NECTAR PRODUCTION,COMPOSITION, ENERGETICS AND POLLINATOR ATTRACTIVENESS IN SPRING FLOWERS OF WESTERN NEW YORK

High Performance Liquid Chromatography (HPLC) was used to determine specific sugar ratios (fructose, glucose and sucrose) in nectars of nine families of flowering plants. All nectars contained all three sugars with the exception of that of Asclepias. Asclepias nectar was nearly a pure sucrose solution. Sucrose/hexose ratio was correlated with flower morphology, with tubular flowers having more sucrose and open flowers having more hexose. Open flowers contained nectar whose concentration was more affected by relative humidity than tubular flowers. Available nectar in unbagged flowers was found in volumes of 0.1 to 5 μl (17.5 to 68.2% sugar). Total sugar available per blossom amounted to 0.07 to 3.54 mg. We note that care must be taken in converting % concentration to mg sugar. Energetic yield was not as variable as the other measures and ranged from 0.72 to 3.58 cal/μl. Total daily nectar production was measured in five families and 24-hr sugar production varied from 0.64 to 5.52 mg per flower. Insect nectar feeders frequently searched many blossoms with little or no reward, but were rewarded sufficiently at rare “lucky hit” blossoms which contained relatively large nectar rewards. Insect pollinators did not seek nectars of specific sucrose-hexose ratios, but instead took nectar where caloric reward and accessibility made it most profitable.     

Pattern of flower size variation along an altitudinal gradient differs between Impatiens textori and Impatiens noli-tangere

Studies of factors influencing spatial variation in flower size offer insights into floral evolution. We investigated altitudinal variations in five flower dimensions of two native Japanese Impatiens species (I. textori and I. noli-tangere) and their interactions with their faunal visitors. These two species have similar floral traits, including flower shape, flowering time, and pollinator species; both species are pollinated mainly by Bombus diversus. In I. textori, all measured flower dimensions were negatively correlated with altitude. In contrast, in I. noli-tangere, no measured flower dimensions correlated with altitude. Thus, the altitudinal pattern of flower size variation differed between these congeneric co-habiting herbaceous plant species. The different patterns suggest that the factors (e.g. altitudinal variations of abiotic factors) that cause variation of flower size differ between these two Impatiens species even though focal species have similar floral traits (e.g. flower shape, flowering time, and pollinator species).     

Modification of flower colour by suppressing β‐ring carotene hydroxylase genes in Oncidium

Oncidium ‘Gower Ramsey’ (Onc. GR) is a popular cut flower, but its colour is limited to bright yellow. The β‐ring carotene hydroxylase (BCH2) gene is involved in carotenoid biogenesis for pigment formation. However, the role of BCH2 in Onc. GR is poorly understood. Here, we investigated the functions of three BCH2 genes, BCH‐A2, BCH‐B2 and BCH‐C2 isolated from Onc. GR, to analyse their roles in flower colour. RT‐PCR expression profiling suggested that BCH2 was mainly expressed in flowers. The expression of BCH‐B2 remained constant while that of BCH‐A2 gradually decreased during flower development. Using Agrobacterium tumefaciens to introduce BCH2 RNA interference (RNAi), we created transgenic Oncidium plants with down‐regulated BCH expression. In the transgenic plants, flower colour changed from the bright yellow of the wild type to light and white‐yellow. BCH‐A2 and BCH‐B2 expression levels were significantly reduced in the transgenic flower lips, which make up the major portion of the Oncidium flower. Sectional magnification of the flower lip showed that the amount of pigmentation in the papillate cells of the adaxial epidermis was proportional to the intensity of yellow colouration. HPLC analyses of the carotenoid composition of the transgenic flowers suggested major reductions in neoxanthin and violaxanthin. In conclusion, BCH2 expression regulated the accumulation of yellow pigments in the Oncidium flower, and the down‐regulation of BCH‐A2 and BCH‐B2 changed the flower colour from bright yellow to light and white‐yellow.     

