Structure and function in asclepiad pollination

Aspects of floral biology in relation to floral structures are investigated in seven species ofAsclepiadaceae. Pollinaria removal and insertion by the insect pollinator is aided by various structural devices such as hairs or corona elements guiding the proboscis towards the entrance of the anther slit. The nectary is situated directly below this opening, thus increasing the probability of the proboscis being introduced into the anther slit during retraction. The anther slit itself often displays structural differentiation in adaptation to the guiding of the proboscis and the insertion of a pollinium in cross pollination. These mechanisms are highly elaborated so that they work as well in closed flower types, in which the pollinator does not enter the flower directly, but only introduces its proboscis (Sisyranthus, Microloma). A special feature of pollinaria removal is the variety of ways of fixing the corpusculum against the force of the proboscis when it is drawn into the cleft. — The stigma of all species exceptSecamone is located on the lower surface of the stigma head where it is appressed to the filament tube. In some cases a direct contact of the inserted pollinium with the stigmatic surface does not seem to be necessary for pollen tube germination, probably moistening by nectar suffices in these flowers. WithTylophora spec., a case of in situ pollen tube germination has been studied. — Concerning the path of the pollen tubes, two different strategies are followed.Secamone andTylophora develop a structurally defined compitum, while the other five species have strictly divided stylar canals. In these higher evolved species, the pollen tubes of one inserted pollinium are all directed into one ovary only, thus enabling the plant to allocate its resources to one carpel only in the frequent case of fertilization by one pollinium.

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Zusammenfassung An sieben Asclepiadaceen-Arten wird der Zusammenhang von Blütenstrukturen mit dem Bestäubungsprozeß untersucht. Die Analyse erfaßt die Bandbreite von einfacher organisierten (Secamone) bis zu hochdifferenziertenAsclepiadaceae, wobei neben dem offenen Blütentyp auch zwei Formen von geschlossenen Blüten (Sisyranthus, Microloma) untersucht werden. Die Blüten weisen unterschiedlich stark elaborierte Strukturen auf, die als Führungshilfen für den Insektenrüssel dienen. Neben der eigentlichen Leitschiene, die teilweise einen hohen Grad an innerer Differenzierung besitzt, sind vor allem Haare und Coronastrukturen an der primären Führung des Bestäuberrüssels beteiligt. Die interstaminale Lage der Nektarien sowie ihr Hinaufreichen bis zur Leitschiene sichert ebenfalls das Zurückziehen des Rüssels durch die Leitschiene. Die Einführung des Rüssels in den Spalt des Klemmkörpers erfordert einen relativ festen Sitz des Corpusculums, der in den hier untersuchten Arten auf drei verschiedenen Wegen erreicht wird: Anheftung an den Antherenrändern, basales Ankleben am Griffelkopf und Fixierung durch einen distalen Vorsprung des Griffelkopfes. MitTylophora spec. wird ein Fall vorgestellt, bei dem Fremdbestäubung wie auch Auskeimen der Pollenschläuche in situ und damit vermutlich Autogamie zugleich vorkommen. — Die übertragenen Pollinien werden gewöhnlich in die Narbenkammer eingezogen, dort keimen die Pollenschläuche aus und dringen an der Unterseite des Narbenkopfes in diesen ein. BeiSecamone erfolgt der Eintritt der Pollenschläuche an der Flanke des Griffelkopfes, beiMicroloma ist eine exakte Positionierung des Polliniums unnötig, auch vor der Leitschiene abgelegte Pollinien keimen und senden ihre Pollenschläuche durch die innere Leitschiene zum Griffelkopf. BeiSecamone undTylophora ist ein gemeinsamer Anfang der Griffelkanäle (Compitum) im Griffelkopf vorhanden, die Pollenschläuche werden hier auf beide Ovarien verteilt, auch wenn sie nur von einem einzigen eingeführten Pollinium stammen. Bei den übrigen fünf Arten sind beide Griffelkanäle isoliert voneinander, eine direkte Zuweisung der Pollenschläuche eines Polliniums zu nur einem Ovar ist hier die Regel. Damit verfolgen diese höher evolviertenAsclepiadaceae eher die Strategie der resource allocation, indem sie bei der vorherrschenden Belegung mit nur einem Pollinium durch die strukturell gelenkte Zuweisung der Pollenschläuche auch nur ein Fruchtblatt zur Reife bringen müssen.

