Pollination intensity and potential seed set in Passiflora vitifolia

Summary Initial seed set and fruit set were pollen-limited in a Costa Rican population of Passiflora vitifolia, a self-incompatible species with 200–350 ovules per flower. Pollination intensity was measured by counting the number of allogamous pollen grains on stigmas of the large one-day flowers. Hand-pollinations demonstrated that 25–50 pollen grains are required for fruit set, and >450 are needed for maximum seed set, with a pollen:seed ratio of about 1.6:1.0. Hummingbirds (Phaethornis superciliosus) delivered sufficient allogamous pollen for maximum seed set to only 28% of the flowers examined. Naturally pollinated flowers yielded fewer fruits and fewer seeds per fruit than those pollinated by hand. Most pollen transferred by humming-birds was self-incompatible; emasculated flowers yielded higher seed set than flowers with intact anthers. Visitation rates did not provide a good index of effective pollination.There were significant differences in ovule number, maximum seed set, and maximum per cent seed set among individual vines. More than half of an individual's flowers failed to set fruit, whether pollinated by birds or by hand. In this population, maximum reproductive potential may be limited by maternal resources for fruit development, but seed set varies with pollination intensity. Pollen-limited seed set may be a disadvantage of self-incompatibility, especially in species with many-seeded fruits.     

Pollination, seed set and seed predation on a landscape scale

We analysed the combined effects of pollination and seed predation on seed set of Centaurea jacea in 15 landscapes differing in structural complexity. In the centre of each landscape, a patch of Centaurea plants was established for standardized measurements of flower visitation, seed predation and seed set. Both the number of flower-visiting bees and the proportion of flower heads damaged by seed predators increased with landscape complexity, which was measured as the proportion of semi-natural habitats. The mean number of seeds per flower head did not increase with the proportion of semi-natural habitats, presumably because of the counterbalancing effects of pollination and seed predation. For a subset of undamaged flower heads, the number of seeds per flower head was positively correlated with the number of flower visits. Further reasons for the unexpected failure to detect a correlation between landscape complexity and seed set appeared to be changes in flower-visitor behaviour and the contrasting responses of honeybees and wild bees to habitat context. Landscape analyses at eight spatial scales (radius of landscape sectors, 250-3000 m) showed that different groups perceived the landscape at different spatial scales. Changes in pollinator numbers could be explained only at small scales (up to 1000 m), while the seed predators also responded to large scales (up to 2500 m).     

Heteropogon-Themeda grasses evolve to occupy either tropical grassland or wetland biomes

Species of the Heteropogon-Themeda clade are ecologically important grasses distributed across the tropics, including widespread species, such as the pantropical Heteropogon contortus and Themeda triandra, and range-restricted species such as Heteropogon ritchiei and Themeda anathera. Here, we examine habitat preferences of the grassland/savanna and wetland species by describing bioclimatic niche characteristics, characterizing functional traits, and investigating the evolution of functional traits of 31 species in the Heteropogon-Themeda clade in relation to precipitation and temperature. The climatic limits of the clade are linked to mean annual precipitation and seasonality that also distinguish seven wetland species from 24 grassland/savanna species. Tests of niche equivalency highlighted the unique bioclimatic niche of the wetland species. However, climatic factors do not fully explain species geographic range, and other factors are likely to contribute to their distribution ranges. Trait analyses demonstrated that the wetland and grassland/savanna species were separated by culm height, leaf length, leaf area, awn length, and awn types. Phylogenetic analyses showed that the wetland species had tall stature with long and large leaves and lack of hygroscopic awns, which suggest selective pressures in the shift between savanna/grassland and wetland. The two most widespread species, H. contortus and T. triandra, have significantly different bioclimatic niches, but we also found that climatic niche alone does not explain the current geographic distributions of H. contortus and T. triandra. Our study provides a new understanding of the biogeography and evolutionary history of an ecologically important clade of C₄ tropical grasses.     

