LIPID RESERVES OF SEEDS OF IMPATIENS CAPENSIS AND I. PALLIDA (BALSAMINACEAE): DEVELOPMENTAL ASPECTS

It has been known since 1948 that seeds of Impatiens species (family Balsaminaceae) contain a polyconjugated 18:4 fatty acid, α-parinaric acid. Content of this acid is high in seeds of the two species of Impatiens native to eastern North America, I. capensis Meerb. and I. pallida Nutt. The cotyledons of these seeds also have an unusual blue pigmentation that disappears when lipids are extracted. Deposition of oil reserves and appearance of cotyledon color was observed during fruit development. Pods of I. capensis and I. pallida were measured daily after anthesis. Length increased linearly to a maximum of about 27 mm (I. capensis) and 30 mm (I. pallida) by day 10 after which increase in girth and fresh weight of pods, and fresh weight of seeds, increased exponentially. At maturity (15 days) seeds of I. pallida were twice the weight of seeds of I. capensis. Lipid reserves, extracted with chloroform-methanol, were detectable by day 6 and increased rapidly to 28% fresh weight (2 mg/seed) in seeds of I. capensis and to 24% (4 mg/seed) in I. pallida. The characteristic ultra-violet (uv) spectrum of α-parinaric acid was already present by day 6, and content of this acid increased to 42% of total lipids in I. capensis and to 30% in I. pallida at maturity. Fatty acids as determined by gas chromatography of oil of mature seeds were 16:0 (13%, I. capensis; 11%, I. pallida); 16:1 (trace); 18:0 (2%); 18:1 (20%, 25%); 18:2 (17%, 14%); 18:3 + 18:4 (48%, 47%). Electron micrographs of mature seeds of I. capensis showed abundant oleosomes and protein bodies. The blue color of the cotyledons was faint at day 11 and then intensified with age. It was somewhat paler in I. pallida than in I. capensis, suggesting that it may be associated with the presence of α-parinaric acid.     

THE RELATIVE COSTS OF SELF- AND CROSS-FERTILIZED SEEDS IN IMPATIENS CAPENSIS (BALSAMINACEAE)

By using a generally applicable technique that involves monitoring the development and survivorship of flowers and seed capsules, I estimated the material and energetic costs of producing self- and cross-fertilized seeds in Impatiens capensis. All flowers and fruits on six plants were censused intensively for the two-month period of reproduction. Cleistogamous (selfing) flowers ripened seed in about 24 days, compared to about 36 days for the chasmogamous (outcrossing) flowers. In terms of dry weight, selfed seeds cost about two-thirds as much as outcrossed seeds: 12.4 versus 18.4 mg dry weight per seed. When adjusted to the currency of calories, and including an independent estimate of pollen and nectar production in outcrossing flowers, I estimate the costs to be about 65 and 135 calories per selfed or outcrossed seed. Sources of error include the accuracy of the estimates of flower and fruit weight, and possible differences among the developmental stages in respiratory costs. The cost discrepancy implies that outcrossed seeds should possess a countervailing fitness advantage large enough to offset their greater energetic cost.     

FACTORS INFLUENCING SEED WEIGHT IN IMPATIENS CAPENSIS (BALSAMINACEAE)

I analyzed within- and between-plant variation in seed weight from one greenhouse and four natural populations of Impatiens capensis Meerb. Most variation in seed size in natural populations occurred within populations and within plants. Weights of seeds from cleistogamous flowers on greenhouse plants increased with height on the plant and proximity to the main stem, a result that may be related to the local supply of photosynthate. Mean seed weight in the natural populations declined during seed set, then increased again. Seeds from chasmogamous flowers were significantly larger in one of the four populations. Maternal parents differed significantly in both mean seed weight and the relative size of the two types of seeds. Combined genetic and general environmental effects accounted for 26% of the total phenotypic variance, which provides an outside estimate of the heritability of this character.     

GAS EXCHANGE OF IMPATIENS PALLIDA NUTT. (BALSAMINACEAE) IN RELATION TO WILTING UNDER HIGH LIGHT

Impatiens pallida, a succulent annual herb of moist temperate forests, typically wilts on summer days after several minutes of direct sunlight. Time courses of gas exchange and leaf water potential were measured to determine if wilting resulted in substantially reduced photosynthesis, stomatal conductance, or leaf internal CO₂ concentrations. Leaves quickly wilted with the onset of high-light, but photosynthesis and stomatal conductance increased markedly. Photosynthetic rates and stomatal conductance declined slightly after several hours of high-light, and from morning to late afternoon shade conditions. Leaf internal CO₂ declined with increased photosynthesis, but there was no evidence that stomatal conductance limited photosynthesis through the day. We propose that rapid wilting is an adaptation that facultatively limits heat loading and extreme water loss under high-light. Further whole plant studies in natural settings are needed to fully evaluate the quantitative significance of wilting in relation to water use and photosynthesis.     