     

Species composition and ecologo-phytocenotic spectra of vascular plants, mosses, and lichens on gold-mining sites in the taiga zone of the lower Amur River basin, the Russian Far East

Two-hundred-three species of vascular plants, 50 species of bryophyte and 41 species of lichens have been found on dredging and hydraulic gold-mining sites in the taiga zone of the lower Amur River basin. The main features of the anthropogenic relief and vegetation have been described. Dredging and hydraulic dumps are supposed to have the biggest similarity in the species composition and set of ecologo-phytocenotic elements, while the biggest difference is observed between dredging and hydraulic dumps on the one hand and pockets and settling pits, on the other. According to the data obtained in different regions, the species diversity of vascular plants growing on dredging and hydraulic gold-mining sites is quite rich.     

Integrating floral scent, pollination ecology and population genetics

1 . Floral scent is a key factor in the attraction of pollinators. Despite this, the role of floral scent in angiosperm speciation and evolution remains poorly understood. Modern population genetic approaches when combined with pollination ecology can open new opportunities for studying the evolutionary role of floral scent.
2 . A framework of six hypotheses for the application of population genetic tools to questions about the evolutionary role of floral scent is presented. When floral volatile chemistry is linked to pollinator attraction we can analyse questions such as: Does floral volatile composition reflect plant species boundaries? Can floral scent facilitate or suppress hybridization between taxa? Can the attraction of different pollinators influence plant mating systems and pollen-mediated gene flow? How is population genetic structure indirectly influenced by floral scent variation?
3 . The application of molecular tools in sexually deceptive orchids has confirmed that volatile composition reflects species boundaries, revealed the role of shared floral odour in enabling hybridization, confirmed that the sexual attraction mediated by floral odour has implications for pollen flow and population genetic structure and provided examples of pollinator-mediated selection on floral scent variation. Interdisciplinary studies to explore links between floral volatile variation, ecology and population genetics are rare in other plant groups.
4 . Ideal study systems for future floral scent research that incorporate population genetics will include closely related taxa that are morphologically similar, sympatric and co-flowering as well as groups that display wide variation in pollination mechanisms and floral volatiles.     

Role of lakes for organic carbon cycling in the boreal zone

We calculated the carbon loss (mineralization plus sedimentation) and net CO₂ escape to the atmosphere for 79 536 lakes and total running water in 21 major Scandinavian catchments (size range 437–48 263 km²). Between 30% and 80% of the total organic carbon that entered the freshwater ecosystems was lost in lakes. Mineralization in lakes and subsequent CO₂ emission to the atmosphere was by far the most important carbon loss process. The withdrawal capacity of lakes on the catchment scale was closely correlated to the mean residence time of surface water in the catchment, and to some extent to the annual mean temperature represented by latitude. This result implies that variation of the hydrology can be a more important determinant of CO₂ emission from lakes than temperature fluctuations. Mineralization of terrestrially derived organic carbon in lakes is an important regulator of organic carbon export to the sea and may affect the net exchange of CO₂ between the atmosphere and the boreal landscape.     

Response of littoral vegetation on climate warming in the boreal zone; an experimental simulation

The impact of climate warming on the littoral zone of a boreal lake ecosystem was studied experimentally for three growing seasons in two artificial ponds (10×27 m) and in replicated chamber experiments. One pond was enclosed in a plastic greenhouse and another untreated pond served as a reference system. During the growing seasons temperature in the greenhouse was maintained at levels 2–3 °C higher than ambient with a computer-controlled ventilation system. One growing season prior to initiation of the experiment, a vegetated littoral zone with equal densities of water horsetail (Equisetum fluviatile) was established in both ponds. Although changes occurred in the species dominance (E. fluviatile - Alisma plantago-aquatica - Sparganium erectum spp. microcarpum - Elodea canadensis) within the three years of the study, the emergent macrophytes emerged earlier and grew better in the warmer conditions of the greenhouse pond compared with those in the reference pond. The difference in above-ground biomass throughout the growing seasons was >2 fold and after three experimental growing seasons the difference in below-ground biomass of macrophytes was 2.5-fold between the ponds. In replicated chamber experiments the biomass growth of E. fluviatile was also significantly higher in a 2–3 °C higher temperature than under ambient conditions. An ecosystem-scale induced change, characterized by a heavy growth of filamentous algae (mainly chlorophytes) was evident in the vegetated littoral zone of the greenhouse pond. A hypothesis that macrophyte rhizomes function as `phosphorus pumps' from the sediment and thus accelerate eutrophication in a warmer climate should be further studied.     