Quantitatively determined self-incompatibility. IV. Pollination and seed set in Borago officinalis

Summary The outcomes of sequential double pollination, mixed double pollination and single pollination are compared. Single pollination leads to lower seed set than double pollination. Systematic differences between female genotypes are shown to be possible. It is also shown that failure to set seed is generally due to pre-zygotic maternal factors.     

Pollination by deceit,floral sex ratios and seed set in dioecious Rubus chamaemorus L.

Summary Male and female flowers of the dioecious perennial herb Rubus chamaemorus L. are similar in general appearance. However, female flowers are somewhat smaller, do not produce any pollen, and contain very small amounts of nectar. Syrphids and bumblebees, which are important pollinators of R. chamaemorus, showed a strong preference for male flowers. Male flowers were also less often rejected by flower visitors than were female flowers, and two different groups of syrphid species stayed longer in male than in female flowers. These observations suggest that female flowers of R. chamaemorus attract pollinators by deceit.Hand-pollination experiments indicated that pollen availability limited seed production of R. chamaemorus in female dominated habitats but not in areas with an equal floral sex ratio. We suggest that the relative importance of factors limiting female reproductive success is not constant, but is influenced by the floral sex ratio of the population. This should apply also to other dioecious species that show variable sex ratios on either a local or regional scale.     

Pollination biology, mating system, and seed set in a Danish population of Saxifraga granulata

Saxifraga granulata is able to reproduce both sexually and vegetatively. In an earlier report from Denmark, seed set was said to be low, the species reproducing mainly vegetatively by means of bulbils. Furthermore, investigations in northern England have shown inbreeding depression in gynodioecious 5. granulata populations. The present study, however, demonstrates that 5. granulata is fully self-compatible in Denmark; seed set per flower in hand-selfings was on average 70%, compared to ca. 80% in control plants. Nevertheless, despite high seed set, reproduction by bulbils is probably the most important mode of reproduction. Saxifraga granulata has typically opportunistic flowers which are white, UV-neutral compared to the background reflexion, and offering only a small standing crop of nectar. Optical attraction by the massive flowering over a relatively short period is regarded as a way of overcoming competition from co-occurring, simultaneously flowering species. Saxifraga granulata flowers received visits from at least 27 different insects species. Based on number of recorded flower visits, records on foraging time, and investigations of body surface pollen loads, Rhamphomyia sulcata (Empididae), Cheilosia spp. (Syrphidae), Polyblepharis opaca (Empididae), and Halictus spp. ( s. I .) (Halictidae) were found to be the most important pollen vectors in the 5. granulata population.     

Genetic modification of wetland grasses for phytoremediation

Wetland grasses and grass-like monocots are very important natural remediators of pollutants. Their genetic improvement is an important task because introduction of key transgenes can dramatically improve their remediation potential. Tissue culture is prerequisite for genetic manipulation, and methods are reported here for in vitro culture and micropropagation of a number of wetland plants of various ecological requirements such as salt marsh, brackish water, riverbanks, and various zones of lakes and ponds, and bogs. The monocots represent numerous genera in various families such as Poaceae, Cyperaceae, Juncaceae, and Typhaceae. The reported species are in various stages of micropropagation and Arundo donax is scaled for mass propagation for selecting elite lines for pytoremediation. Transfer of key genes for mercury phytoremediation into the salt marsh cordgrass (Spartina alterniflora) is also reported here. All but one transgenic lines contained both the organomercurial lyase (merB) and mercuric reductase (merA) sequences showing that co-introduction into Spartina of two genes from separate Agrobacterium strains is possible.     