Large flowers tend to be short-lived in Mediterranean ecosystems: Insights from three Cistus species

Larger and longer lived flowers receive more pollinators, but may also involve increased water maintenance costs under hot, dry environments. Hence, smaller and/or short-lived flowers may buffer such costs. We surveyed floral longevity in three large-flowered Mediterranean Cistus species. We hypothesize that: (1) in Cistus, floral longevity decreases with increasing air temperature and flower size; (2) in C. ladanifer, flower size and longevity increase along an altitudinal gradient; (3) floral longevity is differentially affected by temperature rather than flower size along the gradient; (4) under similar temperature, floral longevity decreases with flower size. For each species, we evaluated the effects of flower size and air temperature on floral longevity. Specifically, floral longevity was surveyed along an altitudinal gradient in the largest flowered species Cistusladanifer. Floral longevity in Cistus species lasted < 1 d and was affected by air temperature, independently of flower size. In C. ladanifer, flower size increased along the gradient but floral longevity decreased. Still, floral longevity decreased with increasing air temperature and, to a lesser extent, with flower size. Together, our findings show a triangular relationship among air temperature, flower size and floral longevity with margins for plasticity to accommodate pollinator attraction with the costs of large-flowered Mediterranean plants.     

Spatial patterns of host exploitation in a larval parasitoid of the predatory dusky large blue Maculinea nausithous

The foraging behaviour of the parasitoid wasp Neotypus melanocephalus and factors affecting parasitism at the population level were studied. This specialised parasitoid attacks caterpillars of the butterfly Maculinea nausithous, which sequentially feed on the plant Sanguisorba officinalis and specific red Myrmica ants. Among M. nausithous populations, there is considerable variation in caterpillar densities. At low M. nausithous densities, foraging might be time consuming for N. melanocephalus. High host densities may not always be advantageous to foraging parasitoids due to the caterpillars’ frequent overexploitation of ant resources and subsequent density-dependent mortality. In order to disperse progeny, we hypothesised that N. melanocephalus should search in a non-random way at the level of the micro-habitat, i.e., single flower heads of S. officinalis. Our analysis of 32 natural populations in the Upper Rhine valley in Germany did not show a density-dependent relationship between M. nausithous caterpillars and parasitism. Furthermore, habitat parameters like patch size and density of the host's food plant did not affect the parasitism rate. Foraging N. melanocephalus females preferred to search on large flower heads. They probed host-occupied flower heads only, visiting non-host-exploited flower heads only briefly. Time spent on a flower head was independent of the number of caterpillars per flower head. This study indicates that N. melanocephalus increases its foraging efficiency by preferring large flower heads that were previously shown to contain more host caterpillars than small flower heads. Furthermore, oviposition increases the likelihood of continuing to search on a flower head, which is an adaptive strategy for parasitoids foraging for aggregated hosts. However, many host-occupied flower heads were not probed by N. melanocephalus. We discuss the possibility that temporal host refuges of M. nausithous caterpillars might contribute to heterogeneity of parasitism, and why spreading offspring might constitute a suitable strategy for a parasitoid of an ant-parasitic butterfly.     

商陆属2种植物花器官发生过程观察

该研究采用扫描电镜观察红蕊商陆(Phytolacca esculenta)和浙江商陆(Phytolacca zhejiangensis)的花器官发生过程,以明确商陆属植物花的基数,以及雄蕊和雌蕊是否具有叶性器官发生的特点,阐明商陆属植物花发生的模式。观察结果显示:(1)红蕊商陆和浙江商陆在花原基发生后,小苞片以2/5圆周相继发生,花被片的发生紧接小苞片的发生进行,花被与小苞片的发生均有顺时针和逆时针方向,且二者的发生方向始终一致。(2)花被发生结束后,雄蕊在花顶端分生组织的环状分生组织上发生,没有明显的发生顺序,近似同时发生;2轮雄蕊时内轮雄蕊先发生;外轮雄蕊有少数有时偶然与花被互生,但因外轮雄蕊数多于花被数,雄蕊与花被常不互生,也没有规律性。(3)红蕊商陆和浙江商陆的心皮都在雄蕊发生后,紧接着开始发生,且雌蕊与雄蕊(或内轮雄蕊)互生发生;心皮没有发生的先后次序,且每个心皮在基部连成一个整体形成雌蕊基部并发育成为子房。(4)红蕊商陆和浙江商陆的花基数为5,雄蕊和雌蕊的发生及数目不符合5基数的特点。研究认为,红蕊商陆和浙江商陆为5基数花,该研究结果不支持商陆属植物为3基数花的发生模式。     

