Programmed cell death promotes male sterility in the functional dioecious Opuntia stenopetala (Cactaceae)

Background and Aims

The sexual separation in dioecious species has interested biologists for decades; however, the cellular mechanism leading to unisexuality has been poorly understood. In this study, the cellular changes that lead to male sterility in the functionally dioecious cactus, Opuntia stenopetala, are described.

Methods

The spatial and temporal patterns of programmed cell death (PCD) were determined in the anthers of male and female flowers using scanning electron microscopy analysis and histological observations, focusing attention on the transition from bisexual to unisexual development. In addition, terminal deoxynucleotidyl transferase-mediated dUTP nick-end labelling assays were used as an indicator of DNA fragmentation to corroborate PCD.

Key results

PCD was detected in anthers of both female and male flowers, but their patterns differed in time and space. Functionally male individuals developed viable pollen, and normal development involved PCD on each layer of the anther wall, which occurred progressively from the inner (tapetum) to the outer layer (epidermis). Conversely, functional female individuals aborted anthers by premature and displaced PCD. In anthers of female flowers, the first signs of PCD, such as a nucleus with irregular shape, fragmented and condensed chromatin, high vacuolization and condensed cytoplasm, occurred at the microspore mother cell stage. Later these features were observed simultaneously in all anther wall layers, connective tissue and filament. Neither pollen formation nor anther dehiscence was detected in female flowers of O. stenopetala due to total anther disruption.

Conclusions

Temporal and spatial changes in the patterns of PCD are responsible for male sterility of female flowers in O. stenopetala. Male fertility requires the co-ordination of different events, which, when altered, can lead to male sterility and to functionally unisexual individuals. PCD could be a widespread mechanism in the determination of functionally dioecious species.     

Going to great lengths: selection for long corolla tubes in an extremely specialized bat–flower mutualism

In a hypothesis that has remained controversial since its inception, Darwin suggested that long-tubed flowers and long-tongued pollinators evolved together in a coevolutionary race, with each selecting for increasing length in the other. Although the selective pressures that flowers impose on tongue length are relatively straightforward, in that longer tongues allow access to more nectar, selective pressures that pollinators impose on flower length are less clear. Here, we test for such selective pressures in the highly specialized mutualism between the nectar bat Anoura fistulata, which can extend its tongue twice as far as other nectar bats, and Centropogon nigricans, which has flowers of a similar length (8–9 cm). We used flight cage experiments to examine the effects of artificially manipulated flower lengths on (i) bat behaviour and (ii) pollen transfer. Increased length produced longer visits, but did not affect the force bats applied during visits. In the second experiment, flower length increased both the male and female components of flower function: long male flowers delivered more pollen grains and long female flowers received more pollen grains. However, pollen transfer was not correlated with visit duration, so the mechanism behind differences in pollen transfer remains unclear. By demonstrating that bats select for increasing flower length, these results are consistent with the hypothesis that A. fistulata evolved its remarkable tongue in a coevolutionary race with long-tubed flowers similar to that envisioned by Darwin.     

Epigenetic control of sexual phenotype in a dioecious plant,Melandrium album

Melandrium album (syn.Silene latifolia) is a model dioecious species in which theY chromosome, present only in heterogametic males, plays both a male-determining and a strict female-suppressing role. We showed that treatment with 5-azacytidine (5-azaC) induces a sex change to androhermaphroditism (andromonoecy) in about 21% of male plants, while no apparent phenotypic effect was observed in females. All of these bisexual androhermaphrodites (with the standard male 24,AA +XY karyotype) were mosaics possessing both male and hermaphrodite flowers and, moreover, the hermaphrodite flowers displayed various degrees of gynoecium development and seed setting. Southern hybridization analysis with a repetitive DNA probe showed that the 5-azacytidine-treated plants were significantly hypomethylated in CG doublets, but only to a minor degree in CNG triplets. The bisexual trait was transmitted to two successive generations, but only when androhermaphrodite plants were used as pollen donors. The sex reversal was inherited with incomplete penetrance and varying expressivity. Based on the uniparental inheritance pattern of androhermaphroditism we conclude that it originated either by 5-azaC induced inhibition ofY-linked female-suppressing genes or by a heritable activation of autosomal female-determining/promoting genes which can be reversed, on passage through female meiosis, by a genomic imprinting mechanism. The data presented indicate that female sex suppression inM. album XY males is dependent on methylation of specific DNA sequences and can be heritably modified by hypomethylating drugs.     

