Experimental evidence for pollination of Banksia spp. by non-flying mammals

Summary The importance of non-flying mammals as pollinators of Banksia integrifolia and B. spinulosa was analysed by examining the effect of pollinator exclusions on fruit-set. Visitation by potential pollinators was also measured by observation and by indirect methods. Nonflying mammals were frequent visitors to inflorescences of both Banksia species. The aluminium sleeves used to exclude non-flying mammals from B. integrifolia trees were associated with a reduction in both the number of infructescences produced and the number of fruit per infructescence, indicating that non-flying mammals were important pollinators. Bird-nets over trees also significantly reduced the number of fruit per infructescence, but had no significant effect on the number of infructescences produced. The results of exclusion experiments using single inflorescences were inconclusive due to low fruitset. No conclusions could be drawn from these experiments with B. spinulosa. However, results for B. integrifolia support the conclusions of whole-tree experiments. Analysis of the genotype frequencies in seed from B. integrifolia provided no support for the hypothesis that the relatively limited mobility of non-flying mammal pollinators would cause inbreeding.     

Flowering dynamics in Arum italicum (Araceae): relative role of inflorescence traits, flowering synchrony, and pollination context on fruit initiation

We studied the relative role of inflorescence traits, flowering synchrony, and pollination context for infructescence and fruit initiation in two Spanish populations of Arum italicum, a species in which inflorescences are the pollination unit. In this species, a specialized inflorescence organ, the appendix, is important for pollinator attraction. However, the short floral longevity and the production of mostly one inflorescence per plant make its pollination potentially dependent on strong flowering synchrony and on external factors not controlled by the plant (the pollination context). The flowering period in both sites lasted >3 mo. Day-to-day variation in simultaneous antheses was high, and 11-50% of antheses occurred on days during which no pollen donor was present. Inflorescence traits, flowering synchrony, and between-plant distance all influenced infructescence and fruit initiation, but their relative importance differed between sites. In one large population, infructescence initiation was positively related to inflorescence traits; in a smaller population infructescence initiation increased with the number of donor inflorescences. In both sites, percentage of fruits initiated per infructescence was dependent on a combination of inflorescence traits, flowering synchrony, and between-plant distance. Plants producing 2-4 inflorescences had higher probability of infructescence initiation and overlapped their antheses with more plants than single-inflorescence ones.     

Gender modification in a monoecious species <Emphasis Type="Italic">Sagittaria potamogetifolia</Emphasis> (Alismataceae)

In order to find out if the inflorescences number variation has influences on the gender modification in plant species, we investigated the gender modification in a cultivated population of the monoecious species Sagittaria potamogetifolia. We also designed two nutrient levels to explore the impact of nutrient on gender modification in S. potamogetifolia. We found that the female and male flowers did not change with increasing plant size for each inflorescence at a low nutrient level. At a high nutrient level, the female flower numbers on each inflorescence did not increase with plant size; however, the male flower numbers had some positive correlation with the plant size. At the ramet level, the total male and female flower numbers increased with the plant size at both nutrient levels. The sex ratio (female to male flower ratio) decreased with the inflorescence numbers and the plant size (Midvein length). Although the nutrient variation had impact on the flower number production, it did not change the gender modification pattern. The high plasticity of inflorescence numbers, which caused the gender variation in S. potamogetifolia, and low plasticity of female and male flowers on a single inflorescence, indicates that the limited modification on gender in a single inflorescence may be compensated by inflorescence number variation at the ramet level.     

Causes of spatial patterns of fruit set in waratah: Temporal vs. spatial interactions between flowers on an inflorescence