Responses to drought and flooding in tropical forage grasses

Seasonal drought and flooding severely limit pasture growth in tropical savannas. The objective of this study is to analyze and compare yield, biomass allocation, leaf growth rate and nutrient concentration of four important perennial C₄ forage grasses to short term flooding and moderate drought under controlled conditions. The grasses studied were the tufted Andropogon gayanus (CIAT 621) and Hyparrhenia rufa and the stoloniferous Echinochloa polystachya and Brachiaria mutica. All grasses were able to adjust their growth and development in response to flooding and drought: leaf growth and total biomass decreased under both treatments but the specific responses to these treatments differed markedly. Considering only total yield and leaf area, A. gayanus and H. rufa were relatively more tolerant to and less affected by drought whereas B. mutica and E. polystachya were more flood tolerant. In A. gayanus and H. rufa, both treatments reduced the proportion of assimilates devoted to roots and culms while increasing that of leaves decreasing the root/shoot ratio. In contrast, in B. mutica and E. polystachya only the proportion devoted to culms or stolons increased under flooding but the root/shoot ratio remained relatively stable under both treatments. All grasses produced adventitious rootlets except A. gayanus which was the most affected by flooding. Waterlogging decreased leaf nutrient concentration in all grasses which contributed to growth reduction. All species were relatively tolerant to both stresses. The results confirm the empirical observation that stoloniferous species B. mutica and E. polystachya are more tolerant to flooding thanks to adaptations typical of wetland plants such as hollow stolons which enhance oxygen diffusion to the roots and the development of adventitious rootlets that promotes water and nutrient absorption.     

Nitric oxide accelerates seed germination in warm-season grasses

The nitric oxide (NO) donor sodium nitroprusside (SNP) significantly promoted germination of switchgrass (Panicum virgatum L. cv Kanlow) in the light and in the dark at 25°C, across a broad range of concentrations. SNP also promoted seed germination in two other warm-season grasses. A chemical scavenger of NO inhibited germination and blocked SNP stimulation of seed germination. The phenolic (+)-catechin acted synergistically with SNP and nitrite in promoting seed germination. Acidified nitrite, an alternate NO donor also significantly stimulated seed germination. Interestingly, sodium cyanide, potassium ferricyanide and potassium ferrocyanide at 200 μM strongly enhanced seed germination as well, whereas potassium chloride was without effect. Ferrocyanide and cyanide stimulation of seed germination was blocked by an NO scavenger. Incubation of seeds with a fluorescent NO-specific probe provided evidence for NO production in germinating switchgrass seeds. Abscisic acid (ABA) at 10 μM depressed germination, inhibited root elongation and essentially abolished coleoptile emergence. SNP partially overcame ABA effects on radicle emergence but did not overcome the effects of ABA on coleoptile elongation. Light microscopy indicated extension of the radicle and coleoptiles in seeds maintained on water or on SNP after 2 days. In contrast, there was minimal growth of the radicle and coleoptile in ABA-treated seeds even after 3–4 days. These data indicate that seed germination of warm-season grasses is significantly influenced by NO signaling pathways and document that NO could be an endogenous trigger for release from dormancy in these species.     

Density‐dependent pre‐dispersal seed predation and fruit set in a tropical tree

Negative density‐dependent demographic processes operating at post‐dispersal seed, seedling, and juvenile stages are the dominant explanation for the coexistence of high numbers of tree species in tropical forests. At adult stages, the effect of pollinators and pre‐dispersal fruit predators are often dependent on the density or abundance of flowers and fruit in the canopy, but each have opposite effects on individual realized reproduction. We studied the effect of density on total and mature fruit set and pre‐dispersal predation rates within individual tree canopies in a common canopy tree species, Jacaranda copaia in a 50‐ha forest census plot in central Panama. We sampled all reproductive sized trees in the plot (n = 188) across three years and estimated fruit set and predation rates. Population‐wide pre‐dispersal seed predation averaged between 6–37% across years. Using linear mixed effects models, we found that increased density and fecundity of conspecific neighbours increased focal tree fruit set, but also the rate of pre‐dispersal predation. An interaction between individual and neighbourhood fruit production predicted lower predation rates at high individual and neighbourhood fecundities, which suggests predator satiation at high fruit abundance levels. However, the rate at which fruit set increased with conspecific neighbour fruit production was greater than the rate at which fruit were lost to predation, resulting in an overall positive effect of neighbour density on mature fruit production in focal trees. Our results run counter to the expectation of a uniformly negative effect of density across all life stages in tropical trees and suggest further exploration of the role of spatial clumping, pollen dispersal limitation, and predation at pre‐dispersal adult stages in maintenance of species diversity in plant communities.     