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.     

DISTANCE BETWEEN MATES AFFECTS SEEDLING CHARACTERS IN A POPULATION OF IMPATIENS CAPENSIS (BALSAMINACEAE)

We tested for the presence of an optimal outcrossing distance in a population of Impatiens capensis, an annual herb that possesses a mixed mating system, by measuring fitness components for offspring of parents that were separated by distances of 2, 10, or 50 m. Heavier seed weights and later seed maturation enhanced probabilities of emergence and subsequent survival, but emergence and survival were unaffected by outcrossing distance. Path analysis showed that most characteristics of seeds and seedlings were positively correlated. Effects of maternal family were strong through the first month of growth. Analyses of covariance reveal that, as anticipated, the effects of maternal family, seed weight, and seed maturation date declined over time. Seeds that were produced late in the season were heavier than early seeds, and late seeds germinated at later dates than early seeds. Surprisingly, although late germination dates were associated with taller plants after 1 and 3 months of growth, late germination resulted in lighter dry weights at maturity. The distance between parents had a negative direct effect on seed weight but a positive effect on height after 1 month. Height at 1 month responded positively to outcrossing distance and negatively to the square of distance when seed weight, collection date, and maternal family were held constant. This implies an optimal outcrossing distance for height at 1 month, which we estimate as 29 m using a quadratic procedure. However, path analysis suggests that the apparent optimum is due to a negative correlation between the interparent distance and seed weight.     

VARIATION IN POPULATION GROWTH RATE IN THE WOODLAND ANNUAL IMPATIENS PALLIDA (BALSAMINACEAE)

Variation in population growth rate over environmental gradients was determined for the annual Impatiens pallida (Balsaminaceae), by monitoring age- and size-specific survivorship and fecundity in five populations of this summer-flowering and primarily woodland species. All I. pallida seedlings emerged within a few days of each other and a size hierarchy was established within a month, and remained unchanged thereafter. Light and initial seedling density together explained 67% of the variance in mean adult plant size. As a result of differences among populations in the amount of disturbance, survivorship to the time of floral bud production ranged from 39% to 93% of the initial seedling total. Survivorship during the reproductive period was most affected by plant size and soil moisture. For plants surviving to flowering time, the probability of setting seed, the type of seed and the number of seeds produced per plant were significantly positively correlated with plant size. More than 82% of all seeds were produced by plants greater than or 1 m tall. Cleistogamy was the major form of reproduction in 4 out of 5 populations monitored. The net reproductive rate (R₀) differed greatly among populations, ranging from 1.23 to 16.27. Population growth rate increased with increasing resource availability and decreasing disturbance during the growing season. The magnitude of R₀ depended primarily on 1) timing, intensity and frequency of disturbance, 2) length of the reproductive period, and 3) population size structure.     

NEIGHBOR RELATEDNESS AND COMPETITIVE PERFORMANCE IN IMPATIENS CAPENSIS (BALSAMINACEAE): A TEST OF THE RESOURCE PARTITIONING HYPOTHESIS

The Elbow Room hypothesis for the maintenance of sex depends upon the assumption that intraspecific resource partitioning occurs and increases in magnitude with decreasing genetic similarity of competitors. This assumption leads to the prediction that plants should have greater fitness in competition with nonrelatives than in competition with siblings. Moreover, if a population displays fine scale genetic structure and genetic similarity declines with distance, then resource partitioning should increase with increasing geographical distance between the source locations of the competing genotypes. We tested these predictions in a greenhouse experiment by subjecting inbred Impatiens capensis seedlings to four types of competitors: 1) inbred full sibs; 2) inbred nonrelatives from the same source location; 3) inbred nonrelatives from a source location 30 m away; and 4) inbred nonrelatives from another population approximately 1 km distant. Plant dry weight at harvest increased significantly with seed weight and earlier emergence date, decreased significantly with seed weight and earlier emergence date of the competitor, and varied significantly among maternal seed families. However, there was no significant effect of competitor relatedness or distance between parental locations. The experiment therefore failed to support the resource partitioning hypothesis.     

