A nodding capitulum enhances the reproductive success of Cremanthodium campanulatum (Asteraceae) at high elevations in the Sino–Himalayan Mountains

Background: Flower orientation is considered an evolutionary response to pollinators and abiotic factors. Several members of the genus Cremanthodium (Asteraceae) growing in alpine habitats in the Sino–Himalayas have conspicuously nodding capitula, the function of which is not understood.

Aims: We investigated the influences of nodding capitula on floral thermal conditions, pollinator visitation rate, pollen viability and achene production in Cremanthodium campanulatum.

Results: (1) the nodding capitulum did not modify internal thermal conditions, but the elongated involucral bracts could shelter pollen from UV-B radiation; (2) water and UV-B radiation seriously reduced the viability of pollen grains; (3) capitula artificially held erect set significantly fewer achenes than nodding ones; and (4) no pollinator preference was observed between capitula that were artificially held erect and natural nodding ones.

Conclusions: This study indicated that the nodding capitulum of C. campanulatum may protect pollen from being damaged by UV-B radiation, being washed away from the anther and stigma, and enable it to avoid rain damage during the wet monsoonal period. Our results suggest that nodding capitula in C. campanulatum possibly confer a selective advantage and may be driven by non-biological agents rather than pollinator attractors.     

Foraging by honeybees on Carduus acanthoides: pattern and efficiency

Abstract.

• 1 The relationship between the fluctuation in total quantity of sugar available in the nectar of Carduus acanthoides inflorescences and the foraging behaviour of honeybees Apis mellifera ligustica was studied in a patch under natural conditions.
• 2 Every day, three inflorescence populations coexisted, one for each day of the inflorescence life-span. The maximum availability of sugar occurred in second-day capitula from 12.00 to 15.00 hours.
• 3 Bee arrivals at the patch were positively correlated with the total availability of sugar as determined by sampling.
• 4 Bees responded to variation in the availability of sugar by visiting the inflorescence stage that was most rewarding at that time of day. Early in the morning, when the frequency of visits was low, the bees worked the third-day capitula, which were the best available choice at that time. Similarly, beginning at 12.00 hours, bees switched to second-day capitula. At the end of the day, some bees switched to third-day capitula where a slightly superior quantity of sugar could be obtained from a small number of florets although other bees continued to work the second-day capitula.
• 5 Individual bees switched to another inflorescence stage after a high proportion of their visits to a small number of capitula of one kind had provided a low reward.
• 6 The number of florets visited per minute per bee varied throughout the day and was positively correlated with the availability of sugar.

     

Pubescence,floral temperature and fecundity in species of Puya (Bromeliaceae) in the Ecuadorian Andes

Summary The effects of inflorescence pubescence on floral temperatures and fecundity were examined in four species of Puya in the Ecuadorian Andes. These species span an elevational range from 1,980–4,000 m and comparisons among these species showed that pubescence production increases significantly with elevation. Flower temperatures of glabrous or slightly pubescent species of Puya from lower elevations closely tracked air temperature, while those of the pubescent páramo species did not. Pubescence removal experiments on Puya hamata, a páramo species, demonstrated that pubescence is an effective insulator, maintaining elevated flower temperatures. In Puya clava-herculis (also a páramo species) elevated flower temperatures were associated with higher seed set. Thus, the greater production of inflorescence pubescence in páramo species of Puya may be an important factor contributing to reproductive success in these higher elevation species.     

Photonastic and thermonastic opening of capitulum in dandelion,Taraxacum officinale andTaraxacum japonicum

The capitula ofTaraxacum officinale andT. japonicum open in response to temperature rise at lower temperatures (thermonasty), and in response to light at higher temperatures (photonasty), as was the case inT. albidum. The capitula ofT. officinale could respond to the same temperature rise more sensitively than those ofT. albidum orT. japonicum. The minimum temperature for photonastic opening is as low as 13 C forT. officinale, while that forT. albidum andT. japonicum is about 18 C. That is why the capitula ofT. officinale opened earlier than those ofT. albidum andT. japonicum in the morning in April under natural conditions. The capitulum continued to be open for about 13–14 hr inT. officinale and about 8–11 hr inT. japonicum and inT. albidum both under natural conditions in April and even under constant light-temperature conditions, suggesting that the time of capitula-closing in these three species is not controlled by changes in environmental factors (light and/or temperature).     