A Sexually Dimorphic Corolla Appendage Affects Pollen Removal and Floral Longevity in Gynodioecious Cyananthus delavayi (Campanulaceae)

The floral traits of bisexual flowers may evolve in response to selection on both male and female functions, but the relative importance of selection associated with each of these two aspects is poorly resolved. Sexually dimorphic traits in plants with unisexual flowers may reflect gender-specific selection, providing opportunities for gaining an increased understanding of the evolution of specific floral traits. We examined sexually dimorphic patterns of floral traits in perfect and female flowers of the gynodioecious species Cyananthus delavayi. A special corolla appendage, the throat hair, was investigated experimentally to examine its influences on male and female function. We found that perfect flowers have larger corollas and much longer throat hairs than female flowers, while female ones have much exerted stigmas. The presence of throat hairs prolonged the duration of pollen presentation by restricting the amount of pollen removed by pollen-collecting bees during each visit. Floral longevity was negatively related to the rate of pollen removal. When pollen removal rate was limited in perfect flowers, the duration of the female phases diminished with the increased male phase duration. There was a weak negative correlation between throat hair length and seed number per fruit in female flowers, but this correlation was not significant in perfect flowers. These results suggest that throat hairs may enhance male function in terms of prolonged pollen presentation. However, throat hairs have no obvious effect on female function in terms of seed number per fruit. The marked sexual dimorphism of this corolla appendage in C. delavayi is likely to have evolved and been maintained by gender-specific selection.     

平基槭花性别分化的细胞形态学观察

平基槭为杂性花,雄花与两性花同株,本文对其花性别分化过程进行了细胞形态学观察。结果发现,在花性别分化的早期,雄花和两性花的花芽中雌、雄蕊原基均具备,只是在花芽发育到一定时期,雄花的雌蕊原基发生选择性败育,败育发生在大孢子母细胞减数分裂为4个大孢子时期。两性花的雌蕊可以正常膨大结实,雄蕊花药虽然可以形成二核花粉,但不能正常开裂,属于不育雄蕊。初步分析认为,两性花雄蕊花药不能正常开裂与花粉囊壁纤维层木质化程度低有关。     

Breeding system and bumblebee drone pollination of an explosively pollen‐releasing plant,Meliosma tenuis (Sabiaceae)

• Explosive pollen release is a mechanism used by some angiosperms that serves to attach pollen to a pollinator's body. It is usually adopted by species with zygomorphic tubular flowers and pollinated by birds and bees. The tree genus Meliosma (Sabiaceae, Proteales) has unique disc‐like flowers that are externally actinomorphic, but internally zygomorphic, and release pollen explosively.
• To elucidate the adaptive significance of explosive pollen release, we observed flowering behaviour, the breeding system and pollinator visits to flowers of the Japanese species Meliosma tenuis in a temperate forest. Flowers bloomed in June and were nectariferous and protandrous. Explosive pollen release was triggered by slight tactile stimuli to anther filaments or staminodes in male‐stage flowers.
• Because pollen cannot come into contact with the pistils enclosed by staminodes, M. tenuis is functionally protandrous. Artificial pollination treatments revealed that M. tenuis is allogamous. The dominant flower visitors were nectar‐seeking drones of the bumblebee species Bombus ardens (Apidae). The drones’ behaviour, pollen attachment on their bodies and fruit set of visit‐restricted flowers suggest that they are the only agent triggering the explosive pollen release mechanism, and are the main pollinator of M. tenuis.
• The finding that bumblebee workers rarely visit these flowers suggests that the explosive pollen release has another function, namely to discourage pollen‐harvesting bumblebee workers.

     

Can floral consumption by fish shape traits of seagrass flowers?