Butterflies show flower colour preferences but not constancy in foraging at four plant species

1. The extent to which flower colour and other visual cues influence butterfly flower choice in the field is poorly understood, especially in comparison with choices by Hymenoptera. 2. Using a novel approach to studies of visitation behaviour by butterflies, flower colour of four Asteraceae species was phenotypically manipulated to decouple the influence of that trait from others (including morphology and nectar rewards) on visitation by Lycaena heteronea, Speyeria mormonia, Cercyonis oetus, and Phyciodes campestris. 3. Flower visits were recorded to experimental flower arrays in subalpine meadows to measure (i) spontaneous preference by butterflies for particular colours and other traits and (ii) flower constancy (longer than expected strings of visits made to flowers of the same species), a behaviour that can reduce interspecific gene flow in plants. 4. Over three field seasons, 3558 individual flower visits in 1386 foraging bouts were observed for free‐flying butterflies. All four butterfly species responded to the phenotypic manipulations of flower colour, although in different ways. Speyeria mormonia and L. heteronea also exhibited preferences based on other flower traits. Lycaena heteronea responded to combinations of traits such that the other traits it preferred depended upon the context of flower colour. 5. None of the butterfly species exhibited flower constancy in any of the arrays employed. 6. The observed preferences show that butterflies, like some other pollinators, are potentially capable of exerting selection on colour and other floral traits. Moreover, these flower preferences can depend on the context of other flower traits. The absence of constancy contrasts with reports of high constancy in many bees.     

Identification and Gene Mapping of a Novel Mutant supernumerary Iodicules (snl) in Rice

In order to gain a better understanding of rice flower development, a rice flower mutant supernumerary lodicules (snl), which was identified from ethyl methane sulfonate (EMS)-treated Jinhui10 (Oryza sativa L. ssp. indica) was used in the present study. In the snl mutant, the palea obtained lemma identity, additional glume-like organs formed, lodicules increased and elongated, stamens decreased, and a few aberrant carpels formed. These phenotypes suggest that SNL is involved in the entire rice flower development. SNL was mapped between two simple sequence repeat markers RM3512 and RM1342 on chromosome 2, an approximate 800 kb region, and it co-segregated with SSR215. We conclude that SNL is a novel gene involved in flower development in rice. The present study will be useful for further cloning of the SNL gene, which will contribute to the elucidation of rice flower development.     

Geographic variation in oviposition preference for male and female host plants in a geometrid moth: implications for evolution of host choice

Several dioecious plant species exhibit sexual dimorphisms in defensive traits. However, the effects of sexual dimorphism on defense against herbivores remain poorly understood. Eurya japonica (Thunb.) (Theaceae) is a dioecious shrub that shows sexual dimorphism in the chemical defense of flower buds. Female calyces contain higher concentrations of total phenolics and condensed tannins than do male calyces. Male flower buds are edible for a florivore moth, Chloroclystis excisa (Butler) (Lepidoptera: Geometridae), whereas the female flower buds are lethal to the moth larvae. The moths prefer to oviposit on male over female E. japonica flower buds. As the moths also occur in areas lacking E. japonica, we tested whether the oviposition preference for E. japonica flower sex differed between moths sympatric and allopatric with E. japonica. The moths sympatric with E. japonica showed a stronger preference for male E. japonica than the moths allopatric with E. japonica. Our phylogeographic study using mitochondrial cytochrome oxidase subunit I gene sequences revealed little genetic differentiation between moth populations sympatric and allopatric with E. japonica. These results suggest that the adaptive oviposition preference for flower sex of E. japonica has evolved rapidly in C. excisa.     

