FUNCTIONAL DIOECY AND ANDROMONOECY IN SOLANUM

Field and laboratory studies of 19 diclinous species endemic to Australia help to clarify the nature and evolution of andromonoecy, androdioecy, and dioecy in the genus Solanum. Ten species are andromonoecious; typically these species bear inflorescences with a single, large basal hermaphroditic flower and 12–60 distal, smaller staminate flowers. We suggest that the andromonoecious condition was derived from hermaphroditic-flowered ancestors in part by hemisterilization of flowers but largely by addition of staminate flowers. The resultant larger inflorescences are hypothesized to serve both to attract and to entrain pollinators, yielding more or higher-quality seed set in hermaphroditic flowers and/or greater dispersion of pollen from staminate flowers. We suggest that andromonoecy may also serve to reduce selling. Nine other species are morphologically androdioecious but functionally dioecious. In these species, staminate flowers, like those of the andromonoecious species, bear anthers with copious tricolporate pollen and a highly reduced gynoecium. The morphologically hermaphroditic flowers are functionally pistillate and borne singly in inflorescences, and they bear anthers with inaperturate pollen. The inaperturate pollen, although viable, never germinates and is hypothesized to be retained in pistillate flowers as a reward to pollinators in the nectarless Solanum flowers. All other species of Solanum studied with pollen dimorphism in which one pollen morph is inaperturate are also best treated as functionally dioecious. We conclude that there is no evidence for androdioecy in Solanum. A review of other families suggests that there is little support for this unusual breeding system in any other angiosperm group either. Preliminary analyses suggest that andromonoecy and dioecy are polyphyletic in Solanum. Furthermore, dioecy is as likely to have arisen from hermaphroditic as from andromonoecious ancestors.     

Pollen morphology and functional dioecy inSolanum (Solanaceae)

Dioecy has evolved independently several times in the large, mostly tropical genusSolanum. In all cases of dioecy inSolanum functionally male flowers have normal anthers, normal pollen and reduced stigmas while functionally female flowers have stigmas and anthers that appear normal but contain non-functional, usually inaperturate pollen. The inaperturate pollen has living cytoplasm, but apparently never germinates and it has been hypothesised that the pollen in these functionally female flowers is retained as a pollinator reward. Pollen morphology is compared in twelve of the thirteen known dioecious species ofSolanum, and some stages in the the development of inaperturate pollen in the anthers of functionally female flowers ofSolanum confertiseriatum of Western Ecuador are examined. Observations on the development and morphology of inaperturate pollen in functionally female flowers ofSolanum are related to hypotheses about the evolution of dioecy in the genus.     

FUNCTIONAL DIOECY IN ECHINOCEREUS COCCINEUS (CACTACEAE): BREEDING SYSTEM,SEX RATIOS,AND GEOGRAPHIC RANGE OF FLORAL DIMORPHISM

The breeding system of the hedgehog cactus, Echinocereus coccineus Engelm. (Cactaceae), was studied at two sites in the northern Chihuahuan Desert, New Mexico. Two distinct flower types, restricted to separate plants, were observed. One was hermaphroditic and the other male-sterile. Hand-pollinations and field observations showed them to be functionally male and functionally female, respectively. There were significant quantitative intersexual differences in floral morphology. The main pollinators of the populations studied are probably two species of halictid bee. Sex ratios based on the number of plants and flowers at the two sites did not differ significantly from 1:1. Significant differences in fruit set existed between functional female plants from the two sites. The floral dimorphism described here appears confined to eastern and northwestern populations, at the periphery of the range of E. coccineus. Floral dimorphism was not observed in three closely related species.     

Reproductive biology of the dioecious Canary Islands endemic Withania aristata (Solanaceae)

We confirmed functional dioecy of Withania aristata via field and greenhouse studies. Male flowers are significantly larger. Female flowers bear stamens with no pollen; males bear 220?000 grains. Stigmata of male flowers senesce in buds. Anatomical observations confirm more ovules in females and an ovarian nectary in both sexes. We detected nectar in female flowers in the greenhouse but found no nectar in males. Thus, males offer pollen and females nectar. Females bear large numbers of fruits and, infrequently, male plants bear few significantly smaller fruits with few seeds. Outcrosses of females (self crosses impossible without pollen) yielded fruits in young buds, older buds, and open flowers. Self crosses of male flowers succeeded only with very young buds. Although functionally dioecious, this species manifests self-compatibility; however, no fruits are produced autonomously. Bee species (Lassioglossum, Amegilla, Apis) visit flowers and mature buds. Bud visits in which bees force petal tips apart, coupled with self-compatibility, may explain infrequent fruit on males. Thus, dioecy in W. aristata seems to have evolved from self-compatible ancestors, that leaky dioecy may have been favored during colonization, and, that despite autogamy and a low floral visition rate, this endemic enjoys a high rate of reproductive success.     