Seagrasses are marine flowering plants with hydrophilous pollination. This abiotic pollination by water assumes absence of flower-animal interaction, but animals can interfere in this process through consumption of reproductive structures. We studied predation on male flowers by fish for three dioecious seagrass species (Thalassia testudinum, Syringodium filiforme and Halodule wrightii) in the Mexican Caribbean. Seagrass flowers have a highly reduced or absent corolla and florivores directly consumed the anthers with pollen. The foliar structures (tepals, bracts or sheaths) protecting the male flower buds were removed by hand in situ. The floral buds were followed by videos or taking pictures at regular intervals and most (56–100 %, depending on seagrass species and experimental setting) artificially denuded male flower buds were consumed within 24 h by juvenile fish of various species. Histochemical analysis showed that the pollen and embedding mucilage were rich in polysaccharides and proteins, thus potentially nutritious. The seagrasses had copious production of pollen (between 0.2 and 1.2 × 10⁶ pollen per flower, depending on the species). But T. testudinum and S. filiforme were often pollen limited, and the probability of fruit set was reduced ~50 % when the females were at the distance of 1 and 5–6 m from the males flowers, respectively. Under natural conditions, depredation on pre-anthesis male flowers in the three species was low because flower bud emergence (few hours) and pollen release (1–4 h) were ephemeral processes. In addition, the release of pollen of T. testudinum occurred at dusk when herbivorous fish became inactive. These life-cycle characteristics aid to avoid excessive pollen consumption by fish, however, whether they are anti-predator strategies or mere adaptations for submarine pollination remains to be established.     

白背叶(大戟科)性系统和传粉生物学研究

为探讨野桐属(Mallotus)雌雄异株的进化和传粉机制,对白背叶(Mallotus apelta)野生居群的性系统和传粉生物学进行了研究。结果表明,所调查的白背叶居群均由雌性(F)、雄性(M)和少数两性(B)个体组成,平均性比为1：0.66：0.18(F：M：B);3种性别植株的基径大小差异不显著;雌株与两性植株的单花胚珠数、单果结籽数和自然结实率无显著差异;雄株与两性植株的花粉萌发率、花粉组织化学和花粉微形态特征也无显著差异,但雄株的单花花粉量是两性植株的1.26~1.63倍,且差异显著;雌株的异交结实率为96.67%,两性植株的异交结实率为76.00%,两者差异显著,说明居群内雌株的潜在种子生产力明显高于两性植株;野外观察到雄株和两性植株上的雄花具有访花者而雌株没有;雌株经套网处理后结实率超过30%而套袋处理不结实。这些表明白背叶具典型的亚雌雄异株性系统,雌株和雄株的适合度均高于两性植株;雌株以风媒传粉结实,两性植株可能兼有风媒和虫媒传粉特征。     

A cascade signal pathway occurs in self-incompatibility of Pyrus pyrifolia

Pear (Pyrus pyrifolia L.) possesses an S-RNase-based gametophytic self-incompatibility (GSI) system and S-RNase, the self-incompatibility (SI) determinant in the pistil, has also been implicated in the rejection of self-pollen and genetically identical pollen. We have demonstrated that S-RNase depolymerises actin cytoskeleton, triggers mitochondrial alteration and DNA degradation in the incompatible pollen tube, which indicates programmed cell death (PCD) may occur in SI response of Pyrus pyrifolia. Recently, we have identified that S-RNase specifically disrupted tip-localized reactive oxygen species (ROS) of incompatible pollen tube via arrest of ROS formation in mitochondria and cell walls in Pyrus pyrifolia. Furthermore, tip-localized ROS disruption not only decreased the Ca²⁺ current and depolymerised the actin cytoskeleton, but it also induced nuclear DNA degradation in the pollen tube. The results mentioned above indicate that a cascade signal pathway may occur in SI of Pyrus pyrifolia and PCD is used to terminate the incompatible pollen tubes growth. In this addendum, we review the cascade signal pathway of Pyrus pyrifolia SI.Key words: S-RNase, programmed cell death, reactive oxygen species, actin cytoskeleton, Ca²⁺ current, nuclear DNA     