Effects of spring flood and water level draw-down on methane dynamics in the littoral zone of boreal lakes

1. The annual dynamics of methane (CH₄) in a temporarily flooded meadow, mire bank, lacustrine sedge fen, temporarily and continuously inundated sedge ( Carex sp.) and reed ( Phragmites australis ) marshes were studied from June to November in the humic mesoeutrophic Lake Mekrijärvi and in eutrophicated parts of the mesotrophic Lake Heposelkä in the southern part of East Finland. The effects of water level and temperature on littoral CH₄ fluxes were determined. Vegetation zonation along the moisture gradient, and associated CH₄ fluxes, were evaluated.
2. The CH₄ flux increased along the moisture gradient from –0.2 to 14.2 mg CH₄ m^–2 h^–1, and was highest in the permanently inundated marshes. The duration of anoxia in the sediment caused differences in the CH₄ flux. Estimated emissions for the period 1 June – 30 September in continuously inundated sparse reed and sedge marshes, drying sedge marsh, and lacustrine sedge fen were 13, 11 and 6 g CH₄ m^–2, respectively.
3. In continuously inundated vegetation, the fluxes were highest in late July/early August. The seasonal CH₄ flux pattern suggested that the fluxes were regulated by the supply of organic matter during the course of the summer and the water level. In the temporarily flooded zone, the seasonal CH₄ flux dynamics was greatly affected by changes in the lake water level, the fluxes being highest during the spring flood in early June.     

Biotic versus abiotic pollination in Oeceoclades maculata (Lindl.) Lindl. (Orchidaceae)

Geographical variation in the reproductive biology of widespread species often occurs at their distributional boundaries. We sought to determine whether such variation has occurred in an invasive orchid, Oeceoclades maculata, across its naturalized range. We compared its reproductive biology in a Brazilian population with that published for a population on the Caribbean island of Puerto Rico. In the state of São Paulo, O. maculata flowers between December and February, at the height of the rainy season. Similar fruit sets were observed in manual self (76%) and cross (70.4%) pollination treatments. The fruit set of plants protected from both pollinators and rainfall was 6.1%, whereas plants exposed only to rainfall had a fruit set of 41.4%, slightly less than the controls (48.3%). Like the Puerto Rico population, reproduction is primarily through rain‐assisted autogamy, but unlike observations made on the island, outcrossing can eventually occur. We observed two butterfly species (Heliconius ethilla narcaea and Heliconius erato phyllis) pollinating O. maculata. Secretory epidermal cells and trichomes of the spur lumen produced 0.7 µL of 25% (sucrose equivalents) nectar per flower each morning, which was stored in a dilated basal portion of the spur and reabsorbed by the afternoon. Thus, geographical variation in reproductive biology exists across the broad invasive range of O. maculata.     

Climate and vegetation controls on boreal zone energy exchange

The boreal forest, one of the world's larger biomes, is distinct from other biomes because it experiences a short growing season and extremely cold winter temperatures. Despite its size and impact on the earth's climate system, measurements of mass and energy exchange have been rare until the past five years. This paper overviews results of recent and comprehensive field studies conducted in Canada, Siberia and Scandinavia on energy exchanges between boreal forests and the atmosphere. How the boreal biosphere and atmosphere interact to affect the interception of solar energy and how solar energy is used to evaporate water and heat the air and soil is examined in detail. Specifically, we analyse the magnitudes, temporal and spatial patterns and controls of solar energy, moisture and sensible heat fluxes across the land–atmosphere interface. We interpret and synthesize field data with the aid of a soil–vegetation–atmosphere transfer model, which considers the coupling of the energy and carbon fluxes and nutrient status. Low precipitation and low temperatures limit growth of many boreal forests. These factors restrict photosynthetic capacity and lower root hydraulic conductivity and stomatal conductance of the inhabitant forests. In such circumstances, these factors interact to form a canopy that has a low leaf area index and exerts a significant resistance to evaporation. Conifer forests, growing on upland soils, for example, evaporate at rates between 25 and 75% of equilibrium evaporation and lose less than 2.5 mm day^?1 of water. The open nature of many boreal conifer forest stands causes a disproportionate amount of energy exchange to occur at the soil surface. The climatic and physiological factors that yield relatively low rates of evaporation over conifer stands also cause high rates of sensible heat exchange and the diurnal development of deep planetary boundary layers. In contrast, evaporation from broad‐leaved aspen stands and fen/wetlands approach equilibrium evaporation rates and lose up to 6 mm day^?1.     