SEED BANK,SURVIVORSHIP, AND SIZE DISTRIBUTION OF A NEBRASKA POPULATION OF IMPATIENS CAPENSIS (BALSAMINACEAE)

The growth and survivorship of a population of Impatiens capensis were followed in eastern Nebraska. Soil cores were taken within the population to estimate the size of the seed bank. The number of seeds in the soil remained relatively constant from April through October, with a mean of 200 seeds/m². The seed bank size corresponded to observed seedling densities in the following years. Twenty permanent plots were established within the population and censused for three years, 1983–1985. Seedling recruitment to the population was similar in 1983 and 1984 but 30% higher in 1985. The seedling densities of plots were uncorrelated from year to year. Survivorship in 1983 followed a Deevey Type I pattern. Almost one-third of the population survived until chasmogamous reproduction. Plant size (stem height) was measured monthly on all individuals during 1983. The frequency distribution of stem height was positively skewed as long as growth was exponential. When the growth rate slowed, the distribution became more normal.     

ABERRANT MICROSPOROGENESIS AND STERILITY IN IMPATIENS SULTANI (BALSAMINACEAE)

Three mutants of Impatiens sultani (named as ‘Orange', ‘Crimson', and ‘Pink') show an unusually wide spectrum of aberrations in microsporogenesis. These aberrations range from premeiotic cytomixis to meiotic and postmeiotic irregularities such as precocious disjunction, tripolar separation of chromosomes, chromosome bridges, various patterns of atypical cytokinesis, supernumerary divisions of the meiotic products, and irregular divisions in the microspore. As a result of these abnormalities Orange is partially and Crimson and Pink are completely pollen sterile. The stigmas of Pink, besides, do not support pollen germination. The failure of pollen germination is attributed to the defective stigmatic fluid of Pink, which, compared to that of Orange and Crimson lacks organic compounds absorbing UV light at 261 nm. This constitutive deficiency in stigmatic fluid appears to be a little investigated facet of female sterility.     

QUANTITATIVE GENETICS OF SURVIVAL AND GROWTH IN IMPATIENS CAPENSIS

When variation in life-history characters is caused by many genes of small effect, then quantitative-genetic parameters may quantify constraints on rate and direction of microevolutionary change. I estimated heritabilities and genetic correlations for 16 life-history and morphological characters in two populations of Impatiens capensis, a partially self-pollinating herbaceous annual. The Madison population had little or no additive genetic variance for any of these characters, while the Milwaukee population had significant narrowsense heritabilities and genetic correlations for several traits, including adult size, which is highly correlated with fitness. All genetic correlations among fitness components were positive, hence there is no evidence for antagonistic pleiotropy among these traits. Dissimilarity of heritabilities in the two populations supports theoretical predictions that long-term changes in genetic variance-covariance patterns may occur when population sizes are small and selection is strong, as may occur in many plant species.     

RELATIVE PERFORMANCE OF SELFED AND OUTCROSSED PROGENY IN IMPATIENS CAPENSIS

This study compares survival and growth of progeny derived from chasmogamous (CH) and cleistogamous (CL) flowers in Impatiens capensis, a forest annual. When progeny were grown in the field, CH seeds had significantly higher survival rates over winter (64% versus 56%), and the survival advantage of outcrossed progeny was not attributable to seed weight differences. No differences in seedling growth were detected. Greenhouse comparisons revealed no difference in seed survival but a 30% growth advantage to CH seedlings. We found no changes in developmental homeostasis of three leaf shape characters between inbred and outbred progeny, nor was there any difference in variability within CH and CL families. The outcrossing advantage observed in these experiments could not have been caused by avoidance of sib competition. Theory predicts that self-pollinated progenies may be more variable than outcrossed progenies if rare, recessive alleles are important contributors to genetic variances. Electrophoretic markers indicate that progeny derived from CH flowers are predominantly outcrossed (at least 54-97%).     

ENHANCEMENT OF INBREEDING DEPRESSION BY DOMINANCE AND SUPPRESSION IN IMPATIENS CAPENSIS

We investigated the effect of intraspecific competition on the magnitude of inbreeding depression in Impatiens capensis by planting seeds from chasmogamous (CH) and cleistogamous (CL) flowers in three experimental greenhouse treatments: in individual pots, in flats in dense pure stands according to seed type, and in flats with the two seed types intermixed in a checkerboard array. The size distributions of plants grown in flats were significantly more hierarchical than those of plants grown individually, indicating that larger plants competitively suppressed smaller plants in the high-density treatments. The magnitude of inbreeding depression at high density depended upon the planting arrangement of CL and CH seeds. CH advantage was greatest when CH and CL seedlings were grown in competition with one another, suggesting that fitness differences between outcrossed and inbred individuals were intensified by dominance and suppression. For plants grown individually, the effects of maternal parent, seed weight, and emergence date on seedling size disappeared with plant age, whereas at high density these effects remained at the final harvest. Thus, plant density may influence patterns of natural selection both on mating system and on juvenile traits in natural Impatiens populations.     