Active pollination favours sexual dimorphism in floral scent

Zoophilous flowers often transmit olfactory signals to attract pollinators. In plants with unisexual flowers, such signals are usually similar between the sexes because attraction of the same animal to both male and female flowers is essential for conspecific pollen transfer. Here, we present a remarkable example of sexual dimorphism in floral signal observed in reproductively highly specialized clades of the tribe Phyllantheae (Phyllanthaceae). These plants are pollinated by species-specific, seed-parasitic Epicephala moths (Gracillariidae) that actively collect pollen from male flowers and pollinate the female flowers in which they oviposit; by doing so, they ensure seeds for their offspring. We found that Epicephala-pollinated Phyllanthaceae plants consistently exhibit major qualitative differences in scent between male and female flowers, often involving compounds derived from different biosynthetic pathways. In a choice test, mated female Epicephala moths preferred the scent of male flowers over that of female flowers, suggesting that male floral scent elicits pollen-collecting behaviour. Epicephala pollination evolved multiple times in Phyllantheae, at least thrice accompanied by transition from sexual monomorphism to dimorphism in floral scent. This is the first example in which sexually dimorphic floral scent has evolved to signal an alternative reward provided by each sex, provoking the pollinator''s legitimate altruistic behaviour.     

Do specialized flowers promote reproductive isolation? Realized pollination accuracy of three sympatric Pedicularis species

Background and Aims

Interest in pollinator-mediated evolutionary divergence of flower phenotype and speciation in plants has been at the core of plant evolutionary studies since Darwin. Specialized pollination is predicted to lead to reproductive isolation and promote speciation among sympatric species by promoting partitioning of (1) the species of pollinators used, (2) when pollinators are used, or (3) the sites of pollen placement. Here this last mechanism is investigated by observing the pollination accuracy of sympatric Pedicularis species (Orobanchacae).

Methods

Pollinator behaviour was observed on three species of Pedicularis (P. densispica, P. tricolor and P. dichotoma) in the Hengduan Mountains, south-west China. Using fluorescent powder and dyed pollen, the accuracy was assessed of stigma contact with, and pollen deposition on, pollinating bumble-bees, respectively.

Key Results

All three species of Pedicularis were pollinated by bumble-bees. It was found that the adaptive accuracy of female function was much higher than that of male function in all three flower species. Although peak pollen deposition corresponded to the optimal location on the pollinator (i.e. the site of stigma contact) for each species, substantial amounts of pollen were scattered over much of the bees'' bodies.

Conclusions

The Pedicularis species studied in the eastern Himalayan region did not conform with Grant''s ‘Pedicularis Model’ of mechanical reproductive isolation. The specialized flowers of this diverse group of plants seem unlikely to have increased the potential for reproductive isolation or influenced rates of speciation. It is suggested instead that the extreme species richness of the Pedicularis clade was generated in other ways and that specialized flowers and substantial pollination accuracy evolved as a response to selection generated by the diversity of co-occurring congeners.     

A positive feedback loop mediated by CsERF31 initiates female cucumber flower development

Sex determination is a crucially important developmental event that is pervasive throughout nature and enhances the adaptation of species. Among plants, cucumber (Cucumis sativus L.) can generate both unisexual and bisexual flowers, and the sex type is mainly controlled by several 1-aminocyclopropane-1-carboxylic acid synthases (CsACSs). However, the regulatory mechanism of these synthases remains elusive. Here, we used gene expression analysis, protein–DNA interaction assays, and transgenic plants to study the function of a gynoecium-specific gene, ETHYLENE RESPONSE FACTOR31 (CsERF31), in female flower differentiation. We found that in a predetermined female flower, ethylene signaling activates CsERF31 by CsEIN3, and then CsERF31 stimulates CsACS2, which triggers a positive feedback loop to ensure female rather than bisexual flower development. A similar interplay is functionally conserved in melon (Cucumis melo L.). Knockdown of CsERF31 by RNAi causes defective bisexual flowers to replace female flowers. Ectopic expression of CsERF31 suppresses stamen development and promotes pistil development in male flowers, demonstrating that CsERF31 functions as a sex switch. Taken together, our data confirm that CsERF31 represents the molecular link between female–male determination and female–bisexual determination, and provide mechanistic insight into how ethylene promotes female flowers, rather than bisexual flowers, in cucumber sex determination.