Spatial patterns of fruit set within inflorescences may be controlled by pollination, nutrient allocation, or inflorescence architecture. Generally, flowers that have spatial and/or temporal precedence are more likely to set fruits. We sought to separate these factors by comparing patterns of fruit set on inflorescences of two species of Telopea (Proteaceae); one that flowers from the tip to the base of the rachis, the other from base to tip. In both species, most fruits were set at the top of the inflorescence (the last flowers to open for T. speciosissima) and this was extreme for T. mongaensis, where the top flowers open first. Fruit set was not generally limited by inadequate pollination for either T. mongaensis or T. speciosissima, as hand pollinations did not increase fruit set and many abscised flowers contained pollen tubes. In T. speciosissima, we tested whether removal of developing topmost fruits would ‘release’ those that had initiated but not yet aborted lower down. There was no significant effect. Plant hormones can increase the degree to which a developing fruit is a sink for nutrients, so we applied cytokinin to the developing lower fruits on some inflorescences. There was no significant effect of the hormone treatment. We conclude that temporal precedence may contribute to the skewed pattern of fruit set in T. mongaensis, because there was an extreme concentration of fruit set on the distal part of the inflorescences, but it cannot explain this pattern of fruit set in T. speciosissima, where the distal flowers are the last to open. Some other process must therefore constrain fruit set to the topmost flowers in an inflorescence. While cytokinin application had no significant effect, the power of this experiment was low and we consider that the hypothesis of hormonal control is worth further exploration.     

Fruit set,fruit reduction,and fruiting strategy in Cornus sanguinea (Cornaceae)

We studied various aspects of the fruiting biology of Cornus sanguinea (Cornaceae), a fleshy-fruit-producing deciduous shrub, in four populations in northwest Spain. One population was studied over a 5-yr period (1989-1994), and the remaining populations in 1994 only. Fruit-set level varied among years (range 11-18%) and among populations (range 8-22%), but was in all cases low. Within plants, fruit-set level did not vary significantly among inflorescences, indicating that inflorescence fruit set is independent of inflorescence position and inflorescence, phenology. To investigate the function of surplus flowers, we carried out flower removal experiments. Inflorescence fruit-set level was unaffected by removal of up to =75% of flowers. These results suggest that Cornus sanguinea regulates its fruit-set level via plasticity in the number of fruits aborted: if flower mortality has been high, fewer fruits will be aborted. Within the inflorescence, surplus flowers thus act as insurance against flower loss.     

In Vitro Culture of Kodo Millet: Influence of 2,4-D and Picloram in Combination with Kinetin on Callus Initiation and Regeneration

Immature inflorescences of kodo millet (Paspalum scrobiculatum L. cv. GPUK-3) were cultured on MS medium. Induction of embryogenic callus and subsequent somatic embryogenesis was possible on both 2,4-D and Picloram alone or with kinetin from spikelets as well as rachis. Immature inflorescence cultured on medium supplemented with lower levels of Picloram in combination with kinetin developed organogenic callus with shoot buds. Direct somatic embryo formation on rachis was observed at higher levels of Picloram in combination with kinetin. Plant regeneration was observed when calluses were transferred to α-napthaleneacetic acid (NAA) plus 6-benzylaminopurine (BA) supplemented MS medium. Histological observations provided a clear evidence for both somatic embryogenesis and shoot organogenesis. Profuse rooting was induced on phenylacetic acid (PAA) supplemented medium. Regenerated plants were successfully transferred to pots under field conditions where most of the plants survived and set normal seeds.     

Do pollinators influence seed-set in Banksia paludosa Sm. and Banksia spinulosa R. Br.?

The consequences of observed variations in pollinator activity for plant reproduction depend on the breeding system of the plant and also on whether pollination is a resource in short supply. These factors are frequently assumed rather than tested. In bagging experiments with Banksia paludosa in heath and B. spinulosa var. spinulosa in woodland, the present study demonstrated that seed-set does not occur by autogamy and that pollinators are indeed necessary. Experimental addition of outcrossed pollen to inflorescences failed to produce any increase in the mean number of seeds per inflorescence, measured for non-barren inflorescences, for either species. However, in B. paludosa, the addition of pollen increased the numbers of inflorescences that set seed from 20% (controls) to 45% (pollen-added treatment). This suggests either unusual patterns of pollen movement among and within plants or control of the pattern of seed-set within a plant. Pollen additions failed to alter seed-set in B. spinulosa. It is possible that a greater abundance of pollinators in the woodland ensures adequate pollination for these plants     

