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1.
A technique is presented that is capable of predicting the motion of airborne pollen grains and the probability of pollen capture by wind-pollinated plants. Equations for the motion of rigid-walled particles (= pollen grains, spores, or Sephadex beads) in a supporting, compressible fluid (= air) are derived from the first principles of fluid dynamics. These equations are incorporated into a computer program (MODEL) which can be used with a desktop computer. The operation of MODEL requires empirical data on the pattern of airflow or the motion of a pollen species around the surfaces of the taxonomically relevant ovulate plant organ. With this information, MODEL can predict the behavior of any pollen species for which physical properties (size and density) are specified or empirically known. The significance of this procedure lies in the quantification of physical phenomena that influence the mechanics and fluid dynamics of pollen capture in wind pollination. The technique is illustrated and tested by its application to two grass species (Setaria geniculata and Agrostis hiemalis) for which velocity fields of pollen motion have been previously reported.  相似文献   

2.
Empirically determined patterns of pollen impaction on the surfaces of pine ovulate cones are correlated with regions of nonlaminar flow created by the spatial arrangement and morphology (aspect ratios) of scale-bract complexes. Results from the serial discharge of pollen, upwind of ovulate cones, indicate that ovules on ovulate cones are preferentially impacted by pollen from their own species. Analyses indicate that while aerodynamic factors dominate the entrapment of pollen by ovulate cones, other factors such as pollen impaction-rebound and rebound-reentrainment are significant. Surface characteristics in addition to the settling velocities of pollen may play important roles in determining pollination efficiency. Wind tunnel analyses of the aerodynamic effects of scale-bract arrangement and aspect ratios indicate that each complex behaves as an aerofoil, deflecting air eddies toward the micropylar ends of ovules. The ovulate cone geometry, as a whole, deflects unidirectional wind into cyclonic vortices around the cone axis, each scale-bract deflecting nonimpacted pollen along orthostichies and parastichies. The morphology of the typical conifer ovulate cone is interpreted as a structure that optimizes anemophilous reproduction.  相似文献   

3.
Comparisons are presented between the three-dimensional airflow patterns created around and by a scale model of a conifer ovulate cone and the trajectories of windborne pollen grains around Picea, Larix, and Pinus ovulate cones. Three general components of the airflow pattern around an ovulate cone model are 1) doldrum-like eddies, rotating over the adaxial surfaces of cone scales and directed toward attached ovules, 2) airflow spiralling around the cone axis along cone scale orthostichies and parastichies, and 3) a complex pattern of vortices (“umbilicus”) directed toward the leeward surface of the ovulate cone. The observed trajectories of pollen grains around cones of Picea, Larix, and Pinus conform to two of these three airflow components: 1) pollen grains are seen to roll along cone scales toward the distal scale margin and to become reentrained in airflow directed backward toward attached ovules, and 2) pollen grains passing around the cone are deflected into the “umbilicus” airflow pattern, where they either settle on or impact with cone scales (approach trajectories), or where they approach the leeward cone surface but are deflected away by airflow passing under the cone (Z-shaped trajectories). Vectoral analyses of pollen grain motion reveal a complex pattern of trajectories influenced by boundary layer conditions defined by ovulate cone geometry and ambient airflow speed. Wind tunnel studies of ovulate cones subtended by leaves and stem indicate that leaves circumscribing the cone act as a snowfence, deflecting windborne pollen toward the cone. Vectoral analyses of airflow patterns and pollen grain trajectories close to ovulate cones indicate that wind pollination in conifers is a non-stochastic aerodynamic process influenced by cone-leaf morphology and the behavior of pollen grains as windborne particles.  相似文献   

4.
Sacci of conifer pollen do not function primarily to increase the efficiency of wind pollination as is widely thought. Rather, they are bladders and cause pollen to float upwards in a liquid drop into the ovules. This observation is seemingly unsupported in the case of oriental spruce (Picea orientalis (L.) Link), which has saccate pollen. Ovulate cones are pendant at the time of pollination, which requires that pollen sink into the ovules. Pollen of oriental spruce floats at first but within 1-2 min sinks into the ovule. As sinking does not occur in saccate pollen of other Pinaceae, a variety of techniques was used to determine anatomical differences leading to this uncharacteristic tendency. Light, scanning electron, and confocal microscopy of the pollen surface yielded no significant appearing difference between pollen of oriental spruce and white spruce. However, transmission electron microscopy of freeze-fixed/freeze-substituted hydrated pollen revealed that the ektexine of oriental spruce pollen sacci is porous compared to that of white spruce. Confocal microscopy allowed examination of pollen hydration dynamics. Water enters pollen at the distal pole between sacci, and resulting rapid expansion of the tube cell forces air out of the saccate space. White spruce pollen remains buoyant because of enclosed air pockets in the saccus ektexine. Evolutionary change in pollen wall anatomy with resultant loss of saccus function is correlated with a change in ovulate strobilus orientation at pollination in oriental spruce. A suite of characters interact in the conifer pollination mechanism, and concerted change in these characters may lead to speciation.  相似文献   

