A gradual reduction of female structures in the pistillate flowers ofRicinus communis L. (castor-bean) with chlorflurenol (morphactin)

Chlorflurenol (morphactin) EMD 7301 W (60, 120 and 240 mg I^?1) when applied at five to seven leaf and 10–12 leaf stage induced hermaphrodite flowers in the inflorescences ofRicinus communis. The hermaphrodite flowers were mostly apical in position unlike the apical female flower in the control. With an increase in the time gap between sowing and emergence of the inflorescence there was a reduction of the female structures in the hermaphrodite flowers. The last formed inflorescences had an apical male flower in place of a female. The largest number of inflorescences with an apical male or hermaphrodite flower were produced with 60 mg 1^?1 applied at five to seven leaf stage. The hermaphrodite flowers failed to set fruits.     

Effects of some natural and synthetic growth regulators on sex-expression of Myrica esculenta Buch-Ham. ex D. Don

Gibberellic acid (GA,), chlorflurenol and ethrel were applied at different concentrations to either male or female trees of Myrica esculenta. GA₃ induced intersexual flowers both on the male and female trees whereas chlorflurenol and ethrel induced similar (intersexual) flowers but only on the male plants. When GA₃ was applied in combination with chlorflurenol, fewer intersexual inflorescences appeared on the male plants along with a significant decline in the number of flowers per inflorescence. In combination, chlorflurenol and ethrel induced copious female and intersexual inflorescences per male plant, especially when two successive treatments of 100 mg/I of chlorflurenol and 1920 mg/l of ethrel were applied. The induced female inflorescences later bore fruits.     

INFLORESCENCE ARCHITECTURE AND FLOWER SEX RATIOS IN SAGITTARIA BREVIROSTRA (ALISMATACEAE)

Two populations of Sagittaria brevirostra from the same lake were sampled 10 years apart and yielded similar data on inflorescence structure and on numbers and ratios of male and female flowers. Larger inflorescences have relatively more male than female flowers than do smaller inflorescences. Pollination success is unrelated to inflorescence size or sex ratio within an inflorescence.     

Hydroxycinnamoyl acid amides and aromatic amines in the inflorescences of some araceae species

Hydroxycinnamoyl acid amides (HCA's) were found to be important components in the inflorescences of different Araceae species. HCA's occurred in large amount in spathes and in the male and female flowers, and were totally absent from the sterile flowers, commonly found on Araceae spadices. Differences in the distribution of HCA's were noted between male and female flowers. Thus the amount of neutral HCA's was always greater in the male than in the female flowers and the female flowers generally contained more basic HCA'S. In the inflorescences of some Araceae species in the Monsteroideae and Philodendroideae (genera Monstera, Raphidophora and Philodendron), the aromatic amines tyramine and dopamine were very abundant, with concentrations ranging from 1 to 4 mg of each amine per g fr. wt.     

Observations on the reproductive ecology of Thalassia testudinum (Hydrocharitaceae) II. Leaf width as a secondary sex character

Short-shoots of Thalassia testudinum Banks ex König were collected from 5 sites in Florida. A total of 284, 625-cm² quadrat samples, containing 6182 shhort-shoots was analyzed for leaf width, inflorescence number and sex (the latter if determinable). Although leaf widths and reproductive densities differed at the 5 sites, leaf width was consistently greater when reproductive structures were present, and when the number of inflorescences increased. The mean number of inflorescences per short-shoot was significantly higher for shoots bearing male inflorescences compared to female shoots. Female inflorescences were normally solitary; male short-shoots usually had 2 or 3 inflorescences. Sex ratios were male-biased for 4 of the 5 sites. Comparisons of leaf widths between the sexes indicated that leaf width constituted a secondary sex character for this species. Female short-shoots tend to have narrower leaves than male short-shoots. This relationship should also be considered when evaluating the significance of morphogeographic and stress-related variation in leaf width for this species.     

