ANDROMONOECY IN ZIGADENUS PANICULATUS (LILIACEAE): SPATIAL AND TEMPORAL PATTERNS OF SEX ALLOCATION

I describe patterns of sex allocation and gamete packaging in the andromonoecious lily Zigadenus paniculatus. In this species, pistil length was continuously, but bimodally, distributed within plants, and smaller pistils contained fewer mature ovules. In hermaphrodite flowers, ovule number per flower increased with blooming rank in small plants but decreased with blooming rank in large plants. Flowers with pistils less than three-fourths stamen length almost never produced fruits and were classified as males. The pedicel, tepals, stamens, and pistil of hermaphrodite flowers were all heavier than those of males. Hermaphrodite flowers were concentrated on the terminal raceme, males on the lower racemes. In combination with acropetal blooming, this spatial separation of flower types resulted in a seasonal decline in the proportion of open flowers that were hermaphrodite. However, individual flowers were protandrous, so that the population sex ratio, initially strongly male-biased, declined as the season progressed. Hand pollinations showed that plants were self-incompatible. Inflorescence size was positively correlated with bulb size, and plants with large inflorescences had a higher proportion of male flowers. Nutrient supplementation had no effect on inflorescence size, but increased the proportion of hermaphrodite flowers. Nutrient-supplemented plants also began blooming earlier than controls. I discuss these patterns in relation to the adaptive significance of andromonoecious breeding systems.     

CHANGES IN PLASTID AND MITOCHONDRION CONTENT DURING MATURATION OF GENERATIVE CELLS OF SOLANUM (SOLANACEAE)

A study was made of the number of plastids and mitochondria present in generative cells of Solanum immediately after microspore mitosis, and the fate of these organelles during development of the pollen was determined. Changes were followed via electron microscopy of anthers of S. chacoense and S. tuberosum Group Phureja × S. chacoense. In earliest stages the generative cells were oval and had one surface along the intine and other surfaces in contact with the vegetative cell. As the pollen matured the generative cells elongated, became spindle-shaped, and were completely engulfed in the vegetative cells. At the earliest stages studied, both mitochondria and plastids were present in the generative cell. Plastids of the generative cell were, in contrast to those of the vegetative cells, fewer, smaller, and lacking in starch. Through the maturation stages the content of these organelles in the vegetative cells remained unchanged. While the generative cells retained mitochondria until anthesis, their plastids disappeared completely during maturation. This selective loss during generative cell maturation could lead to transmission of those characteristics encoded in plastid DNA through the pistillate parent only. The mechanism could explain earlier genetic evidence that plastid characters of Solanum were transmitted uniparentally.     

KARYOTYPIC STUDIES IN SOLANUM SECTION BASARTHRUM (SOLANACEAE)

Mitotic chromosomes of 450 individuals, representing 59 accessions of 18 of the 22 species of Solanum sect. Basarthrum were studied. Statistical analyses of chromosome length, genome length, and centromere position yielded estimates of karyotype composition and asymmetry. A generalized karyotype of the section shows that most of the chromosomes are metacentric (44%) or submetacentric (53%). A general trend emerged: the greater the total genome length, the more asymmetric the karyotype. However, karyotype asymmetry does not seem to be based on major chromosomal rearrangements. There is variation in species groups in the karyotype formula, and chromosome length differences were useful in distinguishing series. Patterns of cytological variation reinforce the systematic arrangement of the taxa based on morphological, chemical, and molecular studies. Chromosomal differences characterize most of the species. Five species bear subtelocentric chromosomes, two have telocentrics, and two have satellites. The genomes of the staminate and pistillate plants of the dioecious S. appendiculatum are identical. Of the species proposed as progenitors of the domesticated cultigen S. muricatum, S. caripense is the most similar karyotypically.     