PHENOTYPIC GENDER IN SEX CHANGING DWARF GINSENG,PANAX TRIFOLIUM (ARALIACEAE)

Dwarf ginseng (Panax trifolium L., Araliaceae) is a diphasic (“sex changing”) species in which one phase has staminate flowers and the other has hermaphroditic flowers. In order to determine the relative allocations of the hermaphroditic gender phase to male and female functions,variation in population gender phase ratios, pollen production and viability, and ovule and seed production were documented. Gender phase ratios are highly male-biased. Dwarf ginseng is self-compatible, and both gender phases have viable pollen capable of effecting fertilization. Males produce more flowers and more viable pollen per anther than hermaphrodites. The phenotypic gender of hermaphrodites is extremely female-biased; it is likely that hermaphrodites function essentially as females. Sexual selection may have a role in the evolution and maintenance of differences between the gender phases in allocation to male function.     

Functional Gynodioecy in Opuntia quimilo (Cactaceae), a Tree Cactus Pollinated by Bees and Hummingbirds

Abstract: The tree cactus Opuntia quimilo is one of three known gynodioecious cacti. Its flowers deviate from most Opuntias in features that are attributable to ornithophily: petals are shiny red in colour, and fleshy in consistency, a nectar chamber is present, and stamen seismonasty is lacking. Pollinators include large matinal bees (predominantly Ptilothrix tricolor and Megachile sp.) and hummingbirds ( Chlorostilbon aureoventris and Heliomaster furcifer ). Hummingbirds rarely visit other local Opuntias. Hummingbirds, which are more common in the afternoon, prefer female flowers whereas bees prefer hermaphroditic flowers. Female flowers have more dilute nectar than hermaphroditic flowers. Under experimental conditions female fertility is as high as that of hermaphrodites, however, seeds from females always result from cross-pollination and from more severe ovule selection (ovule number is higher in female flowers). Under natural conditions female plants are reproductively more successful than hermaphrodites. Known cases of bird pollination in Opuntia and the incidence of ornithophilic features in Opuntia and related genera are discussed.     

Evolution of resupination in Malagasy species of Bulbophyllum (Orchidaceae)

Resupination is the orientation of zygomorphic flowers during development so that the median petal obtains the lowermost position in the mature flower. Despite its evolutionary and ecological significance, resupination has rarely been studied in a phylogenetic context. Ten types of resupination occur among the 210 species of the orchid genus Bulbophyllum on Madagascar. We investigated the evolution of resupination in a representative sample of these species by first reconstructing a combined nrITS and cpDNA phylogeny for a sectional reclassification and then plotting the different types of inflorescence development, which correlated well with main clades. Resupination by apical drooping of the rachis appears to have evolved from apical drooping of the peduncle. Erect inflorescences with resupinate flowers seem to have evolved several times into either erect inflorescences with (partly) non-resupinate flowers or pendulous inflorescences with resupinate flowers.     

Reproductive ecology of dioecious Siparuna (Monimiaceae) in Ecuador–a case of gall midge pollination

The reproductive ecology of the dioecious genus Siparuna , belonging to the primitive family Monimiaceae, was investigated in Ecuador. Thirteen species were studied, and species growing at high altitudes (above 500 m) were compared with species growing at lower altitudes. The genus has specialized flowers pollinated by Cecidomyiidae (Asynapta sp. and other genera) that lay eggs chiefly in male flowers since these are more easily accessible than female flowers due to their morphology. Several traits ensure that pollen transfer is accomplished: 1. long flowering of individuals and single flowers; 2. specific floral morphology (unisexuality, landing platform, restricted pore) allowing the flowers to be visited by Cecidomyiidae seeking egg-laying sites, and 3. a low number of ovules that have to be pollinated in each flower. The low number of ovules per flower results in a high pollen-ovule (P/O) ratio even though pollen production in male flowers is only moderate.
High altitude species had larger flowers, higher pollen-ovule ratios, fewer ovules per female flower, and longer periods of anthesis. Gender differences were seen in the longer functionality of female flowers; the larger numbers of male flowers produced; and the average diameter at breast height (DBH) biased for one of the two sexes depending on the species investigated, with no clear trend in the size distributions of individuals of both sexes. Sex ratios departed significantly from unity both to male and female bias but usually were in favour of males. There were no general differences in the ratio of functional male to female flowers at the time of observation nor in the sizes of male and female flowers. The evolution of unisexual flowers in Siparuna can be explained as a result of the differential predation by larvae: unimportant in male flowers, destructive if occurring in female flowers.     