Pollination ecology of Christmas Bells (Blandfordia nobilis): Effects of pollen quantity and source on seed set

The effect of a number of experimental handpollination regimens on the numbers of seeds set by Blandfordia nobilis flowers was assessed. Individuals were found to be self-incompatible and to require pollinator visits to set fruit. Pollen limited the female component of reproduction in the two populations studied while pollen source did not have a significant effect on the numbers of seeds set. Experiments using a stuffed honeyeater indicated that maximal seed set could be achieved after just one probe by the pollinator. Appreciable heterogeneity in flowering phenology, the number of seeds naturally setting and the number of seeds setting as a result of hand-pollinations existed within a small area across a single flowering season.     

Ensilage of tropical grasses mixed with legumes and molasses

The effects of adding two legumes, Gliricidia sepium and Leucaena leucocephala, cv. Cunningham, and molasses on the fermentation characteristics of silages made from two tropical grasses (Pangola grass, Digitaria decumbens, and Setaria sphacelata cv. Kazungula) were investigated. Pangola grass silages contained significantly higher contents of water-soluble carbohydrates and lactic acid than did setaria silages after 100 days fermentation, but there were no significant differences between the two silages in populations of lactic acid bacteria and contents of total N and NH₃–N. Addition of either species of legume had no significant effect on fermentation acids and NH₃–N contents, and numbers of lactic acid bacteria. Addition of both legumes reduced NH₃–N production in the silages by 59% after 5 days' fermentation. Numbers of lactic acid bacteria were not significantly affected by the different treatments. Enterococcus faecalis represented 60% of the lactic acid bacteria isolated from the treated herbages prior to ensiling. By 100 days of fermentation, only lactobacilli were isolated: 82% homo-fermenters and 18% hetero-fermenters. Lactobacillus mesenteroides subsp. dextranicum was found only in the silage supplemented with 33% (w/w) legume. It was concluded that the low quality of tropical grasses used as feeds for ruminants may be significantly improved by ensiling these grasses with small amounts of molasses and with high-protein tree leaves.M. Tjandraatmadja and B.W. Norton are with the Department of Agriculture. The University of Queensland, Queensland, 4072, Australia; I.C. Mac Rae is with the Department of Microbiology, The University of Queensland, Queensland, 4072, Australia.     

Building global collaboration in tropical wetland research

This article was derived from a SWS symposium held in 2007 and describes research undertaken in the tropics and subtropics by partners in the Global Wetland Consortium. It explores the role of global collaboration in science and tropical wetlands and maps out a pathway for enhanced inter-change and exchange of information and practical experience. Whilst scientific initiatives for a range of earth systems have been previously been developed, a systematic and global research initiative for wetlands that crosses cultural, scientific and economic divides is not yet operational. One option when considering such initiatives is to investigate the utility of using the more than 1800 wetlands designated as internationally important under the Ramsar Convention as the basis of an integrated and multi-node field research and global observation program for detecting and even preempting ecological change in wetlands.     