THE EFFECT OF INVESTMENT IN ATTRACTIVE STRUCTURES ON ALLOCATION TO MALE AND FEMALE FUNCTIONS IN PLANTS

Expressions for male and female fitnesses of partially self-fertilizing cosexual plants are derived, assuming that allocation to pollinator attraction at the time of flowering may decrease resources available for male and female primary structures. The total female fertility is assumed to be controlled by factors at two stages, flowering-time and fruiting-time, with resources for fruit maturation being limited so that maximum seed production may be limited by the availability of these resources. The fitness formulas are used to calculate ESS (evolutionarily stable strategy) allocations at flowering time to primary male and female sex functions and to attractive structures. These are compared with some data that are available for dry weights of different flower parts. The fitnesses of unisexual mutant forms are calculated, assuming that they are introduced into a population consisting mostly of the initial cosexual form and that they obey the same gain curves as that form. When compared with the fitness of the ESS cosexual form, this enables one to ask whether unisexual forms will be favored. We show that the spread of females is unlikely, unless there is high inbreeding depression and a rather high selfing rate, and that in some circumstances a linear relation between number of fertilized ovules and number of seeds matured can be less favorable for the invasion of females than is a highly concave relation. With a nearly linear relation between numbers of fertilized ovules and mature seeds, invasion by females is more likely when investment in attraction is low than when it is high. These effects are discussed in relation to the distribution of dioecy. The spread of male mutants is never likely in these models.     

POLLEN-OVULE RATIOS AND BREEDING SYSTEM EVOLUTION IN SOLANUM SECTION BASARTHRUM (SOLANACEAE)

Pollen-ovule ratio, pollen grains per flower, pollen grain volume, pollen biomass, ovules per ovary, seeds per fruit, seed volume, seed biomass, and corolla size were studied to reveal patterns of variation within a monophyletic group. Solanum sect. Basarthrum includes 22 neotropical species of two clades, one having small, few-seeded red fruits, and the other having larger, many-seeded green fruits. The former includes self-incompatible species and a dioecious species, while the latter includes self-incompatible species, self-compatible (and autogamous) species, and a domesticate (the “pepino”). Although the pollen quantity and ovule quantity of the self-incompatible species are significantly higher in the green-fruited subgroup than in the red-fruited subgroup, the pollen-ovule ratios of the self-incompatible species are not significantly different between the two subgroups, suggesting parallel evolution of the pollen-ovule ratio. Furthermore, the pollen-ovule ratio tracks the breeding system: self-incompatible species have significantly higher pollen-ovule ratios than self-compatible species, resulting both from fewer pollen grains and more ovules of the latter. The pollen-ovule ratio of the dioecious species is among the highest of all, resulting only from fewer ovules. The pepino is self-compatible but has a pollen-ovule ratio like its wild self-incompatible progenitors and shows wide variation in seed production and pollen quality, but not in pollen quantity. Among all species, pollen size and quantity are inversely proportional, as are seed size and quantity.     

THE POLLINATION ECOLOGY OF SOLANUM ROSTRATUM (SOLANACEAE)

The pollination mechanisms and pollen vectors of Solarium rostratum have been examined by greenhouse experiments and field studies. Although the capacity for autogamy exists in this weedy annual, it rarely occurs because of two factors: (1) the morphology of the flower and (2) the foraging behavior of the various species of Bombus, the primary pollen vector in the regions studied. The percentages of geitonogamy and xenogamy are dependent on the flight pattern of the bees and the number of open flowers on a plant.     

DEVELOPMENT OF PLANTS FROM PROTOPLASTS OF SOLANUM (SOLANACEAE)

Experiments were performed to determine conditions critical to the isolation and culture of protoplasts from leaf mesophyll cells of a Solanum Section Tuberarium diploid clone, a S. phureja × S. chacoense F₁ hybrid. The optimum concentration of cellulase (Cellulysin) was 0.4%, while pectinase (Macerase) was inhibitory, even at a low concentration (0.075%). Maximum yields of protoplasts were achieved when the enzyme solution was not changed during incubation, and slow oscillation (60 rpm) on a shaker was used. When detached leaves were held under low light intensities or in the darkness for 3–5 days prior to protoplast production, results were more consistent than when the leaves were used directly from the greenhouse. Following dark or low light treatment the optimum osmolarity of the isolation and growth media was approximately 0.3–0.4 M. Of nine growth media tested only those of Nitsch and Ohyama and of Upadhya supported callus development, but the callus was often loose and did not differentiate roots or shoots. Callus from protoplasts cultured in the Upadhya medium modified by addition of glycine, vitamins, and casein hydrolysate, and subsequently transferred to the medium of Lam, formed roots and shoots when cultures were maintained in light. Mature plants were obtained following transfer to modified White's medium and later transplantation to soil.     