植物繁殖生态学的若干重要问题

发现模式,并根据过程解释模式,是科学研究的主要目标.在植物繁殖生态学领域内,人们已经总结了一些重要的模式,包括普遍的雌雄同体、分离的花果期、较低的结实率、大量的同时开花、胁迫生境更易出现的异交、生活史性状与交配系统的相互作用、花序内的花分化,以及花内的雄蕊分化(异型雄蕊)等.对这些模式如何从机理上予以解释,还需要植物繁殖生态学家的艰苦努力.本文着重强调了植物许多重要繁殖特征都可以通过植株、花序以及花各层次上的资源分配而得到很好的解释,尤其是可以在很大程度上协调其他不同侧重点研究得到的矛盾结论.在今后逐渐强调多因素、多尺度对花部特征与传粉系统的适应与进化的研究中,从直接决定花部特征与交配系统的资源分配出发,将是促进人们准确理解与预测花部特征适应与进化的一个重要研究方向.     

Reproductive ecology of dioecious Siparuna (Monimiaceae) in Ecuador–a case of gall midge pollination

The reproductive ecology of the dioecious genus Siparuna , belonging to the primitive family Monimiaceae, was investigated in Ecuador. Thirteen species were studied, and species growing at high altitudes (above 500 m) were compared with species growing at lower altitudes. The genus has specialized flowers pollinated by Cecidomyiidae (Asynapta sp. and other genera) that lay eggs chiefly in male flowers since these are more easily accessible than female flowers due to their morphology. Several traits ensure that pollen transfer is accomplished: 1. long flowering of individuals and single flowers; 2. specific floral morphology (unisexuality, landing platform, restricted pore) allowing the flowers to be visited by Cecidomyiidae seeking egg-laying sites, and 3. a low number of ovules that have to be pollinated in each flower. The low number of ovules per flower results in a high pollen-ovule (P/O) ratio even though pollen production in male flowers is only moderate.
High altitude species had larger flowers, higher pollen-ovule ratios, fewer ovules per female flower, and longer periods of anthesis. Gender differences were seen in the longer functionality of female flowers; the larger numbers of male flowers produced; and the average diameter at breast height (DBH) biased for one of the two sexes depending on the species investigated, with no clear trend in the size distributions of individuals of both sexes. Sex ratios departed significantly from unity both to male and female bias but usually were in favour of males. There were no general differences in the ratio of functional male to female flowers at the time of observation nor in the sizes of male and female flowers. The evolution of unisexual flowers in Siparuna can be explained as a result of the differential predation by larvae: unimportant in male flowers, destructive if occurring in female flowers.     

Reproductive biology and pollination of Govenia utriculata: A syrphid fly orchid pollinated through a pollen-deceptive mechanism

The reproductive biology and pollination mechanisms of Govenia utriculata (Sw.) Lindl. were studied in a mesophytic semideciduous forest at Serra do Japi, south-eastern Brazil. The floral visitors and pollination mechanisms were recorded, and experimental pollinations were carried out to determine the breeding system of this species. Populations of G. utriculata growing at Serra do Japi are exclusively visited and pollinated by two species of hoverflies in the genus Salpingogaster (Diptera: Syrphidae) that are attracted by deceit to the flowers of this orchid species. The lip apex and the column base present small brownish and yellow to orange spots that mimic pollen clusters. Govenia utriculata is self-compatible, but pollinator dependent. Natural fruit set was low (10%), but similar to that of other non-obligatorily autogamous sympatric orchid species that occur at Serra do Japi and of other fly-pollinated orchid species pollinated through deceptive mechanisms.     