The roles of female and hermaphroditic flowers in the gynodioecious–gynomonoecious Silene littorea: insights into the phenology of sex expression

Some gynodioecious species have intermediate individuals that bear both female and hermaphroditic flowers. This phenomenon is known as a gynodioecious–gynomonoecious sexual system. Gender expression in such species has received little attention in the past, and the phenologies of male and female functions have also yet to be explored. In this study, we examined variations in gender patterns, their effects on female reproductive success and sex expression in depth throughout the flowering period in two populations. The studied populations of Silene littorea contained mostly gynomonoecious plants and the number of pure females was very low. The gynomonoecious plants showed high variability in the total proportion of female flowers. In addition, the proportion of female flowers in each plant varied widely across the flowering season. Although there was a trend towards maleness, our measures of functional gender suggested that most plants transmit their genes via both pollen and ovules. Fruit set and seed set were not significantly different among populations; in contrast, flower production significantly varied between the two populations – and among plants – with consequent variation in total seed production. Conversely, gender and sex expression were similar in both populations. Plants with higher phenotypic femaleness did not have higher fruit set, seed set or total female fecundity. The mating environment fluctuated little across the flowering period, but fluctuations were higher in the population with low flower production. We therefore conclude that the high proportion of gynomonoecious individuals in our studied populations of S. littorea may be advantageous for the species, providing the benefits of both hermaphroditic and female flowers.     

Multiple sex expressions in Cimicifuga simplex: dichogamy destabilizes hermaphroditism

Cimicifuga simplex (Ranunculaceae) has four types of individuals, differing in their sex expression: there are individuals with hermaphroditic, hermaphroditic and male (andromonoecious), only male, or only female flowers. Purely female individuals appear during the early part of the flowering period of the population, and males during the later part. Strong protandry in hermaphroditic flowers causes a shift in functional sex ratio in the population over time. This facilitates the evolution and maintenance of small proportions of unisexual individuals in the population. A hypothesis developed from this case of multiple sex expression predicts dichogamy as a near-universal feature of andro-and gynomonoecious, androdioecious, and gynodioecious species, and also specific temporal distributions for each sex form depending upon the form of dichogamy. Results of tests on large sets of species strongly support the relevance of this hypothesis.     

Polygamy in Atriplex halimus L. (Chenopodiaceae)

It appears that up to now the inflorescence and flower morphologies of Atriplex halimus have been described incompletely. This species has been classified as monoecious or dioecious. Numerous observations and ontogenic studies have pointed to types of flowers morphologically and functionally hermaphroditic, never described until now. One specimen of this species presents both unisexual, male and female flowers and bisexual flowers, so Atriplex halimus is polygam and more precisely trimonoecious. Observation of inflorescences reveals a structure based on the spike and the dichasium. The sex distribution along the inflorescence axis was studied and the existence of a physiological gradient controlling its expression is discussed.     

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.     

Andromonoecy in Solanum lycocarpum A. St. -Hil. (Solanaceae): Floral attributes,visitors and variation in sexual expression over time