The movement of involucral bracts of Syngonanthus elegans (Eriocaulaceae-Poales): Anatomical and ecological aspects

Syngonanthus elegans flowers are distributed in capitula whose involucral bracts open and close in a diurnal rhythm. The anatomy of these bracts was studied to understand how such movements occur and how it influences reproductive ecology of the species. The involucral bracts have a single layered epidermis composed of thick-walled cells on the abaxial surface, which are responsible for the movement. Since they are hygroscopic, these cells swell when they absorb water from the surrounding environment, causing the bracts to bend and the capitula to close. In natural conditions, the capitula open by day, when temperature increases and the relative air humidity decreases, and close at night, when temperature decreases and the relative air humidity increases. The involucral bracts may thus protect the flowers from abiotic factors, exposing them only at the time of the day when temperature is higher and insects are more active, favoring pollination by small insects. The closed capitula do not only protect the flowers, but they also function as a shelter for floral visitors as Brachiacantha australe (Coccinellidae) and Eumolpini sp. (Chrysomelidae). These small Coleoptera pollinate the flowers of S. elegans during the day and remain within the closed capitula during the night, in a possible mutualistic relationship.     

EXPERIMENTAL ECOLOGICAL GENETICS IN PLANTAGO IV. EFFECTS OF TEMPERATURE ON GROWTH RATES AND REPRODUCTION IN THREE POPULATIONS OF PLANTAGO LANCEOLATA L. (PLANTAGINACEAE)

The effects of temperature on growth rates and flowering responses of Plantago lanceolata L. populations sampled from open, shaded, and sunflecked habitats were studied in controlled environments. A large number of characteristics were phenotypically plastic in response to a broad range of temperatures. Low night temperatures substantially reduced reproductive effort, whereas high temperatures produced a reduction in total biomass. Temperature had little effect on leaf or inflorescence production rates, other than at very low temperatures. The phenotypic plasticity observed in the growth chamber correlated well with the observed seasonal phenology. Genetic differences were found in inflorescence height and total plant productivity. The rapid growth rate, the high reproductive effort, and generally broad temperature optimum to moderate temperatures are consistent with the occurrence of P. lanceolata in a wide variety of disturbed habitats in temperate latitudes.     

Contrast adaptation to time constraints on development of two pre-dispersal predators of dandelion (<Emphasis Type="Italic">Taraxacum officinale</Emphasis>) seed

Pre-dispersal seed predators of quickly maturing inflorescences of Asteraceae are constrained by shortage of development time. At seed dispersal, they should pupate or, if still immature, relocate into another inflorescence. To investigate how dominant coleopteran predators of dandelion seed, Glocianus punctiger (Curculionidae) and Olibrus bicolor (Phalacridae), cope with time limitation we combined observation (development and temperature of dandelion capitulum, thermal constants of predator development, age structure of larval populations at seed dispersal) and analogy (“rate isomorphy” in predator development, comparing “model” coleopteran species with similar temperature requirements). Development of a dandelion capitulum takes 21 days. The time available to G. punctiger (140–190 day degrees, development threshold 6.3°C) is sufficient to complete development and pupate after seed dispersal. By contrast, only 30–50 day degrees are available to O. bicolor (threshold 13.5°C) and this is not enough to complete development and consequently immature larvae should move to other capitula to continue feeding until pupation. These contrast strategies which are determined by this thermal adaptation, are accompanied by differences in larval morphology. The “cold adapted” G. punctiger has an apodous larva not capable of migrating between capitula while the “warm adapted” O. bicolor has a mobile campodeiform larva capable of migration.     

THE MORPHOLOGY OF THE INFLORESCENCE OF RAGWEEDS (AMBROSIA-FRANSERIA : COMPOSITAE)

Payne, Willard W. (U. Michigan, Ann Arbor.) The morphology of the inflorescence of ragweeds (Ambrosia-Franseria: Compositae). Amer. Jour. Bot. 50(9): 872–880. Illus. 1963.—The ragweeds possess an inflorescence which is highly modified in relation to anemophily. Several trends from primitive to specialized character expressions may be seen in the morphology of the unisexual capitula and florets. An evolutionary scheme is presented whereby the catkinlike, acropetally maturing, staminate spike and the sessile, centrifugally developing, fruiting involucre cluster are derived from basic inflorescence types within the Compositae. Morphological evidence suggests that: (1) the taxa Ambrosia and Franseria should be combined under the older generic name Ambrosia; (2) the ragweeds and their relatives as a group appear to occupy a position intermediate between the Heliantheae and Anthemideae, as currently delimited within the Compositae.     

