Gender expression and inflorescence structure of Pappea capensis Eckl. and Zeyh. (Sapindaceae)

Gender and the structure of the inflorescence and flowers of Pappea capensis (Sapindaceae) are investigated in a locality around Pretoria (22-27°S and 25-32°E). The trees flower over a long period (December to April) and are basically monoecious, starting with male flowers followed by female flowers towards the end of the flowering period, although some trees may be predominantly male and some predominantly female. The inflorescence is a reduced thyrse with small flowers. Male flowers have five ephemeral petals, eight stamens and a rudimental pistil. Female flowers comprise a 3-lobed ovary, a single style and stigma and eight staminodes.     

Change of Floral Orientation within an Inflorescence Affects Pollinator Behavior and Pollination Efficiency in a Bee-Pollinated Plant,Corydalis sheareri

Vertical raceme or spike inflorescences that are bee-pollinated tend to present their flowers horizontally. Horizontal presentation of flowers is hypothesized to enhance pollinator recognition and pollination precision, and it may also ensure greater consistency of pollinator movement on inflorescences. We tested the hypotheses using bee-pollinated Corydalis sheareri which has erect inflorescences consisting of flowers with horizontal orientation. We altered the orientation of individual flowers and prepared three types of inflorescences: (i) unmanipulated inflorescences with horizontal-facing flowers, (ii) inflorescences with flowers turned upward, and (iii) inflorescences with flowers turned downward. We compared number of inflorescences approached and visited, number of successive probes within an inflorescence, the direction percentage of vertical movement on inflorescences, efficiency of pollen removal and seed production per inflorescence. Deviation from horizontal orientation decreased both approaches and visits by leafcutter bees and bumble bees to inflorescences. Changes in floral orientation increased the proportion of downward movements by leafcutter bees and decreased the consistency of pollinator movement on inflorescences. In addition, pollen removal per visit and seed production per inflorescence also declined with changes of floral orientation. In conclusion, floral orientation seems more or less optimal as regards bee behavior and pollen transfer for Corydalis sheareri. A horizontal orientation may be under selection of pollinators and co-adapt with other aspects of the inflorescence and floral traits.     

Observations of inflorescence patterns and flower sex in a population of Sagittaria papillosa Buch. (Alismataceae)

Sagittaria papillosa Buch. is monoecious with unisexual flowers, pistillate below, staminate above, typically with an unbranched scape. A large population with unusual numbers of staminate and bisexual flowers on the lowest whorl of the inflorescence and many particles was quantitatively evaluated. First-formed inflorescences had more staminate and bisexual flowers than those produced later. Branched scapes were predominantly found to be the second inflorescence produced by a given plant. Genetic crosses between flowers on recemes and panicles produced no branched inflorescences. When grown under greenhouse conditions all tested plants had racemes with pistillate flowers in the lower whorls and staminate ones above. Data from soil parameters, daylengths and air temperatures are compared to reported information on modification of flower sexuality by these factors.     

Floral and inflorescence morphology and ontogeny in Beta vulgaris, with special emphasis on the ovary position

Background and Aims

In spite of recent phylogenetic analyses for the Chenopodiaceae–Amaranthaceae complex, some morphological characters are not unambiguously interpreted, which raises homology questions. Therefore, ontogenetic investigations, emphasizing on ‘bracteoles’ in Atripliceae and flowers in Chenopodioideae, were conducted. This first paper presents original ontogenetic observations in Beta vulgaris, which was chosen as a reference species for further comparative investigation because of its unclarified phylogenetic position and its flowers with a (semi-)inferior ovary, whereas all other Chenopodiaceae–Amaranthaceae have hypogynous flowers.

Methods

Inflorescences and flowers were examined using scanning electron microscopy and light microscopy.

Key Results

Floral development starts from an inflorescence unit primordium subtended by a lateral bract. This primordium develops into a determinate axis on which two opposite lateral flowers originate, each subtended by a bracteole. On a flower primordium, first five tepal primordia appear, followed by five opposite stamen primordia. Simultaneously, a convex floral apex appears, which differentiates into an annular ovary primordium with three stigma primordia, surrounding a central, single ovule. A floral tube, which raises the outer floral whorls, envelops the ovary, resulting in a semi-inferior ovary at mature stage. Similarly, a stamen tube is formed, raising the insertion points of the stamens, and forming a staminal ring, which does not contain stomata. During floral development, the calyces of the terminal flower and of one of the lateral flowers often fuse, forming a compound fruit structure.