Multiple genetic pathways for seed shattering in the grasses

Shattering is an essential seed dispersal mechanism in wild species. It is believed that independent mutations at orthologous loci led to convergent domestication of cereal crops. To investigate genetic relationships of Triticeae shattering genes with those of other grasses, we mapped spike-, barrel- (B-type), and wedge-type (W-type) spikelet disarticulation genes in wheat and its wild relatives. The Br1 gene for W-type disarticulation was mapped to a region delimited by Xpsr598 and Xpsr1196 on the short arm of chromosomes 3A in Triticum timopheevii and 3S in Aegilops speltoides. The spike- and W-type disarticulation genes are allelic at Br1 in Ae. speltoides. The B-type disarticulation gene, designated as Br2, was mapped to an interval of 4.4 cM between Xmwg2013 and Xpsr170 on the long arm of chromosome 3D in Aegilops tauschii, the D-genome donor of common wheat. Therefore, B- and W-type disarticulations are governed by two different orthologous loci on group-3 chromosomes. Based on map position, orthologs of Br1 and Br2 were not detected in barley, maize, rice, and sorghum, indicating multiple genetic pathways for shattering in grasses. The implications of the mapping results are discussed with regard to the evolution of polyploid wheat and domestication of cereals.Supplementary material is available in the online version of this article at     

Dominance of native grasses leads to community convergence in wetland restoration

Wetland restoration is a pressing conservation priority, but there are few replicated field studies that provide a scientific foundation for these activities. We conducted a 3-year, replicated field experiment to examine the effectiveness of initial site preparation techniques (combinations of solarization, herbicide, tilling, and thermal weed control) in restoring native plant biodiversity to an agricultural field in a former wetland prairie in Oregon, USA. Post-treatment, plots were sown with a typical restoration mix of native graminoids and forbs. Treatments were compared to three high-quality managed reference wetlands and the adjacent agricultural field. Site preparation treatments varied in their effectiveness in suppressing extant vegetation and eliminating the residual seed bank. After 1 year, the solarization and fall herbicide application treatments were the most effective at reducing exotic cover. However, after 3 years, plant community composition converged in all treatments due to a loss of annual species and increasing dominance of native perennial bunchgrasses. Plant community composition became more similar to the reference wetlands each year, but diversity and richness diverged, apparently due to a trade-off between the cover of the dominant native bunchgrasses and diversity. Successional theory offers insights into how priority effects and competitive inhibition may influence community trajectories, and offers a useful model for restoring plant communities with high native diversity and dominance. Finding ways to mitigate the tradeoff between native plant cover and diversity by actively managing successional trajectories is an important challenge in wetland restoration that deserves further investigation.     

Pollen limitation and seed set associated with year-to-year variation in flowering of Gmelina arborea in a natural tropical forest

The year-to-year variations in flowering, pollen and fruit production in ten Gmelina arborea individuals in a natural forest were observed over a period of five years, 2009?2013, and pollinator visits were observed closely over two years, 2011 and 2012. A pollen supplementation experiment was also undertaken in two years (2011 and 2012) of contrasting flowering levels. Considerable year-to-year variations were observed in flower, pollen and fruit production. The observed variation represented a normal sequence of good-year and poor-year cycling which is most parsimoniously attributed to resource allocation. The average level of pollen production per individual tree in good flowering years oscillated between 6.6 and 9.62 × 10⁸, which in the poor flowering years was between 1.92 and 3.07 × 10⁸. The magnitude of pollen limitation across years was highly significant (p < 0.0001). Fruit set through supplemental pollination was 73% and 96% greater than that of the open-pollinated branches in the year 2011 and 2012, respectively. Pollen limitation, irrespective of the arrival of outcross pollen, can reduce annual seed set in predominantly bee-pollinated trees. However, various other factors may cause flower abortion in addition to pollen limitation. The results of this study would be very valuable to the seed orchard managers and the silviculturists to manage the seed production areas (SPAs) of G. arborea. The study ultimately recommends supplemental pollination in seed orchards of G. arborea for better genetic gain and good seed yield.     