浙江凤仙花属一新种

本文发表了浙江凤仙花属一新种即泰顺凤仙花Impatiens taishunensis Y.L.Chen et Y.L.Xu, sp.nov.。     

河南凤仙花属一新种

本文发表了河南凤仙花属一新种,即奇异凤仙花Impatiens paradoxa C.S.Zhu et H.W.Yang,sp.nov.     

PATTERN OF PHENOTYPIC VIABILITY AND FECUNDITY SELECTION IN A NATURAL POPULATION OF IMPATIENS PALLIDA

Estimates of viability and fecundity selection of 13 phenotypic characters for 1,536 individuals of Impatiens pallida growing in 24 locations within a single natural population were compared. Directional viability selection of cotyledon area, day of initial leaf production, number of leaves, and stem length was detected throughout this population. Directional fecundity selection of cotyledon area, day of initial flower production, number of leaves present on day of initial flower production, stem length on day of initial flower production, number of leaves, and stem length was also detected. Phenotypic selection of these characters was strong in some cases, and the strength of selection was significantly heterogeneous among locations. For several of the characters, directional phenotypic selection within the population was significantly positive in some locations and significantly negative in others separated by only a few meters. Fecundity selection was stronger than viability selection for some characters, implying that fecundity selection was at least as important as viability selection within this population. Soil moisture levels and light intensities played a larger role than soil nutrient levels in determining the patterns of both viability and fecundity selection, and differences in directional viability selection were more strongly related to environmental variation than were differences in fecundity selection. The pattern of phenotypic selection could not be reliably inferred from the patterns of mortality and reproduction alone.     

GENETIC CONSEQUENCES OF OUTCROSSING IN THE CLEISTOGAMOUS ANNUAL,IMPATIENS CAPENSIS. I. POPULATION-GENETIC STRUCTURE

We examined the genetic structure of 11 populations of Impatiens capensis, a cleistogamous annual herb, using starch gel electrophoresis. We sampled both cleistogamous (CL) and chasmogamous (CH) progeny (if present) from maternal parents in each population to infer maternal genotypes and to estimate the extent and pattern of inbreeding within and among populations. Only eight of 31 loci were polymorphic, with one to six (mean = 3.1) loci varying within each population. Mean heterozygosity per individual is quite low (mean = 3.9%) and comparable to highly self-fertilized species. Gene flow is low, and genetic distances do not parallel geographical distances, suggesting a population structure similar to Wright's Island model with drift among the populations. Fixation indexes within populations (f? or F_IS) span the largest range yet reported for a plant species (0.26 to 0.94, mean = 0.57). Further inbreeding results from population substructuring , resulting in a total average inbreeding coefficient (F? or F_IT) of 0.77. Despite these high overall levels of inbreeding, chasmogamy significantly reduces fixation, which may account for the observed greater fitness of CH progeny.     

REACTION NORMS OF MORPHOLOGICAL AND LIFE-HISTORY TRAITS TO LIGHT AVAILABILITY IN IMPATIENS CAPENSIS

For plants, light availability is an important environmental factor that varies both within and between populations. Although the existence of sun and shade “ecotypes” is controversial, it is often assumed that trade-offs may exist between performance in sun and in shade. This study therefore investigated variation in reaction norms to light availability within and between two neighboring natural populations of the annual Impatiens capensis, one in full sun and the other in a forest understory. Seedlings were collected randomly from both populations and grown to maturity in a greenhouse under two light conditions: full light and 18% of full light. Selfed full-sib seed families were collected from plants from both populations grown in both parental light environments. To characterize family reaction norms, seedlings from each family were divided into the same two light treatments and individuals were scored for a variety of morphological and life-history traits. The maternal light environment had little impact on progeny reaction norms. However, the two study populations differed both qualitatively and quantitatively in plastic response to light availability (indicated by significant population x environment interactions in mixed-model ANCOVA). Much of this difference was attributable to population differences in light sensitivity of axillary meristem allocation patterns, which produced concurrent differences in reaction norms for a suite of developmentally linked traits. Within each population, different sets of traits displayed significant variation in plasticity (indicated by significant family x environment interactions). Thus, the genetic potential for evolutionary response to selection in heterogeneous light environments may differ dramatically between neighboring plant populations. Between-environment genetic correlations were largely positive in the woods population and positive or nonsignificant in the sun population; there was no evidence for performance trade-offs across environments or sun or shade “specialist” genotypes within either population. There was little evidence that population differences represented adaptive differentiation for sun or shade; rather, the results suggested the hypothesis of differential selection on patterns of meristem allocation caused by population differences in timing of mortality and intensity of competition.