The pollination ecology of an assemblage of grassland asclepiads in South Africa

The KwaZulu-Natal region of South Africa hosts a large diversity of asclepiads (Apocynaceae: Asclepiadoideae), many of which are endemic to the area. The asclepiads are of particular interest because of their characteristically highly evolved floral morphology. During 3 months of fieldwork (November 2000 to January 2001) the flower visitors and pollinators to an assemblage of nine asclepiads at an upland grassland site were studied. These observations were augmented by laboratory studies of flower morphology (including scanning electron microscopy) and flower colour (using a spectrometer). Two of the specialized pollination systems that were documented are new to the asclepiads: fruit chafer pollination and pompilid wasp pollination. The latter is almost unique in the angiosperms. Taxa possessing these specific pollination systems cluster together in multidimensional phenotype space, suggesting that there has been convergent evolution in response to similar selection to attract identical pollinators. Pollination niche breadth varied from the very specialized species, with only one pollinator, to the more generalized, with up to ten pollinators. Pollinator sharing by the specialized taxa does not appear to have resulted in niche differentiation in terms of the temporal or spatial dimensions, or with regards to placement of pollinaria. Nestedness analysis of the data set showed that there was predictability and structure to the pattern of plant-pollinator interactions, with generalist insects visiting specialized plants and vice versa. The research has shown that there is still much to be learned about plant-pollinator interactions in areas of high plant diversity such as South Africa.     

Hydrophilous pollination and breeding system evolution in seagrasses: a phylogenetic approach to the evolutionary ecology of the Cymodoceaceae

COX, P. A. & HUMPHRIES, C. J. 1992. Hydrophilous pollination and breeding system evolution in seagrasses: a phylogenetic approach to the evolutionary ecology of the Cymodoceaceae. A phylogenetic analysis of seagrasses of the Cymodoceaceae shows the Cymodoceaceae to be monophyletic and Posidoniaceae to be their sister group. Information on the pollination ecologies and breeding systems of the various genera of the Cymodoceaceae were plotted onto the consensus tree obtained for the group. From this analysis, it is suggested that the clade composed of the Zosteraceae, Posidoniaceae and Cymodoceaceae evolved from a freshwater hydrophilous ancestor that developed submarine pollination and filiform pollen in association with invasion of the marine environment. Dioecism and surface pollination appear to have evolved in the progenitor of the Cymodoceaceae, and hence the seagrasses of the Cymodoceaceae are dioecious due to common descent rather than to convergent evolutionary processes in extant genera.     

Spring phenology in boreal Eurasia over a nearly century time scale

It has been widely reported that tree leaves have tended to appear earlier in many regions of the northern hemisphere in the last few decades, reflecting climate warming. Satellite observations revealed an 8-day advance in leaf appearance date between 1982 and 1991 in northern latitudes. In situ observations show that leaf appearance dates in Europe have advanced by an average of 6.3 days from 1959 to 1996. Modelling of leaf appearance on the basis of temperature also shows a marked advance in temperate and boreal regions from 1955 to 2002. However, before 1955, reported studies of phenological variations are restricted to local scale. Modelling, ground observations and satellite observations are here combined to analyse phenological variations in Eurasian taiga over nearly a century. The trend observed by remote sensing consists mainly in a shift at the end of the 1980s, reflecting a shift in winter and spring temperature. In western boreal Eurasia, a trend to earlier leaf appearance is evident since the mid-1930s, although it is discontinuous. In contrast, the strong advance in leaf appearance detected over Central Siberia using satellite data in 1982–1991 is strengthened by late springs in 1983–1984; moreover, in this region the green-up timing has displayed successive trends with opposite signs since 1920. Thus, such strong trend is not unusual if considered locally. However, the recent advance is unique in simultaneously affecting most of the Eurasian taiga, the leaf appearance dates after 1990 being the earliest in nearly a century in most of the area.     