Biological characteristics of an invasive south African species

Senecio inaequidens, a south African species that has recently invaded parts of Europe, especially human disturbed habitats, was examined under laboratory and glasshouse conditions to assess the germination of achenes, soil seed banks, growth of seedlings and its breeding system. These observations allowed identification of ecological characteristics that may contribute to the invasiveness of the species. Achenes derived from three populations of S. inaequidens were germinated over a wide temperature range (from 14/6 °C day/night temperatures to a constant temperature of 30 °C). They exhibited a rapid achene germination especially when temperatures reached 20 °C. Germination of seeds from soil samples also exhibited a prolonged germination over time suggesting therefore different types of dormancy. Growth response was studied along a nitrogen gradient from 0.33 to 16.0 mmol l^–1 and data were interpreted using growth analysis. Seedlings of S. inaequidens exhibited a slow growth in low nutrient availability and a rapid growth rate when exposed to high nutrient availability, which contributed to producing a large leaf area and biomass. Four types of pollination were studied in S. inaequidens. Hand-pollinations showed that S. inaequidens was mostly self-incompatible. Self-pollinated capitula only showed 7.6% viable achenes, significantly lower than insect-pollinated and cross-pollinated capitula. Some individuals were more self-fertile than others ensuring the production of some offspring in a situation of colonisation. Natural pollination occurred with generalist insects. Most of these results confirm the invasive ability of S. inaequidens.     

Variation in floral morphology within a population of Aster hispidus var. tubulosus (Asteraceae,Astereae)

The family Asteraceae has a particular inflorescence, the capitulum, consisting of ray florets and disc florets. The ray florets function as petals that attract pollinators. Marked variation in the ray floret morphology is known in a natural population of Aster hispidus var. tubulosus (Asteraceae). We analyzed the variation and found two distinct types in the ray florets, the long tubular ray floret and the ligulate ray floret. In this species, therefore, the variation in floral morphology among capitula, each of which is the basic pollination unit, is caused by the variation in the composition of the two ray floret types among capitula. We evaluated the sources of the observed variation in the floral morphology among capitula within a population using a hierarchical analysis that separated within‐individual (i.e. among capitula within each individual) and between‐individual components of the variation. We found that the main source of the variation lay at the between‐individual level, not at the between‐capitulum level nested within individuals. This finding will provide the basic knowledge that enables future study exploring whether the between‐individual variation in floral morphology caused by the compositional variation of the ray floret types leads to differential pollination success of individual plants in species of Asteraceae.     

Ants and extrafloral nectaries: no evidence for plant protection in Helichrysum spp. — ant interactions

Summary Characterstics of Australian endemic Helichrysum bracteratum and H. viscosum suggest that foraging ants act as guards of developing flowerheads, protecting capitula from seed predators: (1) extrafloral nectar is secreted from leaves subtending the capitula and from bracts encircling the floral disc during pre- to post-flowering periods; (2) capitula are attended by ants; and, (3) encounters between ants and other capitula visitors, including predispersal seed predators such as Tephritis sp. (Diptera), can be frequent. In experiments to test the ant-guard hypothesis, exclusion of ants from plants increased abundance of other insects on the developing capitula. The difference between ant-access and ant-exclusion treatments was related to ant abundance on the access plants. These effects were statistically significant in spite of the large variation in insect activity between sites and through the season.The increased abundance of insects on capitula following ant-exclusion did not, however, result in significant increases in the number of adult seed predators observed on capitula, the number of immature seed predators in capitula, or capitula damage as estimated between ant-access and exclusion treatments of either H. bracteatum or H. viscosum. Further, the ant-exclusion treatment on H. bracteatum had no significant influence on pollination as measured by seed set or on the degree of parasitism of Tephritis sp. by Megastigmus sp. Site and season most strongly affected numbers of immature seed predators and damage to capitula.We discuss these findings in relation to the ant-guard hypothesis and suggest that generalization of the protection hypothesis to all plants with extrafloral nectaries is premature.     