Tests of a mechanistic model of one hormone regulating both sexes in Buchloe dactyloides (Poaceae)

A mechanistic model of one hormone regulating both sexes in flowering plants was tested in buffalograss (Buchloe dactyloides). This model assumes that one hormone has male and female cell receptors to inhibit one sex and induce the other independently. Three components—the normal range of hormone level in the plant and the sensitivity levels of the two receptors—interact to regulate sex expression. The study organism, buffalograss, is usually considered dioecious, but natural populations consist of varying proportions of male, female, and monoecious plants. Prior research with growth regulators had shown that only gibberellin (GA) had consistent and significant effects on sex expression in this species. To test the model assumption of a hormone with a dual function, GA and a GA inhibitor (paclobutrazol, PAC) were applied to three monoecious genotypes; in two of the genotypes the GA treatment yielded a significantly higher proportion of male inflorescences, and this transition involved both inducing male and inhibiting female. PAC treatment produced exclusively female inflorescences, illustrating the dual effects of GA. To test the predictability of the model, GA was applied to two dwarf female genotypes. These plants were transformed into neuter and near-neuter plants with normal height and vegetative growth, as predicted by our model for genotypes with a physiologically wide overlapping of male and female sterile regions. The model also predicts that male or female plants would be induced to produce inflorescences of the other sex if the hormone level could be shifted from one side of the overlapping sterile regions to the other. This was verified by applying high levels of GA to a normal female genotype that resulted in the production of male inflorescences. However, this is the only normal female that has responded to GA application by producing male inflorescences, and males lose vigor and/or die without producing female inflorescences at high levels of PAC. The model suggests that the constancy of these males and females is due to the relative location of the sensitivity levels in relation to each other and to the hormone range. We conclude that the one-hormone model can facilitate both applied and basic research.     

Water Deficit and Inflorescence Development in Zea mays L.--The Role of the Developing Tassel

In the normal pattern of development of Zea mays (cv. Iochief)a single mature female inflorescence is produced at node 7.A brief episode of water deficit at the time of terminal maleinflorescence initiation induced the subsequent developmentof two to three mature female inflorescences at nodes 5–7.This growth of the inflorescences at lower nodes was accompaniedby a marked inhibition of the growth of the terminal male inflorescence.Removal of either the developing terminal inflorescence or ofthe axillary inflorescence at node 7 at this time also promotedthe growth of the lower axillary inflorescences. The growthof these inflorescences was further stimulated by a period ofwater deficit when only the inflorescence at node 7 was removed,but removal of the male inflorescence abolished the capacityof these inflorescences to respond to the water deficit Excisionof the male inflorescence immediately before or immediatelyafter the period of water deficit produced the same response.It is concluded that this response of the lower axillary inflorescencesto water deficit is mediated through an effect on the developingterminal male inflorescence. Zea mays, water deficit, inflorescence development, tassel, correlative inhibition     

Flower choice by honey bees (Apis mellifera L.): sex-phase of flowers and preferences among nectar and pollen foragers

Bees foraging for nectar should choose different inflorescences from those foraging for both pollen and nectar, if inflorescences consist of differing proportions of male and female flowers, particularly if the sex phases of the flowers differ in nectar content as well as the occurrence of pollen. This study tested this prediction using worker honey bees (Apis mellifera L.) foraging on inflorescences of Lavandula stoechas. Female flowers contained about twice the volume of nectar of male flowers. As one would predict, bees foraging for nectar only chose inflorescences with disproportionately more female flowers: time spent on the inflorescence was correlated with the number of female flowers, but not with the number of male flowers. Inflorescence size was inversely correlated with the number of female flowers, and could be used as a morphological cue by these bees. Also as predicted, workers foraging for both pollen and nectar chose inflorescences with relatively greater numbers of both male and female flowers: time spent on these inflorescences was correlated with the number of male flowers, but not with the number of females flowers. A morphological cue inversely associated with such inflorescences is the size of the bract display. Choice of flowers within inflorescences was also influenced predictably, but preferences appeared to be based upon corolla size rather than directly on sex phase.     