5.
Wind-tunnel analyses of the behavior of airborne pollen around ovules of two Ephedra species (E. trifurca and E. nevadensis) indicate that at certain airflow speeds (0.5 m/sec and 1.0 m/sec) each species is capable of biasing pollination in favor of conspecific pollen. A computer procedure was designed to evaluate the physical basis for this aerodynamic discrimination. This procedure indicates that differences in size and density confer significantly different inertial properties to the two pollen species. Operating within the specific aerodynamic environments generated around ovules from each species, these differences are sufficient to account for the biases observed in the probability of pollination. Within natural populations, there exists significant variation in pollen size (and possibly in density). Accordingly, it is possible that, under certain ambient wind conditions, ovules from each species can select subsets of the entire airborne population of Ephedra pollen.  相似文献   

6.
In Sequoiadendron ovules are borne inside the ovulate cone, and pollination drops secreted from these ovules collect pollen. We examined: (1) the relation between ovular position and pollen capture; (2) pollen behavior when in contact with a pollination drop; and (3) ultrastructure of ovules during pollination drop secretion. During wet periods a water sheet forms on the surface of the cone due to bract shape and wettability. Pollination drops persist inside the wetted cone, and pollen capture resumes immediately after drying. Pollen landing on a pollination drop is taken inside the drop and carried into the micropyle when the drop contracts. Several notable ultrastructural features appear in the nucellus, integument, chalaza, and bract lamina during pollination-drop secretion. The abaxial surface of the lamina is covered by a membrane that may contribute to the wettable nature of the surface.  相似文献   

7.
Wind tunnel analyses of Simmondsia chinensis (Link) Schneider or “jojoba” were conducted to quantify the behavior of airborne pollen grains around individual branches and leaves and near individual carpellate flowers. Field data (wind velocity) were used to ensure a correspondence between wind tunnel and natural conditions. Based upon the visualization of individual pollen grain trajectories, it is concluded that pollen deposition on stigmatic surfaces is influenced by large-scale aerodynamic patterns, generated by foliage leaves, and small-scale airflow patterns, formed around and by floral parts and stigmas. Leaves are seen to deflect airborne pollen grains into trajectories that can intersect ambient airflow at 90° angles, showering decumbent carpellate flowers with pollen. Similarly, flowers can deflect pollen upward and downwind, toward other flowers. The extent of floral bract and sepal recurvature is shown to influence the extent of pollen deposition by determining the characteristic airflow pattern around stigmas. Available evidence concerning the relatively recent evolutionary transition to anemophily in Simmondsia is interpreted within the context of morphological adaptations and exaptations favoring wind pollination.  相似文献   

8.
This paper discusses the characteristic air mass types over the Carpathian Basin in relation to plant pollen levels over annual pollination periods. Based on the European Centre for Medium-Range Weather Forecasts dataset, daily sea-level pressure fields analysed at 00 UTC were prepared for each air mass type (cluster) in order to relate sea-level pressure patterns to pollen levels in Szeged, Hungary. The database comprises daily values of 12 meteorological parameters and daily pollen concentrations of 24 species for their pollination periods from 1997 to 2001. Characteristic air mass types were objectively defined via factor analysis and cluster analysis. According to the results, nine air mass types (clusters) were detected for pollination periods of the year corresponding to pollen levels that appear with higher concentration when irradiance is moderate while wind speed is moderate or high. This is the case when an anticyclone prevails in the region west of the Carpathian Basin and when Hungary is under the influence of zonal currents (wind speed is high). The sea level pressure systems associated with low pollen concentrations are mostly similar to those connected to higher pollen concentrations, and arise when wind speed is low or moderate. Low pollen levels occur when an anticyclone prevails in the region west of the Carpathian Basin, as well as when an anticyclone covers the region with Hungary at its centre. Hence, anticyclonic or anticyclonic ridge weather situations seem to be relevant in classifying pollen levels.  相似文献   