Reproductive biology of Arum italicum (Araceae) in the south of France

The reproductive success of Arum italicum in south-western France showed a positive relationship with plant vigour at both the plant (number of inflorescences) and inflorescence (number of seeds) levels. However, the infructescence and berry numbers were not related to the number of inflorescences, and two-fold more seeds were produced by individuals flowering several times than by those flowering just once. Moreover, the reproductive success of this species appeared to be limited by the high abortion rate (50%), strong predation pressure on the inflorescences (34%), and low seed set rate of nonpredated inflorescences (36%), suggesting pollination limitation. Nevertheless, a positive relationship was found between the mean number of Psychoda trapped per inflorescence and the number of maturing infructescences, particularly for the most abundant species, Psychoda crassipenis . Therefore, pollinator availability appeared to be an important limiting factor for fructification in the studied population, by contrast with other studies in which the fruit set appeared to be related more strongly to the density of inflorescences. Such different results may have been caused by the relatively low abundance of Psychoda in the present study.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 43–49.     

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.     

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.     

Oviposition preferences and larval performance of a flower-feeding weevil,Coelocephalapion aculeatum, in relation to host development

The oviposition behaviour ofCoelocephalapion aculeatum Fall (Coleoptera: Apionidae), a host specific florivore exploiting an ephemeral resource, was studied in relation to host development. Immature inflorescences of its host, the weedMimosa pigra L. (Mimosaceae), were categorised into 5 developmental stages ranging from first appearance to one day before opening. The duration, weight, percent abortion, and nitrogen content of each stage were measured. Oviposition preferences ofC. aculeatum for all inflorescence stages were determined in multiple choice trials. Newly hatched, larvae were transferred into inflorescences of each stage to determine larval survival as a function of stage. Numbers of eggs laid into inflorescences generally correlated with larval survival except that oviposition was greater in stage 4 inflorescences than expected from the larval survival. Numbers of eggs laid into inflorescences generally correlated with their carrying capacity except that adults avoided stage 5 inflorescences in which larval survival was low. The evolution of oviposition preferences ofC. aculeatum appears to have been influenced by two opposing selective forces. The need to complete development before the inflorescence deteriorated would have selected for oviposition in young inflorescences. The greater availability of food and lower probability of abortion offered by older inflorescences would have selected for oviposition in these. The net outcome of these pressures has resulted in the observed oviposition strategy.     

小叶桦花序生长物候及其生态适应意义

小叶桦(Betula microphylla)是一种典型的荒漠和山地乔木植物,在中国仅分布于新疆。为了明确小叶桦花序生长物候规律及发育特征,该研究对小叶桦进行了花序物候观测,分析其生长规律及果实结籽情况。结果显示:小叶桦雄花序为越年生殖器官,物候周期为345d,其中营养生长期154d、休眠期158d、开花生长期33d;雌花序与果序为当年生殖器官,物候期为105d,其中雌花序生长期24d、果序生长期90d。虽然雄花序比雌花序的生长周期明显较长,但开花授粉均在4月中下旬,雄花序散粉期和雌花序可授粉期之间具有较高的同步性和协调性,表现出集中开花授粉模式,同时这种开花模式在自然条件下,果序的结籽数和结籽率分别为220和76.7%,种子库中具有活力的种子约每平米4万粒,表明小叶桦在荒漠极端环境中能顺利完成有性生殖过程。     

Behavioral responses by bumble bees to variation in pollen availability

Summary Pollen-collecting bumble bees (Bombus spp.) detect differences between individual flowers in pollen availability and alter their behavior to capitalize on rewarding flowers. Specific responses by bees to increased pollen availability included: longer visits to flowers; visits to more flowers within an inflorescence, including an increased frequency of revisits; an increased likelihood of grooming while the bee flow between flowers within the inflorescence; and more protracted inter-flower flights, probably because of longer grooming bouts. The particular suite of responses that a bee adopted depended on the pollen-dispensing mechanism of the plant species involved. Bees buzzed previously-unvisited Dode-catheon flowers longer than empty flowers. In contrast, pollen availability did not significantly affect the duration of visits to Lupinus flowers, which control the amount of pollen that can be removed during a single visit. Simulation results indicate that the observed movement patterns of bumble bees on Lupinus inflorescences would return the most pollen per unit of expended energy. The increased foraging efficiency resulting from facultative responses by bees to variation in pollen availability, especially changes in the frequency and intensity of grooming, could correspondingly decrease pollen dispersal between plants.     