The mating consequences of sexual segregation within inflorescences of flowering plants

Many co-sexual plants segregate female and male function among flowers on an inflorescence through dichogamy or the production of unisexual flowers. Sexual segregation may reduce self-pollination among flowers within inflorescences (geitonogamy), thereby increasing the pollen available for export to other plants. To assess these complementary roles we manipulated the simultaneously hermaphroditic (adichogamous) flowers of Eichhornia paniculata to produce ten-flowered inflorescences with either female above male flowers (female/male inflorescences) or male/female inflorescences, which competed for mating opportunities with five-flowered adichogamous inflorescences. Because of the upward movement of bumble-bees, selfing increased upward in adichogamous inflorescences (overall female selfing rate s+/-s.e.=0.320+/-0.026). Female flowers of male/female inflorescences selfed less than flowers in corresponding positions in adichogamous inflorescences so s fell to 0.135+/-0.027. In contrast, all-female flowers of female/male inflorescences selfed similarly to upper flowers on adichogamous inflorescences, elevating s (0.437+/-0.043). During 1997, male/female inflorescences sired more outcrossed seeds than female/male or adichogamous inflorescences, whereas during 1994 flowers on male/female inflorescences received fewer visits than those of adichogamous inflorescences, reducing their outcross siring success. Hence, sexual segregation limits geitonogamy and enhances outcross siring success when it does not affect pollinator behaviour, illustrating the importance of both female and male function in inflorescence design.     

The floral biology and reproductive system of <Emphasis Type="Italic">Mauritia flexuosa</Emphasis> (Arecaceae) in a restinga environment in northeastern Brazil

Studies of the floral biology of the buriti palm, Mauritia flexuosa, have presented conflicting results with respect to the mechanism of pollination, indicating either cantharophily or anemophily. To resolve this question, the floral biology of M. flexuosa was studied in a coastal restinga environment in northeastern Brazil. The reproductive system was studied experimentally, and floral visitors were collected by bagging inflorescences. In this environment, M. flexuosa, a dioecious species, has several gender-specific floral features that function to attract pollinators, especially beetles. The male flowers produce large amounts of pollen as a reward, and male and female inflorescences produce similar odors that attract pollinators to female flowers, which offer only a nectar secretion as a reward. When feeding on the female flowers, the visitors frequently come into contact with the stigmata. To increase the chances of pollination, the female flowers persist longer than the male ones, and the viability of the pollen grain is very high. A curculionid beetle species of the genus Grasidius was found to be an effective pollinator. We suspect that wind also contributes to the pollination of M. flexuosa in the study area, but in a relatively minor way.     

FLORAL DISPLAY IN PHYLA INCISA: CONSEQUENCES FOR MALE AND FEMALE REPRODUCTIVE SUCCESS

The inflorescences of Phyla incisa consist of flowers in two phases: younger, nectar-containing flowers that have yellow corolla throats and older, nectar-lacking flowers that have dark purple corolla throats. Observations of pollinator visitation patterns to both natural and manipulated inflorescences were made to determine the role of each flower phase in pollinator attraction. The effect of older-phase flowers on male and female reproductive success was determined by comparing stigmatic pollen loads and estimates of pollen removal from inflorescences having different numbers of these flowers. The pollinators of Phyla selected larger inflorescences more often than expected based upon the size distribution of inflorescences available to them. Both younger- and older-phase flowers contributed to the attraction of pollinators, but the latter were less effective in this function. The presence of older-phase flowers significantly increased the visitation rate to inflorescences and the amount of pollen removed but had little effect on pollen deposition on stigmas. The lack of correspondence between pollen deposition and pollinator-visitation rate was not due to stigma saturation, since stigma loads varied greatly. The data indicate that the deposition of pollen on stigmas in this species is a relatively stochastic process, whereas pollen removal from inflorescences occurs at a much more regular rate. Old-phase flower retention appears to contribute to reproductive success through increased pollen donation when pollinator activity is high and may also increase the probability of seed set when pollinators are rare.     