9.
Observation of ovulate cones at the time of pollination in the southern coniferous family Podocarpaceae demonstrates a distinctive method of pollen capture, involving an extended pollination drop. Ovules in all genera of the family are orthotropous and single within the axil of each fertile bract. In Microstrobus and Phyllocladus ovules are erect (i.e., the micropyle directed away from the cone axis) and are not associated with an ovule-supporting structure (epimatium). Pollen in these two genera must land directly on the pollination drop in the way usual for gymnosperms, as observed in Phyllocladus. In all other genera, the ovule is inverted (i.e., the micropyle is directed toward the cone axis) and supported by a specialized ovule-supporting structure (epimatium). In Saxegothaea there is no pollination drop and gametes are delivered to the ovule by pollen tube growth. Pollination drops were observed in seven of the remaining genera. In these genera the drop extends over the adjacent bract surface or cone axis and can retain pollen that has arrived prior to drop secretion (“pollen scavenging”). The pollen floats upward into the micropylar cavity. The configuration of the cone in other genera in which a pollination drop has not yet been observed directly suggests that pollen scavenging is general within the family and may increase pollination efficiency by extending pollination in space and time. Increased pollination efficiency may relate to the reduction of ovule number in each cone, often to one in many genera, a derived condition. A biological perspective suggests that animal dispersal of large seeds may be the ultimate adaptive driving force that has generated the need for greater pollination efficiency.  相似文献   

10.
The air disturbance patterns created by and around the ovules of Taxus cuspidata are quantified for various orientations to the direction of ambient airflow, and are shown to largely dictate the motion (vectoral trajectories) and mode of deposition of windborne pollen on ovule surfaces. Perpendicular orientation to the direction of airflow results in two regions characterized by high densities of adhering pollen — one on the windward surface of the ovule, resulting from direct inertial collision, and another on the leeward surface resulting from non-inertial sedimentation. Parallel and inclined orientations of the ovule to the direction of airflow produce quantitative and qualitative variations in the pattern of adhering pollen resulting from inertial and non-inertial deposition. Direct collision of windborne pollen grains with the micropylar ends of ovules occurs for all orientations to wind direction. The aerodynamics of the ovulate shoot complex of Taxus cuspidata is related to that previously described for conifer ovulate cones, cycad megastrobili, and simulated wind tunnel analyses of archaic Paleozoic ovules based on scale models. Water transport of pollen (adhering to integument and bract surfaces) to micropyles quantitatively alters the distribution of adhering pollen grains on ovule surfaces. Although there is no evidence that pollen grains of this species are osmotically ruptured, observations do not preclude the possibility that water transport of pollen may reduce the number of viable pollen grains reaching the micropyle.  相似文献   

11.
The phenology of pollen release and pollen capture by Pistacia vera was studied in the field and laboratory respectively. Inflorescences of Pistacia vera were examined in a wind tunnel to determine whether the behavior of airborne conspecific pollen around receptive flowers differed as a result of changes in the shape and size of the inflorescence. In addition, the behavior of unclumped (single) and clumped pollen grains was studied to determine differences in the probability of their capture. Wind speeds within a commercial orchard during pollen shedding averaged 0.9–2.2 m/sec and atmospheric pollen concentrations were highest between 0900–1100 hr MST. Each of three stages in inflorescence development (defined on the basis of the number of exserted stigmas) was examined under identical ambient airflow conditions with equal concentrations of airborne pollen (1,000 grains/m3). The general pattern of pollen grain motion involves direct inertial collision by windward surfaces and by sedimentation of pollen onto leeward surfaces; clumped pollen rarely sedimented onto leeward surfaces. Small changes in ambient wind speed (0.5 m/sec to 1.0 m/sec) produced significant changes in the pattern of pollen motion around inflorescences and altered the number of pollen grains captured by leeward surfaces. Thus, wind pollination in P. vera is affected both by windspeed and by the shape or size of flower clusters. Differences in the behavior of clumped and unclumped pollen result from their inertial properties and responsiveness to local changes in the direction and speed of airflow. Unclumped pollen has a higher probability of being captured by leeward surfaces. The apparent insensitivity of pollen motion to differences in inflorescence size may ensure equitable pollination during the acropetal development of flowers.  相似文献   