RESOURCE ALLOCATION AND REPRODUCTION IN POPULATIONS OF AMPHICARPUM PURSHII (GRAMINEAE)

Five New Jersey populations of Amphicarpum purshii (an annual panicoid grass) were investigated as to total allocation of biomass to reproduction and its distribution between aerial chasmogamous and subterranean cleistogamous inflorescences. Germination of both types of seed was documented, for the first time, in not only the field but under three laboratory temperature regimes. Overall, approximately 29 % of the shoot biomass was allocated to reproduction. On a population basis, subterranean inflorescences accounted for 37–100 % of the reproductive biomass, and these seed (5 × heavier but fewer in number) were the source of most surviving seedlings. Subterranean spikelets contained caryopses significantly more frequently than aerial spikelets. The ratio of the number of viable aerial seed to the number of viable subterranean seed increased from 0/4 to 4/2 with recentness and/or frequency of disturbance. As a pioneer species in secondary succession, Amphicarpum purshii produces a larger subterranean propagule, with greater seedling vigor and a higher probability of local reproductive success, and also a smaller aerial propagule in larger numbers with potentially greater genetic variability.     

OUTCROSSING IN THE WEED,SOLANUM PTYCANTHUM (SOLANACEAE): A COMPARISON OF AGRESTAL AND RUDERAL POPULATIONS

Levels of outcrossing, using a Pgm polymorphism, were estimated for individuals from ruderal and agrestal populations of the predominantly monomorphic weed Solanum ptycanthum, at the northern edge of its range, and compared to agrestal populations sampled from the southern portion of its distribution. Individuals were grown in agricultural habitats in Ontario. Outcrossing estimates of the northern agrestals were very low (< 3%) and did not differ from ruderal estimates. Southern agrestals exhibited much higher outcrossing rates (10%–17%). Of the floral traits measured, degree of stigma exsertion and style position showed interpopulation variation and correlated with outcrossing rate. Outcrossing in northern individuals was largely precluded by the complete overlap in female and male functions and stigmatic surfaces that were in direct contact with anther sacs as the flower opened. The observed flexibility in mating system may have played an important role in colonization success of 5. ptycanthum.     

PATTERNS OF RESOURCE ALLOCATION IN A DIOECIOUS CAREX (CYPERACEAE)

Allocation to vegetative growth and sexual reproduction was investigated throughout the growing season in the dioecious sedge, Carex picta, under natural conditions and following experimental manipulations. Measurements taken on unmanipulated plants showed that the sexes did not differ in the total amount of biomass they allocated to either growth or reproduction. The relatively equal investment in reproduction by the two sexes is contrary to other studies, the majority of which show greater investment in reproduction by females. Two features of the reproductive biology of C. picta may account for the equal investment: the fruit are relatively inexpensive because they are uniovulate and nonfleshy, and the stamens are relatively expensive because C. picta is wind pollinated. In contrast to the lack of differences in the amount of allocation, there were differences between the sexes in the timing of allocation to growth and reproduction: males allocated more to reproduction and less to growth up to the time of flowering, whereas females showed this pattern during the time of fruit maturation. Defoliation and inflorescence removal experiments showed that a trade-off within plants between growth and reproduction does exist. In addition, the defoliation experiment revealed a difference in the response of the two sexes: defoliated tillers on males showed a reduction in growth, whereas defoliated tillers on females did not. Overall, the data support the idea that differences in the timing of reproductive expenditure are as important as the amount of expenditure in determining many aspects of the life history strategies of the two sexes.     