Effects of flower number on estimated pollen transfer in natural populations of three hermaphroditic species: an experiment with fluorescent dye

The relationship between the number of open flowers and pollen transfer to stigmas was estimated in three insect-pollinated species (Cynoglossum officinale, Echium vulgare and Oenothera erythrocepala) in 1995 and 1996, using fluorescent dye as a pollen analogue. Dye was applied to an equal number of flowers on a small, a medium and a large individual and its pattern of dispersal to stigmas of conspecifics in local populations was observed. In Cynoglossum officinale and in Echium vulgare large individuals tended to receive more visits per flower. In Oenothera erythrocepala no difference in number of visits per flower was found between small and large individuals. Geitonogamous pollination clearly occurred in all species. In Cynoglossum officinale large individuals transferred more pollen per flower within the same plant (geitonogamy) and an equal or a slightly smaller amount between plants, compared to small individuals. As a result total pollen transfer per flower increased in both years with increasing flower number. In Echium vulgare and in Oenothera erythrocepala large individuals transferred more pollen per flower within the plant (pollen discounting), but a significantly lower amount per flower to other individuals in the populations compared to small individuals. For both these species, there is no difference in total pollen transfer between small and large individuals. In general, the male fitness curve, as estimated by pollen transfer, is a decelerating function of flower number. We discuss this in relation to optimal sex-allocation and the pollen donation hypothesis.     

Honeybees and the nectar of Echium plantagineum L. in southeastern Australia

An account is given of the flower of Echium plantagineum in south-eastern Australia, including stages and timing of flowering, behaviour of raindrops in the flower and aspects of floral microclimate. The concentration of nectar solutes varied with time and site, with means varying from 2 to 62% (as g sucrose/100 g solution). There was a significant negative correlation between nectar solute concentration and ambient relative humidity: the drier the air, the more concentrated the nectar. Rates of nectar secretion per flower varied with the bagging method, with long-term bagging reducing net secretion rates, possibly because of re-absorption. Rates varied with time, day and site, with a temporal pattern of change suggesting a link between rates of photosynthesis and secretion. Maximum nectar secretion rates in short-term bagging experiments were ca. 300 μg sugar/flower/hr (equivalent to > 2 mglflower/24 hr). Secretion rate was correlated with flower density. As flower density increased, secretion rate per flower decreased; rate of sugar production per unit area increased relatively more slowly than flower density. E. plantagineum could produce > 500 mg sugar/m²/day. Honeybees foraged on E. plantagineum only at ambient air temperatures above ca. 17°C unless irradiance exceeded ca. 750 W m^-2. Foragers collected nectar or pollen alone, or both, with the type of visit significantly correlated with nectar solute concentration. Below 35% (as g sucrose/100 g solution) most bees took pollen only; above 40%, most took nectar. Mean standing crop of nectar was generally < 100 μg/flower when most bees were taking nectar, but could exceed 1000 μg/flower when bees were absent or foraging mainly for pollen. Honeybees did not always remove all nectar from flowers they probed. Reabsorption of residual nectar may augment the following day's secretion.     

Flowering patterns of long-lived Heliconia inflorescences: implications for visiting and resident nectarivores

Summary Flowering patterns of four Heliconia (Heliconiaceae) species in Trinidad, West Indies were examined for their predictability and availability to the nectarivores that rely on Heliconia floral nectar. Principal flower visitors are trapling hermit hummingbirds; inflorescences are inhabited by nectarivorous hummingbird flower mites that move between inflorescences by riding in the hummingbirds' nares. Heliconia inflorescences flower for 40–200 days, providing long-term sources of copious nectar (30–60 l per flower), but each Heliconia flower lasts only a single day. As an inflorescence ages the interval increases between open flowers within a bract; wet-season inflorescences produce open flowers more slowly than dry-season conspecifics.Estimated daily energy expenditures for hermit hummingbirds demonstrate that slow production of short-lived open flowers plus low inflorescence density preclude territorial defense of Heliconia by the hermits. Heliconia flowering patterns are viewed as a means of (i) regulating reproductive investment by the plants through staggered flower production over long periods of time, and (ii) maintaining outcrossing by necessitating a traplining visitation pattern by its hummingbird pollinators. I suggest that Heliconia exhibit a two-tiered pollination system by using hermit hummingbirds primarily for outcrossing and using hummingbird flower mites primarily for self-pollination.     

