Density-dependent flowering phenology,outcrossing, and reproduction in Impatiens capensis

Summary We investigated the effects of plant density on cleistogamous (CL) and chasmogamous (CH) flowering phenology and seed production in a natural Impatiens capensis population, by censusing individually marked plants at experimentally reduced and natural densities. CL flowering was earlier at natural density. This plastic density response may have resulted from a stress-related threshold for CL flowering; slower growing plants at natural density flowered earlier. Although apparently triggered by slow early growth, early CL flowering also involved an additional cost for later growth rate. In contrast, CH flowering was unrelated to relative growth rate, but apparently required a size threshold. Experimental density reduction resulted in earlier CH flowering and a dramatic increase in the percentage of plants producing CH flowers. Individual CL and CH flowering duration and flower production were greater at reduced density. These density-dependent effects caused differences between treatments in the shape and location of population flowering phenology curves. Moreover, the percentage of CH seeds produced per individual was much higher at reduced density. At natural density total seed production per plant was lower and more hierarchical than at lower density, suggesting that dominance and suppression shape jewelweed fitness distributions.     

Why is there variation in mean seed size among plants within single populations? test of the fertilization efficiency hypothesis

We tested the fertilization efficiency hypothesis, which attempts to explain mean seed size variation among plants within single populations, by comparing the patterns of seed size variation between chasmogamous (CH) flowers and cleistogamous (CL) flowers in Impatiens noli-tangere and Viola grypoceras, respectively. The fertilization efficiency hypothesis predicts that larger plants produce larger seeds if the number of pollen grains captured by a plant increases with increased allocation of resources to its attractive structures (e.g., corolla and nectar), but with diminishing gains. Thus, seed size should depend on plant size in seeds from CH flowers because of the diminishing gains of capturing pollen in these flowers, whereas seed size should not depend on plant size in seeds from CL flowers because CL flowers need not capture outcross pollen. We found significant positive correlations between mean seed size per plant and plant size for seeds from CH flowers in both species. However, there was no significant positive correlation between these two factors for seeds from CL flowers of both species. The results of the present investigations were thus consistent with the fertilization efficiency hypothesis.     

A TEST OF THE SIB-COMPETITION HYPOTHESIS FOR OUTCROSSING ADVANTAGE IN IMPATIENS CAPENSIS

One of the potential selective mechanisms invoked in discussions of breeding-system evolution is that competition within sibships increases the fitness of outcrossed progeny relative to selfed progeny. We tested this sib-competition hypothesis using cleistogamous (CL) and chasmogamous (CH) seeds of Impatiens capensis in a large greenhouse experiment. The experimental design was a double replacement series which also allowed us to test for inbreeding depression and overall resource partitioning among sibships. We found no evidence for strong inbreeding depression in the study population; although plants from CH seeds had a slight advantage over plants from CL seeds in total flower and pod production, CL plants had slightly higher growth. We also could not detect significant resource partitioning among sibships nor any evidence to support the sib-competition hypothesis for outcrossing advantage. CH sibships were not significantly more variable than CL sibships in any of the phenotypic traits measured. These results suggest that sibling competition may have little importance in the evolution of Impatiens breeding systems.     

Effects of cleistogamy and pollen source on seed production and offspring performance in three endangered violets