12.
Aerodynamic analyses showing characteristic airflow patterns and the potential for wind-mediated pollination are presented for models of Paleozoic (Carboniferous) ovules and ovulate cupules (i.e., Genomosperma kidstoni, G. latens, Salpingostoma dasu, Physostoma elegans, Eurystoma angulare, and Stamnostoma huttonense). Lobes on ovules and cupules are shown to produce localized regions of turbulent flow with a concomitant reduction in airflow velocity. Data based upon models that mimic the characteristics of windborne pollen (= pseudopollen) show that these regions of turbulent flow correspond to those in which suspended pseudopollen impact with ovule and/or cupule surfaces. These data have bearing on a sequence of ovule morphologies purported to show the evolution of the integument by the progressive reduction in length of “preintegumentary” lobes and their acropetal fusion. As the preintegumentary lobes of the models studied consolidate around the megasporangium, regions of turbulent flow and high pseudopollen impact become localized around the pollen chamber or salpinx. The general morphologic trend envisioned for the evolution of the ovule is seen to be associated with an aerodynamic streamlining and an increased potential for wind-mediated pollination. Data for hair-bearing ovules and for ovulate cupules are discussed within the context of possible selective pressures favouring streamlining.  相似文献   

13.
The transport and capture of pollen in ~20% of all angiosperm families occurs in air and water. In other words, pollination is abiotic and occurs via the fluid media, not an animal vector. Whereas some early concepts considered abiotic pollination to be largely a stochastic phenomenon, there is sufficient evidence to indicate that wind pollination (i.e. anemophily) and water pollination (i.e. hydrophily) have deterministic features and are sophisticated fluid dynamic solutions to the problem of pollen release, dispersal, and capture.An abiotic pollination syndrome is defined in which there is spatial or temporal separation of carpellate and staminate flowers, which are drab, a reduction in perianth parts, stigmas and anthers are exposed to the fluid, and typically unclumped pollen may be produced in large amounts. Separate pollination syndromes are defined for anemophilous (i.e. wind-pollinated), ephydrophilous (i.e. surface-pollinated), and hydrophilous (i.e. submarine-pollinated) plants. Distinctions are based on habitat and physical conditions for pollination, pollen size, shape, and ultrastructure, morphology and ultrastructure of stigmas, and outcrossing rates. For example, anemophilous pollen are spherical and small, ephydrophilous pollen are spherical or reniform and large, while hydrophilous pollen are filiform (i.e. filamentous) or functionally filiform. The pollination mechanisms and mechanics associated with these syndromes reveals a strong evolutionary relationship between plant morphology and fluid dynamics.  相似文献   

14.
Pollination systems and associated floral traits generally differ between core and marginal populations of a species. However, such differences are rarely examined in plants with a mixed wind‐ and bumblebee‐pollination system, and the role of wind pollination during range expansion in ambophilous plants remains unclear. We compared floral traits and the contributions of bumblebee and wind pollination in refugium and marginal populations of the ambophilous plant Aconitum gymnandrum. We found that most floral traits differed between the two populations, and those traits associated with the shift to wind pollination were pronounced in the marginal population. Bumblebee visitation rates varied significantly, but were generally low in the marginal population. Wind pollination occurred in both populations, and the efficiency was lower than that of bumblebee pollination. Two types of pollen grains, namely round and fusiform pollen, were transported to a stigma by bumblebees and wind, but fusiform pollen contributed to wind pollination to a larger degree, especially in the marginal population. Our results suggest that wind pollination was enhanced by pollen dimorphism in the marginal population of A. gymnandrum, and wind pollination may provide reproductive assurance when bumblebee activity is unpredictable during range expansion, indicating that ambophily is stable in this species and shift in pollination system could be common when plants colonize new habitats.  相似文献   

15.
Computer simulations are used to predict the behavior of pollen grains with different physical properties within the acceleration field created around the ovules of the gymnosperm Ephedra trifurca. A modelling procedure is given that (1) calculates the number of pollen grains captured by an ovule's pollination-droplet and (2) gives a correlation between pollination efficiency and the physical properties (= mass, size) of different types of pollen. Based on this procedure, the number of Ephedra pollen grains captured by micropyles can be less than the number captured from other species. However, the mass and size of Ephedra pollen grains appear to coincide with those predicted to yield a local maximum of pollination efficiency, i.e. slightly larger or smaller values of either mass or size would decrease the probability of capture. In addition, the properties of Ephedra pollen grains operate synergistically in the aerodynamic environment around ovules and are focused to collide with pollination-droplets. By analogy, the properties of Ephedra pollen coincide with those predicted for a localized adaptive peak. The physical properties of pollen grain types other than E. trifurca that can maximize pollen capture are not generally represented in the aerobiology of Ephedra during the pollination season. Therefore, the phenology of pollen release, community taxonomic-composition, and the physics of particle capture play collectively important roles in the reproductive success of Ephedra trifurca.  相似文献   