Sexual expression in andromonoecious species—those in which a single individual can bear both staminate and hermaphroditic flowers—may vary among reproductive events in the same plant, among individuals and across populations. This variation influences, in turn, the individual contribution of hermaphroditic plants via male and female fitness functions (i.e., Lloydʼs phenotypic gender). However, temporal variation in sexual expression in andromonoecious species and its relationship with seasonal changes in climatic conditions remain poorly understood. Here we analyze floral attributes, visitors and variation in sexual expression in three populations of Solanum lycocarpum A. St. -Hil. Seasonality in the production of floral types, the mating system and floral visitors were also investigated. Hermaphroditic flowers produced more pollen grains, but the pollen of staminate flowers had higher viability. Only hermaphroditic flowers produced fruits, and ovules in staminate flowers were sterile. Solanum lycocarpum is mainly pollinated by large bees with the ability to vibrate flowers. Phenotypic gender varied throughout the year, and the seasonal production of staminate flowers is associated with the local climate. We suggest that the higher and seasonally variable relative abundance of staminate flowers compared to the low and uniform production of hermaphroditic flowers may be explained by (a) the very high energetic costs incurred in producing large fruits, which in turn make hermaphroditic flower production very costly, and (b) the potentially lower energy expenditure of the smaller staminate flowers with reduced pistils and non-viable ovules that allow them to rapidly respond to climate variability.     

Development of male and female flower in Asparagus officinalis. Search for point of transition from hermaphroditic to unisexual developmental pathway

Asparagus officinalis is a dioecious plant. The flowers start to develop as hermaphrodites and later become unisexual. In female flowers the stamens degenerate, while in male flowers the ovary stops growing without degenerating. We have examined young asparagus flowers using SEM and optical microscopy in order to determine the exact moment of transition from hermaphroditic to unisexual development. We defined 13 stages of development, starting from flower primordia up to completely mature flowers and labelled them with numbers from -6 to 7. The first five stages are fully hermaphroditic: a difference between sexes becomes visible at stage — 1 when the style begins to develop in female flowers. Degeneration of stamens in female flowers starts somewhat later. At the stage of transition, some differences between sexes also appear in the bidimensional polypeptide pattern of flowers. RNase activity shows a distinct peak at this stage (in female flowers only), probably related to stamen degeneration.     

Functional Monoecy Due to Delayed Anther Dehiscence: A Novel Mechanism in Pseuduvaria mulgraveana (Annonaceae)

Unlike most genera in the early-divergent angiosperm family Annonaceae, Pseuduvaria exhibits a diversity of floral sex expression. Most species are structurally andromonoecious (or possibly androdioecious), although the hermaphroditic flowers have been inferred to be functionally pistillate, with sterile staminodes. Pseuduvaria presents an ideal model for investigating the evolution of floral sex in early-divergent angiosperms, although detailed empirical studies are currently lacking. The phenology and pollination ecology of the Australian endemic species Pseuduvaria mulgraveana are studied in detail, including evaluations of floral scent chemistry, pollen viability, and floral visitors. Results showed that the flowers are pollinated by small diurnal nitidulid beetles and are protogynous. Pollen from both hermaphroditic and staminate flowers are shown to be equally viable. The structurally hermaphroditic flowers are nevertheless functionally pistillate as anther dehiscence is delayed until after petal abscission and hence after the departure of pollinators. This mechanism to achieve functional unisexuality of flowers has not previously been reported in angiosperms. It is known that protogyny is widespread amongst early-divergent angiosperms, including the Annonaceae, and is effective in preventing autogamy. Delayed anther dehiscence represents a further elaboration of this, and is effective in preventing geitonogamy since very few sexually mature flowers occur simultaneously in an individual. We highlight the necessity for field-based empirical interpretations of functional floral sex expression prior to evaluations of evolutionary processes.     

Observations of the floral biology ofPrasophyllum odoratum (Orchidaceae,Spiranthoideae)

Prasophyllum odoratum is a vernal, nectariferous, terrestrial orchid that flowers profusely six to eight months following cyclical fires that disrupt sclerophyll woodlands. The morphology of the column and pollinarium is indicative of taxa placed within the subfam.Spiranthoideae. The orientation of the pollinaria to the stigma appears to prevent mechanical self-pollination. Both cross- and self-pollination appear to be effected by insects that forage within the brightly-colored, scented, non-resupinate flowers. Ants and drosophilid flies remove nectar, but do not appear to transport pollinaria between flowers. The primary pollinators are polytrophic flies in the fam.Syrphidae and opportunistic male bees in the genusLeioproctus (Colletidae). Approximately 52% of the flowers on a raceme set seed. The comparatively short floral tube ofP. odoratum reflects the dependence of this species on short-tongued insects to effect successful dispersal of pollinaria.     

