therefore obtained from the middle phase of flowering in these annual bean plants. The lower pod yield of later cohorts is interpreted as a result of competition for limited resources between maturing pods and new flowers. We consider possible causes for the various fates of flowers and ovules and discuss flowers as plant modules suitable for demographic examination.     

A COMPARATIVE STUDY OF OVULE AND MEGAGAMETOPHYTE DEVELOPMENT IN FIELD-GROWN AND GREENHOUSE-GROWN PLANTS OF GLYCINE MAX AND PHASEOLUS AUREUS (PAPILIONACEAE)

A study of ovule and megagametophyte development in field- and greenhouse-grown plants of Glycine max (L.) Merrill and Phaseolus aureus Roxb. reveals several consistent features for both species. These features include: a multiple archesporium, enlargement of a primary sporogenous cell directly into a megasporocyte, production of unequal dyad cells, a functional chalazal megaspore, Polygonum-type development, and a hypostase. A filiform apparatus was not observed in either species. Several marked differences in development also occur. Phaseolus usually produces one sporogenous cell per ovule; Glycine produces 2–3 sporogenous cells per ovule. Meiosis II is synchronous in Phaseolus but nonsynchronous in Glycine. Linear tetrads are produced in Phaseolus, whereas linear and T-shaped tetrads are found in Glycine. Starch grains accumulate in the mature megagametophyte of Glycine but are absent at that stage in Phaseolus. The usefulness of the modified clearing fluid, benzyl benzoate-4½, for the study of ovule and megagametophyte development in Glycine max and Phaseolus aureus is here demonstrated. In addition, the study indicates for both species that megagametophyte development in plants grown under field conditions is markedly similar to development in plants grown in the more uniform conditions of the greenhouse. Accordingly, these findings suggest generally that embryological data collected from plants grown under greenhouse conditions will reflect those from plants found in nature.     

THE PHOTOINDUCTIVE CONTROL OF SEED WEIGHT IN CHENOPODIUM RUBRUM L.

The determination of mean seed weight in Chenopodium rubrum L., a short-day, weedy annual, has been examined through the manipulation of photoperiod during the course of reproductive development. Individuals that develop seeds in relatively long photoperiods (15 L: 9 D) form many more seeds of smaller weight than individuals that develop seeds in relatively shorter photoperiods (12 L: 12 D). By altering the inductive photoperiod at different times during the course of reproductive development, it is shown that the effect of photoperiod is most strongly exerted between six and twelve days after the start of floral induction and is coincident with the determination of primordia as floral structures. The ecological significance of this early developmental determination is discussed.     

EFFECTS OF CROSS AND SELF-FERTILIZATION ON PROGENY FITNESS IN LOBELIA CARDINALIS AND L. SIPHILITICA

Inbreeding depression, or the decreased fitness of progeny derived from self-fertilization as compared to outcrossing, is thought to be the most general factor affecting the evolution of self-fertilization in plants. Nevertheless, data on inbreeding depression in fitness characters are almost nonexistent for perennials observed in their natural environments. In this study I measured inbreeding depression in both survival and fertility in two sympatric, short-lived, perennial herbs: hummingbird-pollinated Lobelia cardinalis (two populations) and bumblebee-pollinated L. siphilitica (one population). Crosses were performed by hand in the field, and seedlings germinated in the greenhouse. Levels of inbreeding depression were determined for one year in the greenhouse and for two to three years for seedlings transplanted back to the natural environment. Fertility was measured as flower number, which is highly correlated with seed production under natural conditions in these populations. Inbreeding depression was assessed in three ways: 1) survival and fertility within the different age intervals; 2) cumulative survival from the seed stage through each age interval; and 3) net fertility, or the expected fertility of a seed at different ages. Net fertility is a comprehensive measure of fitness combining survival and flower number. In all three populations, selfing had nonsignificant effects on the number and size of seeds. Lobelia siphilitica and one population of L. cardinalis exhibited significant levels of inbreeding depression between seed maturation and germination, excluding the consideration of possible differences in dormancy or longterm viability in the soil. There was no inbreeding depression in subsequent survival in the greenhouse in any population. In the field, significant survival differences between selfed and outcrossed progeny occurred only in two years and in only one population of L. cardinalis. For both survival and fertility there was little evidence for the expected differences among families in inbreeding depression. Compared to survival, inbreeding depression in fertility (flower number) tended to be much higher. By first-year flower production, the combined effects on survival and flower number caused inbreeding depression in net fertility to reach 54%, 34% and 71% for L. siphilitica and the two populations of L. cardinalis. By the end of the second year of flowering in the field, inbreeding depression in net fertility was 53% for L. siphilitica and 54% for one population of L. cardinalis. For the other population of L. cardinalis, these values were 76% through the second year of flowering and 83% through the third year. Such high levels of inbreeding depression should strongly influence selection on those characters affecting self-fertilization rates in these two species.     

POLLINATION AND SEED SET IN DIERVILLA LONICERA (CAPRIFOLIACEAE): TEMPORAL PATTERNS OF FLOWER AND OVULE DEPLOYMENT

Ovule development in Diervilla lonicera (Caprifoliaceae) is limited by insufficient pollination early in the blooming period and during extensive rainy periods. Production of flowers is skewed in time; an initial burst of flowering is followed by a long period of sparse flower production. Ovule number per flower increases through the blooming period. I discuss the interactions of fruit and seed set, ovule number, and bumble bee pollinator visitation patterns. When certain flowers have a higher a priori probability of successful pollination, it may be advantageous for plants to put more ovules in those flowers. Selective ovule deployment may be a general adaptive phenomenon that has received little attention.     

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 APEX REMOVAL AND NUTRIENT SUPPLEMENTATION ON BRANCHING AND SEED PRODUCTION IN THLASPI ARVENSE (BRASSICACEAE)

The shoot apex of a plant may be damaged by herbivory, weather conditions, or other factors, leading to growth of some axillary buds into branches. This alteration of branching pattern can affect the location and extent of seed production. I examined the effects of removal of the shoot apex and of mineral nutrient addition on branching and seed production in Thlaspi arvense. Individual plants received no, early, or late apex removal and no, early, or late nutrient addition in a two-way factorial experiment. Apex removal led to greater production of secondary branches. Total seed weight per plant was higher for early- than late-removal plants under all nutrient treatments, but intact controls produced the highest total weight of seeds in two of three nutrient treatments, because their seeds were heavier. Addition of nutrients increased the numbers of secondary branches, fruits, and seeds, and the average and total weight of seeds produced. Apex removal resulted in increased seed production only when it occurred early and when nutrients were added at the same time. Resource availability and plant phenology can be influential in determining the effects of apical damage on seed production, and their effects in combination may differ from their individual effects.