Effectiveness of nectarivorous birds and honeybees as pollinators of Banksia spinulosa (Proteaceae)

Abstract The effectiveness of nectarivorous birds and honeybees (Apis mellifera) as pollinators of Banksia spinulosa (Proteaceae) was investigated. Birds visited inflorescences in the early, mid and late flowering seasons. In contrast, honeybees visited only on days in the late flowering period when maximum temperatures exceeded 15°C. Self pollen remained on pollen presenters of flowers for up to 5 days in the early and mid periods. In the late period, when honeybees visited inflorescences, self pollen was removed within 2 days. Pollen removal was similar for caged (birds excluded) and open inflorescences in the late period, indicating that most pollen was removed by honeybees. In the early and mid periods, honeyeaters pollinated 22% and 27% of flowers on open inflorescences, respectively. In the late period, when both birds and bees visited inflorescences, 64–73% of flowers on open inflorescences were pollinated. Foraging by honeybees resulted in pollen deposition as 38% of flowers on caged inflorescences were pollinated. Throughout the flowering season a similar number of pollen grains was deposited per stigma. There were 3.0–3.7 pollen grains per stigma on open inflorescences in the late period, although only 2.0 grains per stigma on caged inflorescences. In the early and mid periods, fewer caged than open inflorescences produced fruits, indicating the importance of honeyeaters to reproductive success at these times. In contrast, in the late period when honeybees visited inflorescences, fruit-set was similar on caged and open inflorescences. Overall, these results indicate that honeybees were effective pollinators of B. spinulosa.     

Analysis of inflorescence organogenesis in eastern gamagrass, Tripsacum dactyloides (Poaceae): the wild type and the gynomonoecious gsf1 mutant

Inflorescence organogenesis of a wild-type and a gynomonoecious (pistillate) mutant in Tripsacum dactyloides was studied using scanning electron microscopy. SEM (scanning electron microscope) analysis indicated that wild-type T. dactyloides (Eastern gamagrass) expressed a pattern of inflorescence organogenesis that is observed in other members of the subtribe Tripsacinae (Zea: maize and teosinte), family Poaceae. Branch primordia are initiated acropetally along the rachis of wild-type inflorescences in a distichous arrangement. Branch primordia at the base of some inflorescences develop into long branches, which themselves produce an acropetal series of distichous spikelet pair primordia. All other branch primordia function as spikelet pair primordia and bifurcate into pedicellate and sessile spikelet primordia. In all wild-type inflorescences development of the pedicellate spikelets is arrested in the proximal portion of the rachis, and these spikelets abort, leaving two rows of solitary sessile spikelets. Organogenesis of spikelets and florets in wild-type inflorescences is similar to that previously described in maize and the teosintes. Our analysis of gsf1 mutant inflorescences reveals a pattern of development similar to that of the wild type, but differs from the wild type in retaining (1) the pistillate condition in paired spikelets along the distal portion of the rachis and (2) the lower floret in sessile spikelets in the proximal region of the rachis. The gsf1 mutation blocks gynoecial tissue abortion in both the paired-spikelet and the unpaired-spikelet zone. This study supports the hypothesis that both femaleness and maleness in Zea and Tripsacum inflorescences are derived from a common developmental pathway. The pattern of inflorescence development is not inconsistent with the view that the maize ear was derived from a Tripsacum genomic background.     

The effect of varying nitrogen and phosphorus availability on nutrient use by Larrea tridentata,a desert evergreen shrub

Summary In a phytotron study of the effects of nitrogen and phosphorus supply ratio on nutrient uptake and use by Larrea tridentata, seedlings responded to increases in N and P availability with increases in leaf size, total biomass, and leaf nutrient concentration, and with decreases in root: shoot ratio. N and P use efficiency decreased with increasing N and P availability, respectively, but increased with increasing availability of the other nutrient, suggesting that Larrea responds both to the absolute and to the relative availability of limiting nutrients. Absolute amounts of N and P resorption, as well as N and P resorption efficiencies did not demonstrate a significant trend with nutrient availability, and there was no evidence of significant interactions between the two nutrients. More studies of the effects of nutrient interactions in the cycling and use of nutrients by different plant species are needed before more general conclusions can be drawn.     