A key regulator promotes female flower development by triggering a positive feedback loop during cucumber sex determination.     

The endangered Iris atropurpurea (Iridaceae) in Israel: honey-bees,night-sheltering male bees and female solitary bees as pollinators

Background and Aims

The coastal plain of Israel hosts the last few remaining populations of the endemic Iris atropurpurea (Iridaceae), a Red List species of high conservation priority. The flowers offer no nectar reward. Here the role of night-sheltering male solitary bees, honey-bees and female solitary bees as pollinators of I. atropurpurea is documented.

Methods

Breeding system, floral longevity, stigma receptivity, visitation rates, pollen loads, pollen deposition and removal and fruit- and seed-set were investigated.

Key Results

The main wild pollinators of this plant are male eucerine bees, and to a lesser extent, but with the potential to transfer pollen, female solitary bees. Honey-bees were found to be frequent diurnal visitors; they removed large quantities of pollen and were as effective as male sheltering bees at pollinating this species. The low density of pollen carried by male solitary bees was attributed to grooming activities, pollen displacement when bees aggregated together in flowers and pollen depletion by honey-bees. In the population free of honey-bee hives, male bees carried significantly more pollen grains on their bodies. Results from pollen analysis and pollen deposited on stigmas suggest that inadequate pollination may be an important factor limiting fruit-set. In the presence of honey-bees, eucerine bees were low removal–low deposition pollinators, whereas honey-bees were high removal–low deposition pollinators, because they removed large amounts into corbiculae and deposited relatively little onto receptive stigmas.

Conclusions

Even though overall, both bee taxa were equally effective pollinators, we suggest that honey-bees have the potential to reduce the amount of pollen available for plant reproduction, and to reduce the amount of resources available to solitary bee communities. The results of this study have potential implications for the conservation of this highly endangered plant species if hives are permitted inside reserves, where the bulk of Oncocyclus iris species are protected.     

The need to re-investigate the nature of homoplastic characters: an ontogenetic case study of the 'bracteoles' in Atripliceae (Chenopodiaceae)

Background and Aims

Within Chenopodioideae, Atripliceae have been distinguished by two bracteoles enveloping the female flowers/fruits, whereas in other tribes flowers are described as ebracteolate with persistent perianth. Molecular phylogenetic hypotheses suggest ‘bracteoles’ to be homoplastic. The origin of the bracteoles was explained by successive inflorescence reductions. Flower reduction was used to explain sex determination. Therefore, floral ontogeny was studied to evaluate the nature of the bracteoles and sex determination in Atripliceae.

Methods

Inflorescences of species of Atriplex, Chenopodium, Dysphania and Spinacia oleracea were investigated using light microscopy and scanning electron microscopy.

Key Results

The main axis of the inflorescence is indeterminate with elementary dichasia as lateral units. Flowers develop centripetally, with first the formation of a perianth primordium either from a ring primordium or from five individual tepal primordia fusing post-genitally. Subsequently, five stamen primordia originate, followed by the formation of an annular ovary primordium surrounding a central single ovule. Flowers are either initially hermaphroditic remaining bisexual and/or becoming functionally unisexual at later stages, or initially unisexual. In the studied species of Atriplex, female flowers are strictly female, except in A. hortensis. In Spinacia, female and male flowers are unisexual at all developmental stages. Female flowers of Atriplex and Spinacia are protected by two accrescent fused tepal lobes, whereas the other perianth members are absent.

Conclusions

In Atriplex and Spinacia modified structures around female flowers are not bracteoles, but two opposite accrescent tepal lobes, parts of a perianth persistent on the fruit. Flowers can achieve sexuality through many different combinations; they are initially hermaphroditic, subsequently developing into bisexual or functionally unisexual flowers, with the exception of Spinacia and strictly female flowers in Atriplex, which are unisexual from the earliest developmental stages. There may be a relationship between the formation of an annular perianth primordium and flexibility in floral sex determination.     