Subdioecy in Consolea spinosissima (Cactaceae): breeding system and embryological studies

The breeding system and the embryology of Consolea spinosissima, a tree-like opuntioid endemic to Jamaica, were investigated. Morphological and embryological studies revealed that the species is subdioecious, with three sexual morphs present in the 150 × 120 m plot studied at Hellshire Hills, Jamaica. The female morph has pistillate flowers with open stigma lobes, no pollen grains, and sets fruit. The male morph has cryptic staminate flowers with closed stigma lobes, viable pollen grains, and a nonfunctional gynoecium that does not set seed. The weak hermaphrodite morph has low fruit set and "perfect" flowers that superficially resemble the functionally staminate flowers of the male morph. These perfect flowers reach anthesis with viable pollen grains, with no or only a few functional ovules, and with the style supporting pollen tube growth. Embryological studies showed that the critical stage for sex determination occurs earlier in pistillate than in staminate and perfect flowers. Anthers of pistillate flowers abort prior to microspore tetrad formation, whereas ovules of the staminate and perfect flowers degenerate after the complete maturation of the embryo sac. Based on flower structure and embryological data, we hypothesize that the ancestor of C. spinosissima is/was hermaphroditic.     

The Development of Gynoecium after Anthesis and Fertilization in Eleutherococcus senticosus (Araliaceae)

The development status of gynoecia in Eleutherococcus senticosus flowers is different from that in ordinary plants. Female gametophytes of E. senticosus have not become mature until the 6th day after anthesis. On the 6th day, 82.25% of embryo sacs in female plants, and 67.25％ of those in hermaphroditic plants become mature, while the rest are sterile, immature or degenerated with no fertilized embryo sacs observed. At the same time, all embryo sacs degenerated and flowers withered in male plants. On the 7th day, a few embryo sacs in female and hermaphroditic plants start being fertilized. Accompanying the differentiation of embryo sacs, styles of female and hermaphroditic flowers start to expand and their nectaries become mature gradually. After the 4th or 6th day of anthesis, stigmatic papillae become conspicious and stigmata become white and open. In the meantime, the stigmata become receptive and the nectaries get active or reactive. By the 9th or 10th day, 40～65 ％ of embryo sacs in female plants and 25～41% of those in hermaphroditic plants have been fertilized. The whole process of fertilization in E. senticosus was observed. About 2 or 3 days after pollination, the two sperm nuclei start to fuse with the egg and the secondary nucleus. The fertilization of E. senticosus belongs to the premitotic type of syngamy. The essential process of the fusion of male and female nuclei during syngamy may be generalized as follows: (1) the contacting of male nucleus with the female one; (2) the fusion of nuclear membranes between the male and female nuclei; (3) the despiralization of male spireme and the appearance of male nucleolus inside the fertilized female nucleus; (4) the dispersion of male chromatin and the mergence with the female chromatin, which is the sign of completion of the fusion of the two nuclei. In addition, degeneration types of mature embryo sacs were observed. And typical polyspermy and a series of cases in which extra sperms enter the em-bryo sac are recorded.     

澜沧舞花姜繁殖生物学特性及其进化意义探讨

 舞花姜属（Globba）植物具有多样的繁殖策略。分布于西双版纳的澜沧舞花姜（Globba lancangensis）自然条件下既能结实,同时又能在花序下部产生珠芽。通过对澜沧舞花姜的花部性状、花粉活力、传粉昆虫的访花行为、结实状况、珠芽产量等多方面的观测,初步了解了澜沧舞花姜的繁殖生物学特性。结果表明,澜沧舞花姜具有雄花及两性花同株的性表达特征。雄花及两性花的花粉在中午12∶00以前活力很高,之后雄花的花粉活力急剧下降,而两性花的花粉到16∶00仍有萌发能力。澜沧舞花姜在花序上产生雄花及两性花,每个花序每天仅开少量的花,其中50%的时间仅开雄花或两性花,在整个种群中形成了一定程度的暂时性的雄花两性花异株现象,从而增加了异交的可能性。澜沧舞花姜雄花的存在为P/O低的两性花提供了花粉补贴,这使得它可以通过调节雄花与两性花比例来调节种群的P/O,从而保证一定的结实能力。人工自交与异交下结果率没有差异,但自交结实率显著低于异交结实率,表明澜沧舞花姜有明显的自交不亲和现象。排蜂（Megapis dorstata）和黄绿彩带蜂（Nomia strigata）是澜沧舞花姜的主要访花昆虫,其中排蜂是它的有效传粉昆虫。澜沧舞花姜可能通过雄花两性花同株与自交不亲和相结合来促进异交。