Basic principles of terminal flower formation

Studies of inflorescences of the mutants bractea and terminal flower1 and double mutant bra tfl1 of Arabidopsis thaliana (L.) Heynh. have shown that the presence of a developed leaf in the node preceding the terminal flower is a necessary condition for the formation of the terminal flower perianth. This means that perianth cannot develop in an abracteose inflorescence of terminal flower. The second necessary condition for the terminal flower formation is a sufficient level of expression of the genes responsible for floral morphogenesis. Combination of these two conditions suffices for the development of a terminal flower with perianth. Since the general principles of organization are common for the majority of Angiosperms, it can be stated that if the abracteose inflorescence is terminated by a flower with perianth, this is a consequence of displacement of the lateral flower into the terminal position.__________Translated from Ontogenez, Vol. 36, No. 2, 2005, pp. 90–95.Original Russian Text Copyright © 2005 by Penin, Choob, Ezhova.     

In vitro flower induction in roses

Summary Vegetatively propagated plantlets of six rose cultivars were induced to flower in vitro on media containing full-strength Murashige and Skoog (MS) inorganic salts, Gamborg's B5 organic elements with 400 mg l⁻¹ myo-inositol, and different phytohormone combinations of 6-benzyladenine (BA) with α-naphthaleneacetic acid (NAA); thidiazuron (TDZ) with NAA; and zeatin (ZT) with NAA. The most efficient flower bud induction (49.1% and 44.1%) was obtained on media supplemented with 0.5 mg l⁻¹ (2.27 μM) TDZ and 0.1 mg l⁻¹ (0.54 μM) NAA or 0.5 mg l⁻¹ (2.28 μM) ZT and 0.1 mg l⁻¹ (0.54 μM) NAA for cultivar Orange Parade. Scanning electron microscopy (SEM) showed that in vitro flower bud induction occurred mostly between 15 and 30 d in induction medium through the normal flower development processes. With TDZ and ZT as the best choice for flower induction in all six cultivars tested, different rose cultivars varied in their responses to phytohormone treatments. Our study also revealed that the total time from original culture and subculture time before flower induction were two very important factors for in vitro flower induction. Plantlets 156–561 d from original culture and subcultured for 45 d were the best for flower induction. These authors contributed equally to this work.     

Symplocos parvibracteata sp. nov. (Symplocaceae) from the Western Ghats,Kerala, India

Symplocos parvibracteata sp. nov. (Symplocaceae) from Chemunji Hills, south Western Ghats, Kerala State, India, is described and illustrated. It is most similar to S. wynadensis and S. tenella, a putative synonym of S. wynadensis, but differs in having shorter racemes, shorter bracts and bracteoles that are persistent in flower and caducous in fruit, narrowly ellipsoid flower buds, and fewer stamens per flower. It is also similar to S. sumuntia, from which it differs by its pubescent branchlets, generally more stamens per flower, and a pubescent floral disk, and to S. macrophylla, from which it differs by the combination of 3 to 5 leaf secondary veins on each side of the midvein, 1–2‐cm long inflorescences, and bracts that are persistent in flower, caducous in fruit, and < 1.5 mm long.     