16.
Evolutionary transitions from animal to wind pollination have occurred repeatedly during the history of the angiosperms, but the selective mechanisms remain elusive. Here, we propose that knowledge of pollen release biomechanics is critical for understanding the ecological and evolutionary processes underpinning this shift in pollination mode. Pollen release is the critical first stage of wind pollination (anemophily) and stamen properties are therefore likely to be under strong selection early in the transition. We describe current understanding of pollen release biomechanics to provide insights on the phenotypic and ecological drivers of wind pollination. Pollen release occurs when detachment forces dominate resistive forces retaining pollen within anthers. Detachment forces can be active or passive depending on whether they require energy input from the environment. Passive release is more widespread in anemophilous species and involves processes driven by steady or unsteady aerodynamic forces or turbulence-induced vibrations that shake pollen from anthers. We review empirical and theoretical studies suggesting that stamen vibration is likely to be a key mechanism of pollen release. The vibration response is governed by morphological and biomechanical properties of stamens, which may undergo divergent selection in the presence or absence of pollinators. Resistive forces have rarely been investigated for pollen within anthers, but are probably sensitive to environmental conditions and depend on flower age, varying systematically between animal- and wind-pollinated species. Animal and wind pollination are traditionally viewed as dichotomous alternatives because they are usually associated with strikingly different pollination syndromes. But this perspective has diverted attention from subtler, continuously varying traits which mediate the fluid dynamic process of pollen release. Reinterpreting the flower as a biomechanical entity that responds to fluctuating environmental forces may provide a promising way forward. We conclude by identifying several profitable areas for future research to obtain deeper insight into the evolution of wind pollination.  相似文献   

17.
应用花粉通量测定新技术,于1992~1994年在小麦(Triticum aestivum L.)、玉米(Zea mays L.)传粉期间,应用库尔式收集仪收集和测定空气中的花粉,记录气象资料,进行统计分析,确定小麦、玉米产量与它们最大传粉期空气中花粉浓度的相对应数据,根据这些数据,试图建立禹城实验区小麦和玉米产量预报图式。  相似文献   

18.
Over 50 genera of bees release pollen from flower anthers using thoracic vibrations,a phenomenon known as buzz-pollination.The efficiency of this process is directly affected by the mechanical properties of the buzzes,namely the duration,amplitude,and frequency.Nonetheless,although the effects of the former two properties are well described,the role of buzz frequency on pollen release remains unclear.Furthermore,nearly all of the existing studies describing vibrational properties of natural buzz-pollination are limited to bumblebees(Bombus)and carpenter bees(Xvlocopa)constraining our current understanding of this behavior and its evolution.Therefore,we attempted to minimize this shortcoming by testing whether flower anthers exhibit optimal frequency for pollen release and whether bees tune their buzzes to match these(optimal)frequencies.If true,certain frequencies will trigger more pollen release and lighter bees will reach buzz frequencies closer to this optimum to compensate their smaller buzz amplitudes.Two strategies were used to test these hypotheses:(i)the use of(artificial)vibrational playbacks in a broad range of buzz frequencies and amplitudes to assess pollen release by tomato plants(Solarium Ivcopersicum L.)and(ii)the recording of natural buzzes of Neotropical bees visiting tomato plants during pollination.The playback experiment indicates that although buzz frequency does affect pollen release,no optimal frequency exists for that.In addition,the recorded results of natural buzz-pollination reveal that buzz frequencies vary with bee genera and are not correlated with body size.Therefore,neither bees nor plants are tuned to optimal pollen release frequencies.Bee frequency of buzz-pollination is a likely consequence of the insect flight machinery adapted to reach higher accelerations,while flower plant response to buzz-pollination is the likely result of its pollen granular properties.  相似文献   

19.

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  相似文献   

20.
鲁先文  马瑞君    孙坤 《生态学报》2008,28(6):2518-2518~2525
连续两年对雌雄异株的中国沙棘(Hippophae rhamnoides L.ssp. sinensis Rousi)的花期物候、开花式样进行了观察,检测了柱头可授性、花粉活力和花粉-胚珠比,并进行了人工授粉、套袋实验,检测结实率.结果显示中国沙棘风媒传粉发生在4月下旬或5月上旬,单花花期约7d;传粉盛期在第3天到第5天,柱头的可授性从第2天到第4天或第5天.中国沙棘花先叶开放、花小、无花冠、花药成熟时萼片从两个侧面的纵缝中裂开,形成有利于花粉散布的对流风洞.花粉活力和花粉-胚珠比都很高,具有典型的适应风媒传粉的花部特征和性状.中国沙棘花粉的传播距离受外界环境影响较大,通过重力玻片法检测可以看出,在风速小于3m/s时,主要集中在15~25m之间,当风速大于3m/s时,顺风向可超过85m以外.传粉效率对结实率的影响较大.  相似文献   

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