In rare plants that often occur in small or isolated populations the probability of selfing between close relatives is increased as a consequence of demographic stochasticity. The mode of pollination (selfing, outcrossing) may have considerable effects on seed traits and offspring performance and hence potential viability. Since current efforts aiming at the restoration of floodplain grasslands through the transfer of plant material from species-rich source stands may lead to the establishment of initially small populations consisting of founders from different populations, the present paper experimentally investigated the effects of pollen source and floral types (i.e. chasmogamous (CH) and cleistogamous (CL) flowers) on seed traits and offspring performance in three highly endangered violet species (Viola elatior, V. pumila, V. stagnina) of these grasslands. We estimated inbreeding depression and tested the performance of selfed and outcrossed offspring in two microbial environments, i.e. in soil inoculated with (i) non-sterile substrate from the same species (‘home’-conditions) and (ii) sterilised substrate.Plants produced more CL capsules than CH flowers. Pollinator exclusion had only small effects on CH seed production. CL seeds had a significantly lower mass per seed than CH seeds. This may be related to constraints in allocation or environmental conditions. Seedling growth was reduced in plants grown under ‘home’-conditions as compared to control soils. Under ‘home’-conditions, relative fitness of selfed seedlings of V. stagnina was significantly higher than that of crossed progeny. Our results suggest that high genetic differentiation among populations as a consequence of isolation may result in outbreeding depression, e.g., through biochemical or physiological incompatibilities between genes or the breaking of coadapted gene complexes. In V. stagnina, offspring fitness differed considerably between environments, but in general we found no indications for inbreeding depression in these rare species.     

Effect of density on magnitude of directional selection on seed mass and emergence time in Plantago wrightiana Dcne. (Plantaginaceae)

Although population density is believed to be an important factor influencing evolutionary processes, surprisingly few studies have documented the existence or nature of density-dependent selection. We quantified the effects of density on directional selection on seed mass (the mass of a sown seed) and emergence time in the greenhouse and field for the annual plant Plantago wrightiana. In the greenhouse, we quantified selection on seed mass and emergence time at each of five planting densities (1 m^–2 to 10,000 m^–2) using the relationship between final plant mass and each trait at each density. We observed no significant selection on either seed mass or emergence time when plants were grown alone. At all higher densities, there was significant selection favoring early emergence and large seed mass, but there were no significant differences among the selection gradients determined at densities greater than individually grown plants. In the field, we detected no relationship between the magnitude of selection for early emergence and density. Our results suggest that selection on seed mass and time of emergence is density dependent, but the relationship between density and the magnitude of directional selection on these traits is not continuously increasing. Over broad ranges of density in the greenhouse and in the field, there was no detectable relationship between density and the magnitude of directional selection.     

EFFECTS OF POSTDISPERSAL SEED PREDATION ON SPATIAL INEQUALITY AND SIZE VARIABILITY IN AN ANNUAL PLANT,ERODIUM CICUTARIUM (GERANIACEAE)

By decreasing seed density, ants introduced into flats of uniformly sown seeds of Erodium cicutarium (Geraniaceae) created differences in the neighbor-free area available to individual plants. The changes in spatial patterns brought about by the ants were greater when a higher proportion of seeds was removed but were independent of initial seed density. These spatial changes and differences in seed density were examined for their effects on plant size and reproduction. Gini values were calculated to determine inequalities. As the inequality in space among individual plants increased, the variation in final biomass increased. The number of individuals reproducing was constant among treatments, and yet seed production per plant was significantly greater for populations in which the spatial pattern was influenced by seed predation. The decrease in density and changed spatial pattern, due to previous seed predation, resulted in a few individuals having much more space than others and consequently producing many more seeds. The increase in reproductive effort per flat was much greater than could be explained by the changing density alone. Our experiment demonstrates that spatial inequality, such as that generated by seed predators, can be more important than density in generating size inequalities in plant populations. This result can profoundly alter the competitive interactions between plants and determine which plants produce seed for the next generation.     

Seed set in Guayule (Parthenium argentatum,asteraceae) in relation to insect pollination

Guayule (Tarthenium argentatum, Asteraceae) is one of two major plant species grown for natural rubber. Studies were conducted to determine the effect of honey bee (Apis mellifera) pollination and season on seed set and total seed yield/ha. The experiments involved four pollination treatments: plants caged with bees; plants caged without bees; plants open pollinated (uncovered); and plants individually covered. Seeds were harvested monthly July–September 1984, and May–September 1985. Plots with bees produced at least 150% more seeds than plots without bees, and there were no qualitative differences in the seed weights among treatments. Highest seed yield was in May and September. Results indicate that (1) insect pollination in guayule increases seed yield and (2) fewer seeds are produced in the warmest months.     