Bottom‐up effects on a plant‐endophage‐parasitoid system: The role of flower‐head size and chemistry

The effects of water and nutrient addition on a trophic chain were studied in a plant‐endophage‐parasitoid system comprised of insects associated with flower heads of Chromolaena squalida (Asteraceae). Nine species of endophages associated with C. squalida flower heads were found, belonging to two families of Diptera – Tephritidae (Cecidochares sp1, Cecidochares sp2, Xanthaciura biocellata, X. chrysura, X. sp. and Neomyopites sp.) and Agromyzidae (Melanagromyza sp.), and two families of Coleoptera – Apionidae (Apion sp.) and Anthicidae (Anthicidae sp.). A factorial field experiment with water and nutrient addition showed that resource availability can affect the developmental process of flower heads. Fertilization increased flower‐head diameter and nitrogen and alkaloid concentrations. Although nutrient availability affected the size and chemistry of flower heads, endophage species did not respond consistently to the experimental treatments. This is contrary to other studies where endophages showed preference for larger flower heads. Thus, the plant vigour hypothesis was not corroborated for our study system. Our results also showed that coupled responses of plants to resource availability (i.e. tissue nutritional quality and investment in growth of the structure that serves as shelter for endophages) can represent distinct kinds of indirect interactions with opposing effects on the herbivore‐parasitoid interaction.     

VARIATION AND COVARIATION AMONG FLORAL TRAITS WITHIN AND AMONG FOUR SPECIES OF NORTHERN EUROPEAN PRIMULA (PRIMULACEAE)

Phenotypic and genetic variation and correlations among floral traits within and among four Primula species were measured to seek evidence for potential constraints on the independent evolution of floral characters, to examine the relationship between mating system, ploidy level, and sex allocation, and to determine whether some traits are more conservative than others within and across these congeners. We measured mean flower diameter, corolla depth, pollen production, modal pollen grain volume, ovule number per flower, and pollen: ovule ratios for 64 field-collected genotypes from northern Europe. These represented one heterostylous (P. farinosa: 2n = 18) and three homostylous (P. scotica: 2n = 54, P. scahdinavica: 2n = 74, and P. stricta: 2n ~ 126) species. All traits differed significantly among species and among the six taxon/morph categories identified (including three morphs of P. farinosa: pin, thrum, and homostylous). Pollen production per flower was significantly higher (and individual pollen grain volume lower) in the outcrossing P. farinosa than in any of the homostylous species; also, pin morphs produced significantly more pollen per flower than thrums in P. farinosa. Among the homostylous species, there were significant differences in all traits except modal pollen grain volume. Ovule number per flower and flower size were less variable among taxa than pollen production and pollen volume. Within species, there were several strong negative correlations among genets between pairs of traits, but each species exhibited a unique set of inverse relationships. We detected only one significant positive genetic correlation; in P. stricta, ovule number and pollen production per flower were positively correlated among genets. Among species means, two pairs of traits were negatively correlated: mean ovule number per flower vs. flower diameter (but P = 0.0587), and mean pollen production per flower vs. modal pollen grain volume. These negative correlations within and among taxa suggest that there may be intrinsic genetic constraints on the independent evolution of these floral characters, but that these constraints differ among species.     

Flower morphology evolution in Jaborosa (Solanaceae): shape and size variation associated with contrasting pollination modes

• Flower morphology is considered an important factor in species diversification because it may influence the efficiency of pollination in different ways (e.g. attraction and mechanical fit with different groups of pollinators). In the present study, we quantified the variation in flower morphology (i.e. shape and size) of the diverse South American genus Jaborosa Juss. (Solanaceae) in relation to contrasting pollination modes: rewarding pollination either by moths or by generalist small insects versus brood-site deceptive pollination by saprophilous flies.
• We examined variations of flower morphology in frontal (pollinator attraction) and sagittal (functional fit with pollinators) views in 12 Jaborosa species using geometric morphometric methods and comparative approaches to infer whether flower shape evolution, not attributable to flower allometry or phylogenetic relationship, is associated with shifts in pollination modes.
• We found remarkable variation in flower morphology among both Jaborosa species and pollination modes, largely in sagittal view. Evolutionary trends in shape of fly-pollinated flowers were mainly attributable to changes in developmental trajectories. Variation in flower architecture facilitated differential pollen placement – on the proboscis of moths, and either on the back or ventral region of saprophilous flies – promoting diversification of the genus.
• Diversification of shape, independent of size, in most of the studied Jaborosa species would indicate adaptation to contrasting pollination modes.