Size and sex of floral displays affect insect visitation rates in the dioecious Australian alpine herb, Aciphylla glacialis

Differences in floral display size between male and female plants in dioecious species are often considered to be the result of competition within and between male and female plants to attract insects, although this hypothesis is rarely tested. In a field study of the alpine dioecious herb Aciphylla glacialis , insect visitation rates were found to increase with increasing floral display, as predicted. Dipterans were the most common visitors to male and female inflorescences, both in terms of individuals visits (2083 out of 2581 total visits by insects), and species number (20 morphospecies, potentially 22 species). Larger male inflorescences attracted 1.7 times as many insects in total, and 3.1 times more visits by the most common flower visitor, Musca vetustissima than did female inflorescences, but equal number of visits by the next most common flying visitors, Poecilohetaerus aquilius and Tephritis poenia. There was no difference in the amount of time M. vetustissima individuals spent on male and female inflorescences, even though male inflorescences have more flowers. Larger displays (four and eight inflorescences) of either sex attracted more visits than smaller ones (one inflorescence), although the rate of increase in visitation rate with increasing floral display declined. The response of insects to increasing floral display does not support the predicted escalating or proportionate increasing rate of visitation of some models for the evolution of dimorphism in floral displays in dioecious plants.     

Plant design and non-random foraging by ants on Croton bonplandianum Baill. (Euphorbiaceae)

Croton bonplandianum bears female flowers in inflorescences which are arranged in a regular pattern. Foraging individuals of the ant Camponotus sericius feed on the nectar in the nectary glands of female flowers. The pattern of movement of ants while foraging between these regularly placed inflorescences was highly systematic and distinctly exploratory. The ants tended to visit new inflorescences at each move and avoided revisiting inflorescences. They followed this foraging pattern by adopting circular movements either to the left or to the right and avoiding other complex but equally effective moves. This was probably due to memory constraints. The ants predicted the resource quality of the subset of a patch by sampling a part of it. The inflorescences within a cluster were similar in quality. Using this relation ants could make decisions about continuing to forage in a cluster after sampling any one inflorescence. If the first inflorescence visited was of poor quality they abandoned the cluster. This ensured efficient foraging with minimal expenditure of time and energy.     

Phenylacetaldehyde attracts male and female apple clearwing moths,Synanthedon myopaeformis,to inflorescences of showy milkweed,Asclepias speciosa

Synanthedon myopaeformis Borkhausen (Lepidoptera: Sesiidae) is a diurnal clearwing moth native to Eurasia that was recently introduced into British Columbia (BC) and Ontario, Canada, where it has become a serious pest in apple orchards. In BC, these moths commonly feed on nectar of inflorescences, particularly that of showy milkweed, Asclepias speciosa Torrey (Apocynaceae). We investigated the relative importance of visual and olfactory cues, and the key floral semiochemical(s) mediating attraction of S. myopaeformis to A. speciosa. In field experiments, inflorescences left exposed or enclosed in cheesecloth bags dyed green induced similar visitation rates by moths, indicating that olfactory cues are attractive. Among the >10 floral odourants that elicited responses from moth antennae in coupled gas chromatographic‐electroantennographic detection analyses, phenylacetaldehyde induced the most frequent proboscis extension reflexes of male and female moths. Among eight floral odourants that were field‐tested singly, phenylacetaldehyde attracted 35 times more male and female moths than any other candidate semiochemical. Attractiveness of phenylacetaldehyde could not be enhanced by admixture with other floral odourants at the ratios or concentrations tested indicating that it alone may mediate attraction of S. myopaeformis to the inflorescences of A. speciosa. The potential use of phenylacetaldehyde as bait to monitor or mass‐trap populations of male and female S. myopaeformis should be investigated.     