Safflower susceptibility and response to feeding by grasshoppers

Feeding by three grasshopper species, Camnula pellucida, Melanoplus packardii and Melanoplus sanguinipes, on three safflower (Carthamus tinctorius) lines for a 6-wk period from anthesis was monitored under field conditions. Ratings of feeding damage to different plant parts (leaves, floral parts, capitula, and peduncles) and measurements after termination of feeding (dry weight, seed yield, seed weight, seeds per capitulum, and capitula per row) were compared among grasshopper species and safflower lines. The Melanoplus species fed preferentially on leaves, floral parts, and capitula, while C. pellucida exhibited only peduncle feeding, which resulted in head clipping. Defoliation of 20 to 30% was associated with significant increases in total dry matter, seed yield, and number of capitula. Further defoliation resulted in decreases. The safflower lines differed in response to grasshopper feeding. S-208 was most susceptible to defoliation by grasshopper feeding, exhibiting decreased dry weight, seed yield, and capitula number. Lesaf 34C-00 was most tolerant and only M. packardii caused significant dry weight and seed yield reductions. Feeding by C. pellucida on this line resulted in an overall seed yield increase. Feeding by M. sanguinipes on Seedtec-5 resulted in yield increases of up to 16%. It appears that certain grasshopper species can increase seed yield in some safflower lines by stimulating the production of additional capitula. Therefore, moderate populations of such grasshoppers in fields of appropriate safflower cultivars do not necessarily require control.     

Experimental investigation on weather cues for mast seeding of Fagus crenata

Although a recent study has suggested that the minimum temperature from late April to mid-May in the year preceding flowering causes mast seeding in Fagus crenata, no direct evidence is available to support this finding. The aim of the present investigation was, therefore, to test – in a field experiment – whether the minimum temperature determines mast seeding in F. crenata. We examined the effect of nighttime temperatures on flower-bud initiation in F. crenata by enclosing fruit-bearing branches in heated bags at night, thereby maintaining average nighttime temperatures of approximately 2°C above the ambient temperatures. Heating was applied at night from 21 April to 20 May, 21 May to 19 June, and 21 April to 19 June in 2001. Female inflorescence initiation was inhibited by the nocturnal heating in the period 21 April to 20 May and 21 May to 19 June. However, nocturnal heating from 21 April to 20 May was the more important based on the odds ratio of the former being much lower than that of the latter in a logistic regression model. Male inflorescence initiation was also inhibited by nocturnal heating from 21 April to 20 May. We therefore conclude that flower-bud initiation in F. crenata was controlled by nighttime temperatures between 21 April and 20 May.     

Hormone-regulated inflorescence induction and TFL1 expression in Arabidopsis callus in vitro

To study hormone-regulated inflorescence development, we established the in vitro regeneration system of Arabidopsis inflorescences in the presence of cytokinin and auxin. Media containing a combination of thidiazuron (TDZ) and 2,4-dichlorophenoxyacetic acid (2,4-D) were used to induce callus formation. Higher frequencies of calli were obtained by using the inflorescence stems as explants. After transferring the calli to media containing a combination of zeatin and indole-3-acetic acid (IAA), the inflorescences were induced from the calli. The morphology of regenerated inflorescences was similar to that of inflorescences in plants; however, flowers of regenerated inflorescences often lacked a few floral organs. Furthermore, TFL1, a gene involved in floral transition in Arabidopsis, was activated during the inflorescence induction. Our results suggest that the TFL1 gene plays an important role in hormone-regulated inflorescence formation.     

Selective advantage of ray florets in Scalesia affinis and S. pedunculata (Asteraceae), two endemic species from the Galápagos

The presence of neuter ray florets in species within Asteraceae is generally believed to increase pollinator attraction. In the endemic Galápagos genus Scalesia (Asteraceae) a natural variation in the presence/absence of neuter ray florets is found. To evaluate whether the presence of ray florets plays a selective role on female reproductive success we chose two species of Scalesia, Scalesia affinis that carries ray florets and S. pedunculata that is rayless. On Santa Cruz Island capitula of S. pedunculata were equipped with fake ray florets while others were untouched. On Isabela Island ray florets were removed on half of the capitula of S. affinis. In S. affinis rayed capitula received more pollinators and more pollen, which resulted in a significantly higher embryo production. In S. pedunculata no effect on embryo production was found. The disagreement between the two species may correspond to a difference in visitation frequency, S. pedunculata receiving many more visit than S. affinis. Thus, ray floret development proved beneficial in pollinator-restricted localities.     