Vascular development of the grapevine (<Emphasis Type="Italic">Vitis vinifera</Emphasis> L.) inflorescence rachis in response to flower number,plant growth regulators and defoliation

The grapevine inflorescence is a determinate panicle and as buds emerge, shoot, flower and rachis development occur simultaneously. The growth and architecture of the rachis is determined by genetic and environmental factors but here we examined the role of flower and leaf number as well as hormones on its elongation and vascular development. The consequences of rachis morphology and vascular area on berry size and composition were also assessed. One week prior to anthesis, Merlot and Cabernet Sauvignon field vines were exposed to manual flower removal, exogenous plant growth regulators or pre-bloom leaf removal. Manual removal of half the flowers along the vertical axis of the inflorescence resulted in a shorter rachis in both cultivars. Conversely, inflorescences treated with gibberellic acid (GA₃) and the synthetic cytokinin, 6-benzylaminopurine (BAP) resulted in a longer rachis while pre-bloom removal of all leaves on the inflorescence-bearing shoot did not alter rachis length relative to untreated inflorescences. Across the treatments, the cross-sectional areas of the conducting xylem and phloem in the rachis were positively correlated to rachis girth, flower number at anthesis, bunch berry number, bunch berry fresh mass and bunch sugar content at harvest. Conversely, average berry size and sugar content were not linked to rachis vascular area. These data indicate that the morphological and vascular development of the rachis was more responsive to flower number and plant growth regulators than to leaf removal.     

Foliar and twig macronutrients (N,P, K,Ca and Mg) in selected species ofEucalyptus used in rehabilitation: sources of variation

Summary The macronutrient variation within four 6 year oldEucalyptus saligna and four 5 year oldE. wandoo growing on rehabilitated bauxite pits was determined. Significant differences in mean nutrient concentrations were generally recorded between good soil condition sites and poor soil sites, between tree individuals, branch height, and plant organ type; but mean nutrient values were not different among canopy aspects. Fully expanded leaves of the current year provided the most uniform nutrient levels among the plant organs and showed major differences between sites with good soil nutrient conditions and those with poor conditions. Differences in foliar and branch levels of N, P, K, Ca and Mg, the variation between sites, canopy heights and plant organ types, and the use of foliar nutrient levels to indicate deficiencies are discussed.     

Reproductive biology of the rare and endangered Banksia brownii Baxter ex R. Br. (Proteaceae)

Abstract Banksia brownii is an endangered species, now limited to ～ 15 disjunct populations in southwestern Western Australia. Data on flowering phenology, plant size, fruit set, pollination and the mating system were gathered for two of these populations between March and October 1993. Flowering for both populations followed a similar pattern, with open flowers first evident in April, and the number of inflorescences with open flowers peaking in June. At both locations, trees differed considerably with respect to their size, the total number of inflorescences produced and the length of their flowering season. Fruiting success was typically low, with approximately half of all inflorescences failing to develop into infructescences. Only 1. 8% of the flowers originally present on inflorescences developed into follicles. The distribution of follicles along each infructescence was non-random, with most forming in the middle third of the infructescence for reasons relating to nutrient supply and pollinator behaviour. More flowers opened during the day than at night, although pollen was lost from individual flowers during both periods. Honeyeaters such as Phylidonyris novaehollandiae were common at the two study sites, and often carried large loads of B. brownii pollen. Though less frequently caught, the nocturnal mammals Rattus fuscipes and Tarsipes rostratus also bore substantial amounts of pollen. Most inflorescences from which these mammals and birds were excluded remained barren. Fruiting success was further reduced when invertebrates such as Apis mellifera were also prevented from visiting inflorescences. The ability of B. brownii to set at least some fruit in the absence of biotic poli-nators indicates that the species is partially self-compatible. Honeyeaters foraged preferentially at inflorescences with one to two thirds of their flowers open, probing mainly along the ‘advancing front’ of open flowers. These animals moved more frequently between inflorescences on the same plant than between those on different plants, and were often recaptured in the same locations. Mammals also appeared to be sedentary. Both B. brownii populations had mixed mating systems, with genetically determined outcrossing rates of ～0.7. The unusually high level of selfing in each population is presumably a reflection of the species’ self-compatibility and the foraging behaviour of its pollinators.