The distribution of sex inMyrica gale

The distribution of male and female inflorescences among stems ofMyrica gale L. is strongly bimodal and indicates a genetical dimorphism in gender. Predominantly or strictly male stems are about twenty times as frequent as predominantly or strictly female stems. There is considerable variation among male and female stems in their precise gender. Quantitative values of the phenotypic sex (the observed gender based on the relative proportions of male and female inflorescences) and functional sex (the relative probabilities of transmitting genes to descendants through male and female gametes) of samples of stems are estimated.Sexual Strategies in Plants, V.     

Floral and seedling morphology and anatomy of Thalassodendron pachyrhizum den Hartog (cymodoceaceae)

Thalassodendron pachyrhizum den Hartog is dioecious with inflorescences on short laterals from upright stems. The male inflorescence consists of two flowers which are morphologically identical but developmentally different. Each male flower has two laterally fused anthers, each of which contains four loculi surrounding a vascular bundle. Filiform pollen grains are arranged in coils. The walls of pollen grains contain cellulosic microfibrils embedded in a protein and carbohydrate matrix, and lack an exine layer. The female inflorescence produces two morphologically and developmentally identical flowers, each having an ovary with a short style containing two vascular bundles and leading to two long, slender stigmas. Both male and female inflorescences are enclosed in several alternating bracts. The innermost bract differs from the others by lacking a ligule. Squamulae intravaginales are present in all inflorescences. In each inflorescence, only one ovary develops into the seed which germinates on the parent plant. Young seedlings have an aril-like structure which disappears at a later stage of seedling development. The seedling produces, firstly, an aberrant seedling leaf and a scarious seedling sheath, then several true foliage leaves and finally several root primordia. The mature seedling separates from its protecting bract and detaches from the parent plant. The floral and seedling morphology and anatomy are compared with other closely related genera in the Cymodoceaceae and unique features are assessed. The frequency of floral and seedling production is discussed in relation to the distribution of T. pachyrhizum.     

Sexual specialization in two tropical dioecious figs

Ficus species (figs) and their species-specific pollinator wasps are involved in an intimate mutualism in which wasps lay eggs in some ovaries of the closed inflorescences (syconia), and mature, inseminated offspring carry pollen from mature syconia to fertilize receptive inflorescences. In monoecious species, each syconium produces seeds and wasps. In functionally dioecious fig species, making up approximately half the figs worldwide, male and female functions are separated; hermaphrodite (functionally male) trees produce wasps and pollen only, while female trees produce seeds only. This sexual separation allows selection to act independently on the reproductive biology of each sex. Examining sexual specialization in a tight mutualism allows us to determine aspects of the mutualism that are flexible and those that are canalized. In this study, we quantified the phenology of two species of dioecious figs, F. exasperata and F. hispida, for 2 years by following the fates of several thousand syconia over time. In studying each of these species in a dry and a wet site in south India, we tested specific predictions of how dioecious figs might optimize sexual function. On female trees of both species, more inflorescences matured during the wet (monsoon) season than in any other season; this fruiting period enabled seeds to be produced during the season most suitable for germination. In F. exasperata, functionally male trees released most wasps from mature syconia in the dry season, during peak production of receptive female syconia, and thus maximized successful pollination. In F. hispida, “male” trees produced more syconia in the dry and monsoon seasons than in the post-monsoon season. In both species, male and female trees abscised more unpollinated, young inflorescences than pollinated inflorescences, but abscission appeared to be more likely due to resource- rather than pollinator- limitation. The phenology of F. exasperata requires that male inflorescences wait in receptive phase for scarce pollinators to arrive. As expected, male inflorescences of this species had a longer receptive phase than female inflorescences. In F. hispida, where pollinators are rarely scarce, duration of receptive phase was the same for both sexes. Duration of developing phase was longer in female syconia of both species than in male syconia, most likely because they need a longer period of investment in a fleshy fruit. Variation in developing phase of female syconia in one species (F. exasperata) was also greater than that in male syconia, and enabled female trees to sample a variety of germination environments in time. The strong sexual differences in both fig species support the hypothesis that selection for sexual specialization has strongly influenced the reproductive biology of these species. Received: 28 May 1997 / Accepted: 2 February 1998