The Bizarre Inflorescence of Norantea brasiliensis (Marcgraviaceae): Visits of Hovering and Perching Birds1

The pollination biology of Norantea brasiliensis (Marcgraviaceae) was studied in the rain forest of southeastern Brazil. This plant presents bizarre, brush-type racemous inflorescences bearing numerous flowers and extrafloral cup-shaped nectaries. Flower anthesis is diurnal, nectar production is continuous and copious, and the sticky pollen is readily removed by visitors during the first morning hours. The ruby-coloured inflorescences were visited by eight species of hummingbirds (Trochilidae), and 10 species of passerine birds (three Coerebidae and seven Thraupidae). Hummingbirds hovered while probing for nectar and touched flowers occasionally, whereas passerine birds perched and made contact with flowers habitually. Due to differences in flower-visiting and general foraging behaviour, perching birds act as better pollen vectors than hovering birds. The inflorescence of Norantea brasiliensis seems well fitted for pollination by passerine birds, and the hexose-dominated nectar supports this idea. Pollination syndrome trends within Marcgraviaceae may stem from insect-pollinated, condensed and spike-like inflorescences which would give rise to bird-pollinated, brush-type inflorescences. From the same basic condensed inflorescence, bat-pollinated umbelliform inflorescence may be derived from bird-pollinated, pendulous and corymb-like inflorescences. These postulated inflorescence types are found among the extant species of Marcgraviaceae.     

Effects of high temperatures,ash and seed position in the inflorescence on the germination of three Spanish grasses

Abstract. Seed germination of three perennial grasses, Agrostis curtisii, Avenula marginata and Pseudarrhenatherum longifolium was studied in relation to the position (basal, intermediate, apical) in the panicle and two factors typically associated with fire, high temperatures and ash. Two similar germination responses were observed in A. curtisii and A. marginata: (1) germination rates were lower for seeds from higher positions in the inflorescence; and (2) watering with a plant-ash solution improved the ability of apical seeds to germinate, but had the opposite effect on basal seeds. In A. curtisii, the latter response was also dependent on whether or not the seed was exposed to high temperatures. In P. longifoliium the exposure to high temperatures and the position of the seeds in the inflorescence influenced the effect of ash. This could be due to the different degrees of development of the seeds.     

Fruit size decline from the margin to the center of capitula is the result of resource competition and architectural constraints

Plants produce repeated structures, such as leaves, flowers, and fruits, which differ in size and shape. One example of this is fruit size, which is commonly observed to decrease from proximal to distal positions within an inflorescence. The resource limitation hypothesis proposes that because proximal fruits usually develop first, they have temporal priority on access to resources over distal fruits. The non-mutually exclusive architectural effects hypothesis suggests that these position effects in fruit size may also be due to inherent architectural variation along infructescence axes. We separated out the effects of resource competition and inflorescence architecture by removing the outer or the inner flowers within capitula of Tragopogon porrifolius. We also studied if fruit position influenced germination and seedling performance in order to assess fitness consequences of position effects. Inner fruits were significantly heavier when outer flowers were removed. However, outer fruits did not significantly increase when inner flowers were removed, suggesting later fruits were limited by the development of early fruits. Our findings also suggest that architectural constraints restricted the size of inner fruits in comparison with outer ones. We found that both resource competition and inflorescence architecture affected the fruit size of T. porrifolius, even though this species does not have linear, indeterminate inflorescences. We advance the hypothesis that, when such effects on fitness occur, resource competition-mediated position effects could turn, in evolutionary time, into architectural position effects.     

影响黄帚橐吾种子生产的因素Ⅰ.生境和花序结构

 通过对黄帚橐吾(Ligularia virgaurea)自然种群在不同生境和种群密度下的种子生产的研究,结果显示：1）黄帚橐吾的种子生产无论是在种群水平,还是在总花序上同一部位头状花序水平,均受到生境条件的影响,其种子数量、种子均重和种子总重量等特征在不同生境间表现出沙地＞滩地＞坡地,在同一生境内则为高密度种群＞低密度种群和植被盖度较低＞植被盖度较高的趋势（滩地例外）;2）种子生产中的种子数量和种子总重与总花序大小、头状花序数量以及无性系株大小多呈显著正相关,与个体密度无关,但种子均重与上述因素（除无性系株大小外）的关系不显著;3）在总花序内不同部位头状花序间表现出的种子生产变化趋势（顶部＞基部）不受生境条件、种群密度等因素的影响; 4）种子生产在总花序上所表现出的这种部位依赖性可能是由于不同部位头状花序间开花时序上的差异引起种子从母体获取资源上的不同所致。