ADVENTIVE ORGANOGENESIS FROM SOMATIC TISSUE CULTURES OF SOLANUM CAROLINENSE: ORIGIN AND DEVELOPMENT OF REGENERATED PLANTS

The development of adventitious shoot formation from cultured somatic tissues of Solanum carolinense was studied using light microscopy. For purposes of comparison, callus initiation and proliferation were also followed. When stem segments of this plant were cultured on medium supplemented with 6 mg/l benzyladenine (BA), cell division was first observed after 48 hr in the external phloem and inner cortex of the segments. This division activity gave rise to meristematic zones which subsequently formed shoot primordia within 6 days of culture. While organogenic potential appears to be limited to specific regions, all tissues of the explant were capable of callus formation when cultured on medium containing 3 mg/l 2,4-dichlorophenoxyacetic acid (2,4-D). In conjunction with a previous study, it appears that unlike somatic embryogenesis in this species, organogenesis does not require an intermediate callus phase for differentiation to occur.     

黄瓜雄花形成过程中心皮的发育

黄瓜(Cucumis sativus L.)为重要的经济作物,雌雄同株异花,是研究植物性别分化的经典材料。人们对黄瓜性别分化进行了广泛的研究。Astmon和Galun、任吉君和王艳对黄瓜性别分化的形态特征和器官发生进行了初步研究,表明黄瓜单性花分化和发育过程中经历了无性期、两性期和单性期,最终只有一种性别的性器官原基发育成有功能的性器官,从而形成单性花,而对单性花中未形成有功能器官的相反性别原基的研究报道甚少。我们对雄花发育过程进行了连续的形态学分析,并对不同时期雄花中的心皮进行了细胞计数和同工酶电泳分析,以期从性器官发育的角度探讨黄瓜性别表现的机理。     

FEMALE CHOICE AND SEXUAL SELECTION OF MALE WING MELANIN PATTERN IN PIERIS OCCIDENTALIS (LEPIDOPTERA)

Using field experiments, I examined the role of 13 melanin-pattern elements in mate choice by female Pieris occidentalis butterflies. Males that mated successfully differed significantly from unsuccessful males in the extent of marginal forewing melanization but not on the basis of the entire forewing pattern. Deletion of the marginal forewing melanin characters significantly reduced the mating success of manipulated males relative to controls under field conditions. Female choice in P. occidentalis may act to maximize divergence of male color pattern from that of its close relative Pieris protodice. Sexual selection for increased melanization of the marginal dorsal forewing is generated by the strong preference of females for males with such wings and may be constrained by correlations between forewing melanin characters that have different functional roles.     

TRACKING POLLEN FLOW OF SOLANUM ROSTRATUM (SOLANACEAE) USING BACKSCATTER SCANNING ELECTRON MICROSCOPY AND X-RAY MICROANALYSIS

A technique using micronized metal powders was developed for both general labeling of pollen and marking of individual pollen grains. After labeling, pollen flow is analyzed by the use of backscatter scanning electron microscopy and X-ray microanalysis. To test the efficiency and efficacy of the technique, we assessed differences in pollen distribution in Solanum rostratum, an enantiostylous species with dimorphic anthers which are putatively feeding and pollinating anthers. Pollen from each set of anthers was labeled using different micronized metal powders. We could not confirm the differentiation of functional anthers in S. rostratum. This technique provides an efficient and convenient method for tracking pollen movement within and between flowers, and anthers within a single blossom can be differentially marked.     

CYTOLOGICAL EXPRESSIONS OF A CYTOPLASMIC MALE STERILITY IN SOLANUM

Anther contents of plants expressing the cytoplasm-gene interaction that results in anther indehiscence were studied under the light microscope. Plants could be classified in four groups on the basis of anther content: (1) A blockage resulted from cessation of development principally during the stages of meiosis of apparently normal sporocytes. This produced anthers in open flowers that usually contained monads or dyads. (2) Disorganization of sporocytes during first meiotic prophase resulted in irregularity of disjunction during the two meiotic divisions. Sporocytes produced quartet-stage clumps having more than four cells. Microspores failed to grow beyond early stages of exine development. (3) Abnormal small pollen having a very thick exine. (4) Normal pollen present in anthers that lacked terminal pores. The variety of anther content types resulted from presence of modifying genes rather than from differing actions of genes that conditioned indehiscence itself. Expression was not modified by environmental fluctuations, and plants did not show chimeral sectors having changed anther contents or dehiscent anthers.     