Conclusions

In Beta vulgaris, the inflorescence is compound, consisting of an indeterminate main axis with many elementary dichasia as inflorescence units, of which the terminal flower and one lateral flower fuse at a later stage. Floral parts develop starting from the outer whorl towards the gynoecium. Because of the formation of an epigynous hypanthium, the ovary becomes semi-inferior in the course of floral development.Key words: Beta vulgaris, Chenopodiaceae, floral ontogeny, gynoecial development, epigynous hypanthium, semi-inferior ovary, inflorescence ontogeny, LM, SEM     

What makes a fig: insights from a comparative analysis of inflorescence morphogenesis in Moraceae

Background and AimsMoraceae, the family of mulberry and fig trees, displays small homogeneous flowers but extremely diverse inflorescences ranging from simple and branched to complex and condensed. Inflorescences also vary in flower organization in the receptacle, in the degree of flower condensation and in receptacle shape. Thus, the objective of the present study was to compare the inflorescence morphogenesis of Moraceae species, to investigate whether clades with a similar pollination mode share the same patterns of inflorescence development and the developmental stages at which we observe the key changes resulting in the diversified inflorescence architecture that culminates in the Ficus syconium.MethodsInflorescences at different developmental stages were sampled from Brosimum gaudichaudii, Castilla elastica, Clarisia ilicifolia, Ficus pertusa, Maclura tinctoria and Morus nigra and processed for surface and anatomical analyses.Key ResultsThe inflorescence morphogenesis of the studied species is highly variable. The shape of the inflorescence meristem (bulging, hemispheric or elongated), the initiation order and arrangement of flowers along the receptacle and the occurrence of bracts vary between related species. This diversity originates early during inflorescence development. Brosimum gaudichaudii, C. elastica and F. pertusa have flowers enclosed or immersed within the receptacle, although inflorescences begin their development as flat and open structures, as occurs in the other three study species.ConclusionComparison of the inflorescence morphogenesis in Moraceae species allows us to infer that evolutionary ontogenetic changes driven by pollinators culminate in the enclosure of flowers inside the receptacle, as occurs in the Ficus syconium.     

Honeyeater foraging: A test of optimal foraging theory

Honeyeaters (Meliphagidae) were observed foraging for nectar from Lambertia formosa inflorescences, each of which has seven flowers. The frequency distribution of numbers of flowers probed per visit to an inflorescence was found to be bimodal, with one peak at two and the other at seven. It is hypothesized that this frequency distribution results from a rule of departure from inflorescences that maximizes the net rate of energy gain. Patterns of nectar distribution were determined for a large sample of inflorescences. In addition the extent to which the honeyeaters re-probe flowers during a visit to an inflorescence was estimated. From these data and from field measurements of the times required by the honeyeaters to perform the various foraging behaviours, computer simulations of honeyeater foraging were constructed. These simulations led in turn to optimal frequency distributions of numbers of flowers probed per inflorescence that were bimodal but had peaks at 1 and 7 instead of 2 and 7. Although the observed and predicted behaviour were consequently similar, the difference between them was nevertheless significant. This difference could have been due to the birds' transient occupancy of the study area.     

Inflorescences: concepts,function, development and evolution

Background

Inflorescences are complex structures with many functions. At anthesis they present the flowers in ways that allow for the transfer of pollen and optimization of the plant''s reproductive success. During flower and fruit development they provide nutrients to the developing flowers and fruits. At fruit maturity they support the fruits prior to dispersal, and facilitate effective fruit and seed dispersal. From a structural point of view, inflorescences have played important roles in systematic and phylogenetic studies. As functional units they facilitate reproduction, and are largely shaped by natural selection.