Effects of simulated herbivory on tillering and reproduction in an annual ryegrass,Lolium remotum

This study examined the overall impact of simulated herbivory on tillering and reproductive performance of an annual ryegrass,Lolium remotum. The interaction between herbivore damage and intraspecific competition and the effect of the timing of damage were also studied. The experimental plants were sown at two densities and were randomly assigned to eight different damage treatments consisting of artificial leaf area removal by clipping with scissors or removal of one-third of the ripening seeds. The treatments were executed at two flowering stages. The pattern of tiller development differed significantly among treatments and between densities. At the lower density, earlier treatments delayed tiller development more than the same treatments executed later. At the higher density, all treatments delayed tiller development. The density effect was significant for all reproductive traits measured. The reproductive output of plants grown at the higher density was lower and the negative treatment effects were stronger than at the lower density. The treatment effect was significant for seed dry weight per plant and individual seed weight but not for number of seeds per plant. There were no statistically significant interaction effects between the damage treatments and density, suggesting that the plants responded to the damage similarly, irrespective of the density. The plants did not totally compensate for losses due to damage at either density, even though they slightly increased their resource allocation to sexual reproduction at the higher density.     

The role of seed depth,litter, and fire in the seedling establishment of amphicarpic peanutgrass (Amphicarpum purshii)

Summary Amphicarpum purshii is an annual grass which mostly grows in disturbed areas of the New Jersey Pine Barrens, USA. It is amphicarpic, producing spikelets (and seeds) both above and below the soil surface. Previous research has shown that subterranean seed production ensures reproduction in the event of a major disturbance such as fire and results in rapid post-burn colonization of these sandy habitats. The effects of fire, litter, and seed depth were further examined by planting subterranean seeds at four depths in 16 litter-covered flats buried at ground level and comparing plants arising from burned flats with those in undisturbed litter-covered flats. At 0 and 1 cm depth, rates of seedling emergence were lowest in burned flats. Surface-sown seeds produced seedlings more likely to desiccate. Sowing depth had a greater influence on most measured characters than burning treatments. The mean depth of subterranean seed placement by Amphicarpum is 3.5 cm and this coincides with the seed depth from which plants showed the greatest height growth, shoot biomass, and reproductive output. In a second experiment, subterranean seeds on the bare soil surface in clay pots were more likely to lose viability and less likely to germinate than seeds protected by litter or burial in soil. In addition to providing protection from fire, placement of seeds below ground in the sandy habitat of peanutgrass provides conditions more suitable for seed survival and subsequent seedling establishment.     

Offspring performance in three cleistogamous Viola species

Seed germination and seedling/juvenile fitness in the cleistogamous perennials Viola hirta, V. mirabilis, and V. riviniana were investigated during three growing seasons, to compare the performance of chasmogamously (CH) and cleistogamously (CL) derived progeny. For V. hirta and V. mirabilis the effects of sibling competition were examined, for V. riviniana the effects of interspecific (grass) competition. Seed abortion and seed weight were also taken into account as fitness measures.In none of the species, seed abortion rate differed between CH and CL capsules. In V. mirabilis and V. riviniana, CL seeds had a lower germination rate than CH seeds. In V. hirta the two seed types did not differ in germinability. Mortality did not differ between the two seedling types in any of the species. In V. hirta and V. riviniana, CL progeny had shorter mean length of largest leaf than CH progeny. In V. mirabilis plant size did not differ between progeny types. Sibling competition had little effect on offspring performance, but grass competition increased mortality and reduced plant size of V. riviniana progeny. The two progeny types did not differ in their response to sibling and grass competition.The differences in performance between progeny types could be attributable to inbreeding depression in the CL phase, but the slightly lower fitness of CL offspring is probably balanced by their lower production costs. It is suggested that a dimorphic reproductive system is maintained in perennial Viola species to maximize total seed output in the face of environmental variation.     