Pre-zygotic embryological characters ofPlatycrater arguta, a rare and endangered species endemic to East Asia

Platycrater arguta Sieb. et Zucc. is a rare and endangered species endemic to East Asia. It produces two floral morphs viz. bisexual and male flowers. For bisexual flowers, simultaneous cytokinesis in the microsporocyte meiosis leads to a tetrahedral tetrad. The mature pollen grain is shed at 2-cell stage. The young anther wall is composed of epidermis, endothecium that develops fibrous thickenings at maturity, 1–2 middle layers and tapetum. The tapetum with uninucleate to binucleate cells, disintegrates in situ (glandular tapetum), yet in a small percentage of the anthers (about 37.6%), the tapetum does not disintegrate, causing complete male sterility. The ovules are anatropous, unitegmic, tenuinucellar and the formation of the embryo sac follows the monosporic, Polygonum type. Antipodal cells are lacking in the mature embryo sacs. Before fertilization, two polar nuclei fuse into a secondary nucleus. The formation of microsporangial wall, microsporogenesis and male gametogenesis in male flowers are analogous to those in the bisexual. Prezygotic embryological characters ofP. arguta were reported for the first time, revealing that its endangerment is correlated with the abortion of pollen of a part but not to the female development that is normal.     

Evidence of trimonoecy in Phyllanthaceae: Phyllanthus acidus

Pollen from staminate flowers and pistillate flowers with ??staminodes?? of Phyllanthus acidus Skeels were analyzed under scanning electron microscopy, and tests of pollen viability and in?vitro germination were carried out to verify possible similarities between the three types of flowers. The results show that pistillate flowers with ??staminodes?? are bisexual, indicating the occurrence of trimonoecy in this species.     

Size-dependent gender modification in Lilium apertum (Liliaceae): does this species exhibit gender diphasy?

Background and Aims

Variation in the relative female and male reproductive success of flowering plants is widespread, despite the fundamental hermaphroditic condition of the majority of species. In many hermaphroditic populations, environmental conditions and their influence on development and size can influence the gender expression of individuals through the formation of hermaphroditic and unisexual flowers. This study investigates the hypothesis that the bulbous, animal-pollinated, perennial Lilium apertum (Liliaceae) exhibits a form of size-dependent gender modification known as gender diphasy, in which the sexual expression of individuals depends on their size, with plants often changing sex between seasons.

Methods

Variation in floral traits was examined in relation to their size using marked individuals in natural populations, and also under glasshouse conditions. Measurements were taken of the height, flower number, floral sex expression, flower size, flower biomass and pollen production of individuals over consecutive years between 2009 and 2012 in seven populations in south-west China.

Key Results

Flowers of L. apertum are either perfect (hermaphroditic) or staminate (male) and, in any given season, plants exhibit one of three sex phenotypes: only hermaphrodite flowers, a mixture of hermaphroditic and male flowers, or only male flowers. Transitions between each of these sex phenotypes were observed over consecutive years and were commonly size-dependent, particularly transitions from small plants bearing only male flowers to those that were taller with hermaphroditic flowers. Hermaphroditic flowers were significantly larger, heavier and produced more pollen than male flowers.

Conclusions

The results for L. apertum are consistent with the ‘size advantage hypothesis’ developed for animal species with sex change. The theory predicts that when individuals are small they should exhibit the sex for which the costs of reproduction are less, and this usually involves the male phase. L. apertum provides an example of gender diphasy, a rare sexual system in flowering plants.     

Asymmetrical conspecific seed-siring advantage between Silene latifolia and S. dioica

Background and Aims

Silene dioica and S. latifolia experience only limited introgression despite overlapping flowering phenologies, geographical distributions, and some pollinator sharing. Conspecific pollen precedence and other reproductive barriers operating between pollination and seed germination may limit hybridization. This study investigates whether barriers at this stage contribute to reproductive isolation between these species and, if so, which mechanisms are responsible.

Methods

Pollen-tube lengths for pollen of both species in styles of both species were compared. Additionally, both species were pollinated with majority S. latifolia and majority S. dioica pollen mixes; then seed set, seed germination rates and hybridity of the resulting seedlings were determined using species-specific molecular markers.