Scope

The papers in this Special Issue bridge the gap between structural and functional approaches to inflorescence evolution. They include a literature review of inflorescence function, an experimental study of inflorescences as essential contributors to the display of flowers, and two papers that present new methods and concepts for understanding inflorescence diversity and for dealing with terminological problems. The transient model of inflorescence development is evaluated in an ontogenetic study, and partially supported. Four papers present morphological and ontogenetic studies of inflorescence development in monophyletic groups, and two of these evaluate the usefulness of Hofmeister''s Rule and inhibitory fields to predict inflorescence structure. In the final two papers, Bayesian and Monte-Carlo methods are used to elucidate inflorescence evolution in the Panicoid grasses, and a candidate gene approach is used in an attempt to understand the evolutionary genetics of inflorescence evolution in the genus Cornus (Cornaceae). Taken as a whole, the papers in this issue provide a glimpse of contemporary approaches to the study of the structure, development, and evolution of inflorescences, and suggest fruitful new directions for research.     

Diversity of inflorescences in the Boutelouinae subtribe (Poaceae: Chloridoideae: Cynodonteae)

The Boutelouinae subtribe is comprised of one monophyletic genus, Bouteloua, with 57 species inhabiting the semi-arid regions of the New World. The inflorescences show significant structural variations, which provides an interesting system to examine their morphological evolution and identify characters and processes that may help to understand the group systematics. The structure of inflorescences was studied in 25 species of Bouteloua. All the species covered under this study have truncated polytelic inflorescences. Structural variations in the inflorescence unit among species may be accounted for by: (1) symmetry of the inflorescence unit, (2) total number of long primary branches, (3) total number of spikelets per branch, (4) number of perfect flowers per spikelet, (5) number of rudimentary flowers, and (6) reproductive system. Homogenization and truncation processes account for the diversity of mature inflorescences that exists in Bouteloua. In this work, we discuss the systematic and taxonomic value of the inflorescence in the Boutelouinae subtribe.     

Unisexual flowers as a robust synapomorphy in Cariceae (Cyperaceae)? Evidence for bisexual flowers in Schoenoxiphium

Cariceae, the largest tribe within Cyperaceae, comprises about 2000 species in five genera. Cariceae is usually considered to be distinct from other Cyperaceae by the presence of exclusively unisexual flowers and by the arrangement of the pistillate flowers in single-flowered spikelets that are enclosed by the flask-like spikelet prophyll (utricle or perigynium). The nature of several morphological features of the Cariceae inflorescence remains controversial. The staminate reproductive units, as well as earlier reported bisexual reproductive units in Schoenoxiphium have been considered to be reduced partial inflorescences, or flowers. Aims of this study are to test both interpretations, based on a floral ontogenetic investigation. Moreover, for the first time, detailed SEM micrographs are presented of the inflorescence and floral development and of bisexual flowers in Schoenoxiphium. We propose that ‘inhibition of bisexuality’ is a more robust synapomorphy in Cariceae than ‘presence of only unisexual flowers’.     

Ontogeny and structure of the acervulate partial inflorescence in Hyophorbe lagenicaulis (Arecaceae; Arecoideae)

Background and Aims

The palm tribe Chamaedoreeae displays flowers arranged in a complex partial inflorescence called an acervulus. This type of partial inflorescence has so far not been reported elsewhere in the largest palm subfamily Arecoideae, which is traditionally characterized by flowers predominantly arranged in triads of one central female and two lateral male flowers. The ontogenetic basis of the acervulus is as yet unknown and its structural diversity throughout the genera of the Chamaedoreeae poorly recorded. This study aims to provide critical information on these aspects.

Methods

Developmental series and mature inflorescences were sampled from plants cultivated in international botanical gardens and wild populations. The main techniques employed included scanning electronic microscopy and serial anatomical sectioning of resin-embedded fragments of rachillae.

Key Results

Inflorescence ontogeny in Hyophorbe lagenicaulis demonstrates that the acervulus and the inflorescence rachilla form a condensed and cymose branching system resembling a coenosome. Syndesmy results from a combined process of rapid development and adnation, without or with reduced axis elongation. Acervulus diversity in the ten taxa of the Chamaedoreeae studied is displayed at the level of their positioning within the inflorescence, their arrangement, the number of floral buds and their sexual expression.

Conclusions

The results show that a more general definition of the type of partial inflorescence observed within the large subfamily Arecoideae would correspond to a cyme rather than to a floral triad. In spite of their common cymose architecture, the floral triad and the acervulus present differences with respect to the number and arrangement of floral buds, the superficial pattern of development and sexual expression.     

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.     