Sibling competition, size variation and frequency-dependent outcrossing advantage in Plantago coronopus

Many plants display limited seed dispersal, thereby creating an opportunity for sibling competition, i.e. fitness-determined interactions between related individuals. Here I investigated the consequences of intra-specific competition, by varying density and genetic composition of neighbors, on the performance of seedlings derived by selfing or outcrossing of the partially self-fertilizing plant Plantago coronopus (L.). Seedlings from eight plants, randomly selected from an area of about 50 m² in a natural population, were used in (i) a density series with either one, four or eight siblings of each cross type per pot and (ii) a replacement series with eight plants per pot where selfed and outcrossed siblings were grown intermixed in varying frequencies. Density had a pronounced effect on plant performance. But, except for singly grown individuals, no differences were detected between selfed and outcrossed progenies in vegetative and reproductive biomass. When grown intermixed, selfed offspring were always inferior to their outcrossed relatives. The magnitude of reduction in performance was dependent on the number of outcrossed relatives a selfed seedling had to compete with, giving rise to a frequency-dependent fitness advantage to outcrossed seedlings. The major result of this study is (i) that the relative fitness of inbred progeny is strongly affected by the type of competitors (inbred or outbred) and (ii) that inbreeding depression varies according to the density and frequency of outbred plants and could be considered as a density- and frequency-dependent phenomenon. It is argued that sibling competition, due to the small genetic neighborhood of P. coronopus, might be an important selective force in natural populations of this species.     

CLEISTOGAMY IN MICROLAENA POLYNODA (GRAMINEAE): AN EXAMINATION OF SOME MODEL PREDICTIONS

A population of the cleistogamous grass, Microlaena polynoda, was investigated to determine some of the factors responsible for adjusting the balance between reproduction by chasmogamous (CH) and cleistogamous (CL) flowers, the tissue costs associated with the two reproductive modes, the fate of progeny produced by each mode, and the genetic diversity of the progeny. Cleistogamous flower production begins earlier in the season than CH flower production. There is a distinct threshold of low light intensity below which mostly CL flowers are produced. Paternal expenditure per plant is an order of magnitude larger for the CH than the CL component. The opposite relation holds for maternal expenditure. Increased maternal expenditure in the CL component may be due to greater fertilization success and retrieval of paternal costs. Cleistogamous seeds are dispersed later than CH seeds. Following dispersal, the spikelet encloses the CL seed but not the CH seed, and is responsible for inducing dormancy. The ratio of seedlings arising from CH seeds to that from CL seeds in a natural habitat is significantly lower than the ratio of estimated numbers of CH to CL seeds produced. There were no detectable polymorphisms among ten presumptive enzyme loci assayed. Many of the features associated with CH and CL reproduction in M. polynoda are in accord with the theoretical requirements for the evolution of closed flower self-pollination and the maintenance of two distinct methods of reproduction.     

Differential seed dispersal in Oxalis acetosella,a cleistogamous perennial herb

Explosive seed dispersal in the cleistogamous perennial forest herb Oxalis acetosella was studied during two growing seasons, to determine whether seeds derived from chasmogamous (CH) and cleistogamous (CL) flowers differ in dispersal distance. Seed dispersal distance, seed weight, and height of fruits were measured for both flower types, and the effects of phenology and year were also taken into account. The dispersal experiment was performed indoors, using plants transplanted from natural populations to pots. CL seeds were thrown significantly further than were CH seeds, though there was a considerable overlap in dispersal distances. There was also a significant positive relationship between seed weight and dispersal distance. No relationship was found between fruit height and dispersal distance. The results of this study contradict the common view that CL progeny should always be dispersed closer to the mother plant than CH progeny. The ecological implications of the dispersal difference are unclear, especially since it is uncertain whether CH seeds are generally outcrossed or not. Variation in dispersal distance in O. acetosella seems to be mainly dependent on a combination of reproductive mode and variation in seed weight.     