Pollination, breeding system and seed abortion in some African acacias

TYBIRK, K., 1993. Pollination, breeding system and seed abortion in some African acacias . Studies of the flower visitors and pollination ecology of Acacia albida Del., A. nilotica (L.) Willd. ex Del. A. tortilis (Forssk.) Hayne and A. Senegal (L.) Willd. in Senegal and Kenya showed a high diversity of floral foragers. One hundred and eighteen taxa of insects mainly from Hymenoptera, Lepidoptera, Coleoptera and Diptera were collected. The most important pollen vectors were bees from the families Megachilidae and Halictidae, and wasps from the families Scoliidae and Eumenidae. Beetles, flies and butterflies were secondary pollen vectors. Diversity and frequency of flower visitors of species with floral nectar ( A. Senegal and A. albida ) were not clearly different from species without floral nectar ( A. tortilis, A. nilotica ). Acacia tortilis was almost exclusively outcrossed (Index of Self Incompatibility = 0.2), with 5.5% of the inflorescences (0.13% of the flowers) developing fruits. The number of seeds per pod was positively correlated with dispersal type. Wind dispersed species had fewer seeds per pod than animal dispersed species. About 5% of the seeds were aborted in A. tortilis and A. nilotica , while 22–48% of the seeds were aborted in A. Senegal, A. ataxacantha DC. and A. polyacantha Willd. Most abortions in the latter three species occur in the proximal end of the pods, indicating selective seed abortion.     

Differences in transpiration rates between tropical and temperate grasses under controlled conditions

Summary Two experiments were conducted to determine the transpiration rates of tropical and temperate grasses under a range of environmental conditions. In dense populations, three temperate grasses lost considerably more water per unit leaf area than did four tropical grasses, though tropical grasses tended to produce more dry matter per unit leaf area. The efficiency of production in relation to water use was thus greater in tropical than in temperate grasses. Wheat, a temperate grass, lost water at an average rate 2.25 times that of sorghum, a tropical grass, on a unit leaf area basis when single leaves were exposed to temperatures from 17 to 32° and light intensities from 1.7 to 4.4×10⁴ ergs cm^-2 sec^-1 at 0.55 (1,100 to 2,800 ft.-c.). The measurement of transpiration and leaf temperature indicated that latent heat loss was much more important in wheat, and sensible heat loss was more important in sorghum as means of dissipating excess energy absorbed. These findings were attributed to the greater resistance to gas diffusion offered by sorghum than by wheat stomata in each environment.     

Pollination and fruit‐set of Grevillea robusta in western Kenya

The floral visitors of silky oak, Grevillea robusta A. Cunn. ex R.Br., their foraging behaviour and their effects on fruit‐set were studied at Malava, western Kenya. Grevillea robusta is a popular tree for farm plantings in the eastern and central African highlands. Yield of seed has been disappointingly low in some areas and a lack of appropriate pollinators has been suggested as a possible cause. Investigations involved the monitoring of visitors on active inflorescences, assessment of the rewards available to potential pollinators, and exclusion experiments to establish the effects of various visitors on fruit‐set. The flowers are visited mainly by birds and insects. The likely pollinators of G. robusta are sunbirds (Nectarinia amethystina, N. cyanolaema, N. olivacea, N. superba and N. venusta) and white‐eyes (Zosterops kikuyuensis and Z. senegalensis). Very little aggressive behaviour between birds was recorded. No nocturnal pollinators were observed. Nectar was the major floral reward for pollinators, but is likely depleted by ants and honey bees, the foraging behaviour of which confirmed them to be nectar‐robbers. These insects hardly ever touched stigmas during their visits. Eighty‐nine per cent of bird visits were in the morning (07.00–10.00 hours) when nectar volume was highest. Inflorescences bagged to exclude birds set no fruits, and unmanipulated flowers and flowers bagged with self‐pollen set no fruits, indicating a self‐incompatibility mechanism. Control cross‐pollinated flowers displayed greatly increased fruit‐set (25.1%) compared with natural open‐pollination (0.9%). All these findings confirm the importance of cross‐pollen transfer to flowers and the necessity of pollinators for fruit‐set. Effective seed production requires activity of pollinators for self‐pollen removal and cross‐pollen deposition. Seed production stands for G. robusta should be established where flowering is prolific and bird pollinators are abundant.