Direct costs of forest reproduction,bee-cycling and the efficiency of pollination modes

The bee guild represents direct primary costs of angiosperm reproduction. Tropical flower visitors take an amount comparable to herbivores, exceeding 3% of net primary production energy. Therefore herbivory and aboveground net primary production have been underestimated. Comparing pollinators to other herbivores, harvest in mature forest by tropical bees is greater than leafcutter ants, game animals, frugivores, vertebrate folivores, insect defoliators excluding ants, flower-feeding birds and bats, but not soil organisms. The ratio of total aboveground net primary production to investment in pollen, nectar and resin used by pollinators suggests wind pollination is several times more efficient in temperate forests than is animal pollination in neotropical moist forest. Animal pollination may be favoured by habitat mosaics and an unpredictable or sparse dispersion of conspecifics — consequences of fluctuating abiotic and biotic environments. Natural selection evidently favours diminished direct reproductive costs in forests, for example by wind pollination, regardless of latitude and disturbance regime. An example is “wind pollination by proxy” of dominant trees in seasonal southeast Asian forests. They flower only occasionally and their pollen is dispersed by tiny winged insects that are primarily carried by the wind — rather than the nectar-hungry bees, bats, birds and moths used by most tropical flora. Increasing evapotranspiration is associated with greater net primary production; I show its correlation with species richness of social tropical bees across the isthmus of Panama, which may indicate increasing forest reproductive effort devoted to flowering, and its monopolization by unspecialized flower visitors in wetter and less seasonal lowland forests.     

Relationships between DOC concentration and epilithon stable isotopes in boreal lakes

1. Littoral biota in boreal lakes are known to assimilate epilithon. Being able to predict the stable isotopic composition of these alga will help to identify those systems in which δ¹³C and δ¹⁵N analysis can be used in foodweb investigations of allochthony and biomagnification.
2. In a survey of 15 boreal lakes, the concentration of dissolved organic carbon (DOC) explained 76% of the variation in epilithon δ¹³C, and 86% of the variation in epilithon δ¹⁵N.
3. Because both δ¹³C and δ¹⁵N values were depressed and similar to terrestrial values in humic (high DOC) lakes, it will be more difficult to successfully employ stable isotopic techniques for estimating allochthony in such systems. Lower δ¹⁵N values in humic lakes also indicate that trophic positions estimated by stable isotopes are not directly comparable to those of similar biota inhabiting clearwater lakes, unless autochthonous baseline corrections are made.     

Wind of change: new insights on the ecology and evolution of pollination and mating in wind-pollinated plants

Background

The rich literature that characterizes the field of pollination biology has focused largely on animal-pollinated plants. At least 10 % of angiosperms are wind pollinated, and this mode of pollination has evolved on multiple occasions among unrelated lineages, and hence this discrepancy in research interest is surprising. Here, the evolution and functional ecology of pollination and mating in wind-pollinated plants are discussed, a theoretical framework for modelling the selection of wind pollination is outlined, and pollen capture and the occurrence of pollen limitation in diverse wind-pollinated herbs are investigated experimentally.

Scope and Conclusions

Wind pollination may commonly evolve to provide reproductive assurance when pollinators are scarce. Evidence is presented that pollen limitation in wind-pollinated plants may not be as common as it is in animal-pollinated species. The studies of pollen capture in wind-pollinated herbs demonstrate that pollen transfer efficiency is not substantially lower than in animal-pollinated plants as is often assumed. These findings challenge the explanation that the evolution of few ovules in wind-pollinated flowers is associated with low pollen loads. Floral and inflorescence architecture is crucial to pollination and mating because of the aerodynamics of wind pollination. Evidence is provided for the importance of plant height, floral position, and stamen and stigma characteristics in promoting effective pollen dispersal and capture. Finally, it is proposed that geitonogamous selfing may alleviate pollen limitation in many wind-pollinated plants with unisexual flowers.Key words: Wind pollination, reproductive assurance, pollen limitation, geitonogamy, sex allocation, inflorescence architecture, mating systems     

Beaver population fluctuations and tropospheric methane emissions in boreal wetlands

Measurements of net methane flux were made during the 1988 ice-free season (May–October) at a beaver-meadow complex in northern Minnesota, USA. The site included upland boreal forest, sedge meadow, submerged aquatic plants, and the open water of a beaver pond. Annual fluxes were 8–11 g C/m² in the permanently wetted zones and 0.2–0.4 g C/m² at the occasionally inundated meadow and forest sites. These data, when coupled with long-term (46 yr) data on beaver (Castor canadensis) population size and habitat alteration, suggest that about 1% of the recent rise in atmospheric methane may be attributable to pond creation by beaver in North America.