Optimal foraging in hummingbirds: Rule of movement between inflorescences

The movements of hummingbirds between inflorescences of scarlet gilia (Ipomopsis aggregata) were studied. These movements exhibited the following patterns: (1) Although the hummingbirds appeared to avoid moving to the previous inflorescence, no significant correlation was found between the directions of successive inter-inflorescence movements. (2) The frequency distribution of inter-inflorescence flight distances was found to be leptokurtic. (3) The hummingbirds were more likely to move to an inflorescence the larger and/or closer it was. (4) The hummingbirds moved to inflorescences of greatest apparent size (i.e. ratio of number of flowers available to distance from present inflorescence) more often than they moved to the largest inflorescence, the closest infloresence, or the inflorescence estimated to yield the greatest rate of energy gain. (5) The frequency distribution of moves to the inflorescence having the ith greatest apparent size is well fitted by a geometric distribution. This is consistent with the hummingbrids choosing the inflorescence of greatest apparent size (excluding the previous inflorescence) from within some scanning sector. These movement patterns are consistent with the expectations of optimal foraging theory only if the hummingbirds cannot or do not determine the directions of possible inflorescences relative to the direction of arrival at the present inflorescence and if they cannot assess independently the sizes and distances of possible inflorescences.     

Factors affecting the Abscission of Reproductive Organs in Yellow Lupins (Lupinus luteus L.): I. THE EFFECT OF DIFFERENT PATTERNS OF FLOWER REMOVAL

1. The effect of various patterns of flower removal on pod settingwas investigated in Lupinus luteus L. Four-fifths, three-fifths,or two-fifths of the flowers of the main inflorescence wereremoved according to ten different patterns. 2. All flowers could produce pods but later ones were less efficientin doing so. Developing pods had an abscission-inducing effecton later flowers, which became increasingly effective towardsthe apical part of the inflorescence. More pods were retained when flowers on each consecutive whorlwere arranged in a spiral than when the same number was arrangedvertically. Pod setting was incomplete when the number of flowers per inflorescencewas reduced well below the total number of pods normally present. 3. The number of ovules in consecutive flowers gradually decreasedfrom an average of 5.7 at the base to 4.3 at the top of theinflorescence. The ratio of seeds to ovules fluctuated irregularlybetween 65 and 94 per cent, and did not indicate a general trendin embryo abortion. 4. The growth-rate of pods at the top of the inflorescence wasmuch slower than at the bottom. Vascular differentiation wasalmost absent at the top of the inflorescence when the flowerswere fertilized, and further vascular tissue was produced onlywhen flowers produced pods.     

Organographic and ontogenetic studies on the inflorescence ofLespedeza cuneata (Dum.-Cours.) G. Don (Leguminosae)

Structure of inflorescence and its variation were organographically and ontogenetically studied inLespedeza cuneata (Dum.-Cours.) G. Don. An axillary inflorescence of the species forms a compound inflorescence which is composed of three or four component inflorescences. Each component inflorescence bears four (rarely six), three, two, or one flowers. Based on the arrangement of inflorescence phyllomes, the component inflorescence with four flowers is interpreted as a pseudoraceme bearing two shortened lateral shoots (partial inflorescences) each of which has two flowers. The component inflorescence with one flower appears to be terminated by the flower and to compose the cyme. Organographic observations revealed that the terminally located flower is not truly terminal, but axillary in origin. Ontogenetic observations showed that the apices of component inflorescence and partial inflorescence exist in early developmental stages in spite of variation in the form of component inflorescence. The terminally located flower in the cyme-like inflorescence was thus demonstrated to be laterally borne on the partial inflorescence axis. The component inflorescence composing the cyme-like one inL. cuneata is a reduced form in the number of partial inflorescences and of flowers from the pseudoraceme. The cyme-like inflorescence inL. cuneata resembles the inflorescence ofKummerowia.     