THE BALANCE BETWEEN CHASMOGAMY AND CLEISTOGAMY IN COLLOMIA GRANDIFLORA (POLEMONIACEAE)

The extent of cleistogamy (CL) and chasmogamy (CH) was investigated in natural populations of Collomia grandiflora Dougl. ex Lindl. The ubiquitous occurrence of CL flowers was documented in 18 populations from throughout the geographic range of the species. The interrelationship of environmental perturbation, vegetative weight, and the balance between CH and CL flowers was investigated by means of a factorial experiment which varied plant density, incident light and soil mix. An ANOVA revealed that total plant weight increased significantly under conditions of low density, low light and low sand content, with density accounting for most of the variance in plant weight. Interactions involving density were not significant. The number of CH and CL flowers and estimated percent outcrossing increased significantly with increased plant weight, low density, low light, and low sand content, although an analysis of covariance showed that plant weight and density accounted for most of the variance in these traits. Among large, multibranched plants, the proportion of CL flowers increased in a basipetal pattern, whereas most of the CH flowers were restricted to the terminal and upper inflorescences. The smallest plants produced only a single terminal inflorescence comprising entirely CL flowers. The proportion of reproductive effort allocated to outcrossing was dependent upon plant size as estimated from total weight, whereas the proportion allocated to ***selfing was relatively independent. Studies of outcrossed vs. inbred progenies of five populations revealed little evidence for immediate inbreeding depression with respect to plant weight and the number of CH or CL flowers, but indicated that significant quantitative genetic differences exist for these traits among populations of diverse ecological or geographical origin.     

Density Studies on Lupins. II. Components of Seed Yield

Components of seed yield of cv. Ultra (Lupinus albus L.) andcv. Unicrop (L. angustifolius L.) were measured when grown atthree densities. The low density (10 plants m^–2) Unicropyield (34 g seed per plant) was 1.8 times that of Ultra as ithad more branches, pods and seeds per pod. Ultra seeds (310mg per seed) were heavier than Unicrop seeds (180 mg). The branchingpattern of Ultra was less dependent on plant density, henceat 93 plants m^–2 it gave a higher per plant yield (7.4vs 6.4 g) than Unicrop at lower densities (83 plants m^–2).Density had most influence on pod formation and only small effectson seeds per pod and seed weight. Yield components on the main-steminflorescence were influenced less by density than componentson branch inflorescences. Later formed, higher order generationsof inflorescences were most affected by increased inter- andintra-plant competition. Pod numbers on the main-stem were similarfor both species. Pods formed at higher flower nodes in Unicrop,but the lower flower nodes were less fertile than those in Ultra.Node position of flowers had no influence on seed set in main-stemUnicrop pods, but pods from higher nodes in Ultra formed fewerseeds. Seed weights in Unicrop were similar among main-stemnodes but in Ultra seed weights tended to increase at highernodes. Lupinus spp, lupins, seed yield, planting density     

Establishment of chasmogamous and cleistogamous seedlings of an ant-dispersed UNDERSTORY HERB,CALATHEA MICANS (Marantaceae)

This study compares the establishment success of cleistogamous (CL, obligately selfed) and chasmogamous (CH, potentially outcrossed) seeds of the neotropical understory herb, Calathea micans (Marantaceae). In this species, CH and CL seeds are morphologically identical and are both dispersed by ants. I measured dispersal distances and analyzed seedling recruitment and seedling size for ant-planted CH and CL seeds, a protocol made possible by the fact that the seed coat of C. micans can be indelibly marked and remains attached to the base of the seedling for a few months. Seeds were taken by ants from experimental depots placed in natural conditions. In a second experiment, I planted CH and CL seeds in gap and understory sites to determine the effects of seed type and light on germination percentage, date of emergence, and seed and seedling survivorship. Ant-planted CH and CL seedlings did not differ in dispersal distance. CH and CL seeds did not differ significantly in recruitment probability and seedling size in either experiment. However, for the ant-planted seedlings, CH recruitment was higher in the understory than in the gap, while CL recruitment was uniform across light environments. In conclusion, I found a difference in the establishment success of CH and CL seeds in this understory herb, only after dispersal by ants in field conditions.     