Key Results

The longest pollen tubes were significantly longer for conspecific than heterospecific pollen in both species, indicating conspecific pollen precedence. Seed set but not seed germination was lower for flowers pollinated with pure heterospecific versus pure conspecific pollen. Mixed-species pollinations resulted in disproportionately high representation of nonhybrid offspring for pollinations of S. latifolia but not S. dioica flowers.

Conclusions

The finding of conspecific pollen precedence for pollen-tube growth but not seed siring in S. dioica flowers may be explained by variation in pollen-tube growth rates, either at different locations in the style or between leading and trailing pollen tubes. Additionally, this study finds a barrier to hybridization operating between pollination and seed germination against S. dioica but not S. latifolia pollen. The results are consistent with the underlying cause of this barrier being attrition of S. dioica pollen tubes or reduced success of heterospecifically fertilized ovules, rather than time-variant mechanisms. Post-pollination, pre-germination barriers to hybridization thus play a partial role in limiting introgression between these species.     

Mutual reproductive dependence of distylic Cordia leucocephala (Cordiaceae) and oligolectic Ceblurgus longipalpis (Halictidae, Rophitinae) in the Caatinga

Background and Aims

The close relationship between distylic Cordia leucocephala and the bee Ceblurgus longipalpis, both endemic to the Caatinga, north-east Brazil, was investigated, emphasizing reproductive dependence, morphological adaptations of the partners, and pollen flow.

Methods

In the municipality of Pedra, in the Caatinga of Pernambuco, the breeding system and reproductive success of C. leucocephala, its interaction with flower visitors and inter- and intramorph pollen flow were determined.

Key Results

The bee Ceblurgus longipalpis, the unique flower visitor and effective pollinator of self-incompatible Cordia leucocephala, presents morphological features adapted to exploit hidden pollen and nectar in the long and narrow corolla tubes. Pollen of low-level anthers is collected with hairs on prolonged mouthparts and pollen of high-level anthers with clypeus, mandibles, and labrum, showing pollen removal from both levels with the same effectiveness. In both morphs, this results in similar legitimate, i.e. intermorph cross-pollen flow. Illegitimate pollen flow to stigmas of pin flowers, however, was much higher than to stigmas of thrum flowers. Moreover, more illegitimate pollen was transported to stigmas of pin and less to those of thrum flowers when compared with legitimate pollen flow.

Conclusions

The study reveals a one-to-one reproductive inter-dependence between both partners. Data indicate that this relationship between bee species and plant species is one of the rare cases of monolecty among bees. Monotypic Ceblurgus longipalpis, the only rophitine species of Brazil, evolved prolonged mouthparts rare among short-tongued bees that enable them to access pollen from flowers with short-level anthers hidden for bees of other species, and nectar at the base of the flower tube.     

Gender variation in Actinidia deliciosa,the kiwifruit

Summary Flower and fruit characters were measured in ten female, five male and five fruiting male selections of A. deliciosa var deliciosa (A. Chev) Liang and Ferguson. Flowers from female vines had functional pistils, which contained many ovules. Stamens appeared to be fully developed but produced only empty pollen grains. Flowers from male vines had functional stamens that produced high percentages of pollen grains with stainable cytoplasmic contents. Pistils did not contain ovules and were generally small with vestigial styles. Fruiting male vines had both staminate and bisexual flowers. Staminate flowers were similar to those found on strictly male vines. Bisexual flowers produced ovules and stainable pollen. Pistils were smaller than in pistillate flowers. Although the three flower sexes differed in style length, ovary dimensions and ovules per carpel, staminate and bisexual flowers were similar in number of flowers per inflorescence, stamen filament length, pollen stainability, inflorescence rachis length and carpel number, and differed from pistillate flowers in these characters. The three flower sexes had similar sepal and petal numbers. The fruit of fruiting males were considerably smaller than those of females. Low ovule number appears to be the major factor limiting fruit size in the fruiting males studied. Prospects for developing hermaphroditic kiwifruit cultivars through breeding are discussed.