Early inflorescence and floral development in Cocos nucifera L. (Arecaceae: Arecoideae)

Palms are generally characterized by a large structure with a massive crown that creates difficulties in anatomical studies. The flowering behaviour of palm species may be a useful indicator of phylogenetic relationships and therefore evolutionary events. This paper presents a detailed histological study of reproductive development in coconut (Cocos nucifera L.), from initiation up to maturation of staminate and pistillate flowers. Reproductive development in coconut consists of a sequence of individual events that span more than two years. Floral morphogenesis is the longest event, taking about one year, while sex determination is a rapid process that occurs within one month. The inflorescence consists of different ultimate floral structural components. Pistillate flowers are borne in floral triads that are flanked by two functional staminate flowers. The staminate flowers are born in floral diads towards the base of the rachilla followed by solitary flowers in the middle to top of the rachilla. Three primary phases were identified in reproductive development, namely, transition of axillary bud into inflorescence bud, formation of floral buds, and sexualisation of individual flower buds. All developmental events with respect to stage or time of occurrence were determined.     

Floral development of Urospatha: merosity and phylogeny in the Lasioideae (Araceae)

In this paper we study merosity in the genus Urospatha within the framework of a resolved phylogeny of the Araceae. We analyse how a transition from dimerous or tetramerous merosity to pentamerous or hexamerous merosity can occur developmentally in the Lasioideae. In Urospatha, initiation of floral primordia along the inflorescence is acropetal, while development of flowers is basipetal. This indicates the presence of two distinct phases in the development of the Urospatha inflorescence. The first phase corresponds to initiation of flowers and establishment of the phyllotactic pattern, and the second phase to differentiation of floral organs. Urospatha is characterized by the presence of trimerous, tetramerous, pentamerous and rarely hexamerous flowers. In all types of flowers, the stamens are closely associated and opposite to the tepals. Pentamerous flowers are formed by addition of a sector comprising a stamen and tepal. Likewise, in the case of hexamerous flowers, two sectors are added. In the Lasioideae, the increase in the number of tepals and stamens is linked with two developmental processes that have appeared independently in the subfamily: (1) addition of one or two stamen?Cpetal sectors (Anaphyllopsis and Urospatha), and (2) independent increase in the number of tepals and stamens on whorls, more or less organized and inserted in alternate position (Dracontium). Tetramerous whorls as they occur in basal Lasioideae would be homologous to two dimerous whorls from an evolutionary point of view.     

Direct and indirect selection on floral pigmentation by pollinators and seed predators in a color polymorphic South African shrub

The coexistence of different color morphs is often attributed to variable selection pressures across space, time, morph frequencies, or selection agents, but the routes by which each morph is favored are rarely identified. In this study we investigated factors that influence floral color polymorphisms on a local scale in Protea, within which approximately 40% of species are polymorphic. Previous work shows that seed predators and reproductive differences likely contribute to maintaining polymorphism in four Protea species. We explored whether selection acts directly or indirectly on floral color in two populations of Protea aurea, using path analysis of pollinator behavior, nectar production, seed predation, color, morphology, and maternal fecundity fitness components. We found that avian pollinators spent more time on white morphs, likely due to nectar differences, but that this had no apparent consequences for fecundity. Instead, the number of flowers per inflorescence underpinned many of the reproductively important differences between color morphs. White morphs had more flowers per inflorescence, which itself was positively correlated with nectar production, seed predator occurrence, and total long-term seed production. The number of seeds per plant to survive predation, in contrast, was not directly associated with color or any other floral trait. Thus, although color differences may be associated with conflicting selection pressures, the selection appears to be associated with the number of flowers per inflorescence and its unmeasured correlates, rather than with inflorescence color itself.     

Can honey bees change foraging patterns?

Abstract. 1. Foraging patterns were studied using honey bees on artificial flower patches to determine if given individuals could change behaviours under differing conditions.
2. Two types of flower patches were used; those simulating a population of flowers, dimorphic for colour, and grids simulating a single colour-dimorphic inflorescence.
3. In the simulated population of flowers bees were individually constant to colour over a range of reward volumes and flower patch sizes.
4. Each bee remained individually constant to a flower morph when visiting a population-type grid but changed to random visitation on the simulated inflorescence.
5. On the simulated inflorescence, with morphs providing unequal qualities of reward, most bees foraged on the higher molarity morph.
6. Most, but not all bees, failed to minimize uncertainty on the simulated inflorescence.
7. On the simulated inflorescence, bees failed to optimize when one morph provided a greater reward volume than did the other.
8. In the population of flowers bees flew from flower to flower, whereas, they walked on the simulated inflorescence.