VARIATION IN MALE REPRODUCTIVE INVESTMENT AND MALE REPRODUCTIVE SUCCESS IN WHITE SPRUCE

An experimental population composed of ramets of white spruce clones was studied to determine the influence of different levels of clonal male reproductive investment on clonal male reproductive success, where the male reproductive success of different clones in the population was defined as the proportion of the seed crop sired by each clone. We present a multilocus estimation procedure which provides unbiased estimates of the proportion of seed sired by each parental genotypic class from pollen gamete frequency data, whenever gametic segregation frequencies are known. Significant linear and quadratic components were found for the relationship between the number of male cones produced and the estimated proportion of the seed crop sired. A clone that produced an exceptionally large number of male cones, however, did not have the greatest male reproductive success. In general, the results support an assumption made in theoretical models of selection on mating system modifiers regarding the relationship between male investment and male reproductive success. They also support the hypothesis that male reproductive success in wind-pollinated species continues to increase substantially with increasing male reproductive investment.     

TEMPERATURE DETERMINATION OF MALE OUTCROSSING ABILITY IN A SIMULTANEOUS HERMAPHRODITE

Phally, a genital dimorphism found in some species of self-compatible simultaneous hermaphrodites, presents an opportunity to examine factors maintaining outcrossing within an animal species in the presence of recombination. Both aphallics and euphallics can self-fertilize but only euphallics develop a functional penis and prostate allowing them to donate sperm. Previous studies of phally in the gastropod Bulinus truncatus (Mollusca: Pulmonata) suggest that phally may be under direct genetic control in some populations and strongly influenced by environmental factors in others. Experiments reported here identify temperature as a cue affecting phally determination in two populations of B. truncatus. In both populations, a higher proportion of euphallics was produced at low temperature (22 ± 1°C) than at high temperature (30 ± 1°C). Temperatures experienced by parents did not affect the proportion of euphallics they produced. Instead, phally was sensitive to temperature during the egg stage postoviposition and during the hatchling stage; the relative influence of temperature before and after hatching varied between populations. The total number of hatchlings reaching maturity at high and low temperature did not differ, but at low temperature, snails took longer to hatch and mature, and had lower survivorship. Just as studies of environmental sex determination have shed light on selective pressures influencing sex ratio evolution, we suggest that temperature-sensitive phally determination may shed light on the selective pressures maintaining outcrossing in B. truncatus.     

ANDROMONOECY AND VARIATION IN PHENOTYPIC GENDER OF PASSIFLORA INCARNATA (PASSIFLORACEAE)

We studied the perennial vine Passiflora incarnata Linnaeus (Passifloraceae) in Alachua County, Florida, from May to August 1983 to determine the breeding system and investigate seasonal changes in phenotypic gender of individual plants. Passiflora incarnata is an obligate outcrosser, incapable of self-fertilization. The major pollinators were carpenter bees (Xylocopa sp.). The proportion of flowers setting fruit was not limited by pollinator service, but the weight of individual fruits and number of seeds/fruit was increased over naturally pollinated flowers by manually performing cross-pollination. Morphological differences in style position among flowers caused some flowers to function primarily as males and others to function as hermaphrodites. Although some of the flowers that functioned as males set fruit when manually cross-pollinated, the proportion of these male flowers capable of setting fruit was lower than the proportion of hermaphroditic flowers setting fruit when manually cross-pollinated. Further, some male flowers had atrophied ovary and styles and were completely incapable of setting fruit. Passiflora incarnata is thus functionally andromonoecious. The relative production of male versus hermaphroditic flowers varied among individuals and over the course of the flowering season. Unmanipulated plants in the population became increasingly male-biased in floral sex ratios as the reproductive season progressed. We attempted to modify phenotypic gender in experimental plants by limiting the ability of plants in some treatment groups to set fruit. Treatment groups significantly affected production of hermaphroditic flowers, but production of male flowers was not affected by treatment. Treatment did not significantly affect fruit weight, number of seeds per fruit or the percentage of hermaphroditic flowers that successfully matured fruit. These results suggest that andromonoecy in P. incarnata is a mechanism for adjusting allocation of reproductive effort to male and female function, and that maternal investment in this species is regulated primarily by varying production of hermaphroditic flowers.