The effects of seed weight on growth and competitive ability of Rumex acetosella from two successional old-fields

Summary The effects of seed size on growth, biomass allocation and competitive ability of Rumex acetosella plants grown either individually or in competition were studied in two populations (6 months and 15 years old respectively) sampled from a postcultivation successional gradient. For plants grown individually there were highly significant effects of seed weight on growth after 43 days, with a higher relative growth rate (RGR) observed for plants raised from heavier seeds. However at the end of the experiment, seedlings developed from lighter seeds had a RGR 2 times greater than those from heavier seeds. Final biomass of the two types was not significantly different after 73 days of growth. When plants were grown individually, there were only slight differences between populations, but when grown in monocultures of 4 plants per pot, plants from the old population had higher root and leaf biomass per pot whereas those from the young population had a higher reproductive effort per pot. This suggests that a trade-off between allocation to sexual and vegetative reproduction occurs over successional time. In mixtures of light and heavy seeds, plants from light seeds were shorter, had fewer leaves and lower biomass than plants from heavy seeds, which were also taller and produced more dry matter than plants grown from heavy seeds in monoculture. The significant effects of seed weight and population on biomass parameters persisted unit the end of the experiment. Seedlings from heavy seeds were strong competitors: those from the young population grew better in the presence of neighbors than in monoculture and those from the late successional population suppressed the more the growth of their partners. Seedlings from light seeds were subordinate competitors. These results suggest that seedlings from seeds of different sizes benefit from contrasting ecological conditions and that selection acts on reproductive output along successional gradients.     

FIELD GERMINATION AND SEEDLING GROWTH OF CH AND CL PROGENY OF IMPATIENS CAPENSIS (BALSAMINACEAE)

The germination behavior and early growth of chasmogamous (CH) and cleistogamous (CL) progeny of Impatiens capensis were investigated in two eastern Nebraska stands. Field germination of families of buried seeds was scored in mid-April. In one stand a significantly higher proportion of CL seeds germinated than CH seeds while no significant differences were observed in the second stand. Among-family variation in germination rates was significant in both stands. The time course of field germination was similar for CL and CH seeds. A sample of the germinated CL and CH seedlings, matched in size, was planted individually in pots and grown in the greenhouse for five wk. Early growth of CL and CH seedlings was similar. In addition, CL seedlings were as variable as CH seedlings for four morphological traits. Overall, a large fitness advantage for CH progeny was not detected in the germination and early growth of Impatiens capensis seedlings.     

EFFECT OF SEED DIMORPHISM ON THE DENSITY-DEPENDENT DYNAMICS OF EXPERIMENTAL POPULATIONS OF ATRIPLEX TRIANGULARIS (CHENOPODIACEAE)

The importance of seed size and density in determining individual plant performance and plant population dynamics in experimental populations of the halophyte Atriplex triangularis was studied. Two distinct seed morphs—large, light seeds and small, dark seeds—are produced by individual A. triangularis plants. Experimental populations consisting of seed size monocultures (large or small seeds) and seed size mixtures were established at three different densities, and the time of germination, plant size, plant survivorship, and plant fecundity were monitored. Marked variation in time of germination was observed among treatments and between seed sizes, but germination within any given treatment occurred over a five- to ten-day period. Large seeds produced larger plants than small seeds did, and this dichotomy was maintained over the course of the entire experiment. Germination date and seed size interacted such that larger plants grew from seeds which germinated earlier than those which germinated later, regardless of seed size. Germination date had a more pronounced effect than seed size did on plant mortality in high density populations. At high density, large seed monocultures experienced greater mortality than small seed monocultures did, but in seed size mixtures, the mortality was evenly distributed between plants from the two seed sizes. Regardless of density conditions and parentage, large and small seeds were produced in equal proportion by the plants. Total seed production, however, was dramatically affected by plant density, and to a lesser degree by germination date. Although seed size effects alone did not appear to affect directly final plant biomass and fecundity, effects of seed size early in ontogeny may have contributed to differences in fecundity.     

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.