Contrasted pollen capture mechanisms in Phyllocladaceae and certain Podocarpaceae (Coniferales)

Comparative study shows that Phyllocladus and representative Podocarpaceae differ in the mechanism by which pollen is introduced into the pollen chamber and onto the apex of the nucellus ("pollen capture"). Both types involve a pollination drop, but only in Podocarpaceae is it consistently inverted and in contact with adjacent surfaces. Phyllocladus has functionally nonsaccate pollen (although a vestigial saccus has been claimed); its pollen is wettable and sinks in water. Podocarpaceae (except Saxegorhaea) have saccate pollen, which is nonwettable and floats on water. In Phyllocladus the pollination drop receives the pollen directly and presence of pollen stimulates complete drop withdrawal, which may be a metabolic process. Once pollinated, an ovule does not resecrete a pollination drop. In Podocarpaceae the drop usually receives the pollen indirectly via pollen scavenging and saccate pollen is preferentially captured. The retraction of the drop appears to be the result of evaporation and is presumably nonmetabolic. Drop secretion can be repeated in the presence of pollen. A major consequence of these contrasted mechanisms is that in Phyllocladus the entire contents of the pollination drop are ingested, whereas in Podocarpaceae only that part of the drop that includes saccate pollen is ingested. Because of differences in repeatability of the secretion process, Podocarpaceae are likely to capture more pollen. In neither mechanism does the process favor 'own" pollen. but in Podocarpaceae all but saccate pollen is excluded. We thus have further evidence for differences in pollen capture mechanisms in conifers with a pollination drop, and differences in the behavior of the pollination drop itself.     

Reconstruction of the Jurassic conifer Sewardiodendron laxum (Taxodiaceae)

Compressed seed cones and pollen cones of Sewardiodendron laxum are described from the Middle Jurassic of Yima, Henan, central China. They are either organically attached to or associated with leafy shoots. Seed cones are terminally borne. Each cone is ovate to elongated, up to 6.5 cm long and 3.5 cm wide, and consists of a stout axis and numerous helically arranged bract-scale complexes. The bract protrudes beyond and is partially fused with the reduced ovuliferous scale. The ovuliferous scale bears approximately six inverted, small, and flattened seeds. Pollen cones are borne in terminal clusters. Microsporophylls are helically arranged, each bearing three abaxial, basally fused pollen sacs. Pollen is assaccate, rounded, and with an inconspicuous pore. Morphological, structural, and cuticular features of seed cones, pollen cones, and leafy shoots of S. laxum are compared with those of fossil and extant conifers. S. laxum is included in Taxodiaceae and believed to have its closest affinities with a Mesozoic conifer Elatides and a group of Cunninghamia-like conifers. It is reconstructed as a half-evergreen tree that lived in a humid, warm-temperate climate.     

Pollination drop in Juniperus communis: response to deposited material

BACKGROUND AND AIMS: The pollination drop is a liquid secretion produced by the ovule and exposed outside the micropyle. In many gymnosperms, pollen lands on the surface of the pollination drop, rehydrates and enters the ovule as the drop retracts. The objective of this work was to study the formation of the pollination drop in Juniperus communis, its carbohydrate composition and the response to deposition of conspecific pollen, foreign pollen and other particulate material, in an attempt to clarify the mechanism of pollination drop retraction. METHODS: Branches with female cones close to pollination drop secretion were collected. On the first day of pollination drop exposure, an eyelash mounted on a wooden stick with paraffin was used to collect pollen or silica gel particles, which were then deposited by contact with the drop. Volume changes in pollination drops were measured by using a stereomicroscope with a micrometer eyepiece 3 h after deposition. The volume of non-pollinated control drops was also recorded. On the first day of secretion, drops were also collected for sugar analysis by high-performance liquid chromatography. KEY RESULTS: The pollination drop persisted for about 12 d if not pollinated, and formed again after removal for up to four consecutive days. After pollination with viable conspecific pollen, the drop retracted quickly and did not form again. Partial withdrawal occurred after deposition of other biological and non-biological material. Fructose was the dominant sugar; glucose was also present but at a much lower percentage. CONCLUSIONS: Sugar analysis confirmed the general trend of fructose dominance in gymnosperm pollination drops. Complete pollination drop withdrawal appears to be triggered by a biochemical mechanism resulting from interaction between pollen and drop constituents. The results of particle deposition suggest the existence of a non-specific, particle-size-dependent mechanism that induces partial pollination drop withdrawal. These results suggest that the non-specific response may decrease the probability of pollen landing on the drop, reducing pollination efficiency.     

Anatomy and relationships of petrified seed cones of the Cupressaceae,Taxodiaceae, and Sciadopityaceae

Morphological features and the affinities of petrified seed cones of the Taxodiaceae, Cupressaceae, and Sciadopityaceae are reevaluated. TheCunninghamia-like plants are the earliest record of the families.Parataiwania explains the divergence ofTaiwania from aCunninghamia-like ancestral form by loss of ovuliferous scale and sclerenchyma.Sequoia-like allies probably diverged from a Cretaceous transitional plant likeYezosequoia and are most diversified in the Late Cretaceous.Yubaristrobus andArchicupressus suggest monophyly ofTaxodium and its allied genera of the Taxodiaceae and the Cupressaceae. The Cupressaceae may be derived from a taxodiaceous plant with orthotropous seeds, acquiring a reniform arrangement of the vascular bundles of the bract-scale complex, decussate or whorled phyllotaxis, and spherical cones consisting of a small number of bract-scale complexes. The Cretaceous fossils of the Sciadopityaceae suggest the ancient origin of the family.     

野慈姑(Sagittaria trifolia)人工授粉的有效性:授粉工具与花粉保存

人工授粉作为一项关键性技术在生态研究与农业生产领域广泛应用,但是授粉过程中的工具有效性以及花粉保存尚未受到研究者的关注。以植物野慈姑为试验材料,研究了人工授粉方式和花粉保存条件对繁殖的影响。两年的结果表明,相较于直接接触式授粉,借助毛刷工具授粉显著降低了野慈姑的座果概率和种子数。花粉经过半小时野外高温保存后,其授粉的座果概率显著高于对照组,但单果种子数和种子大小保持稳定。综上,在野慈姑的人工授粉中应避免使用毛刷等工具,离体花粉可置于野外环境适当保存。在相关应用中,研究人员应当警惕工具授粉造成的负面影响,注意比较工具与其他授粉方式的等效性,保证实验的科学性。     

Seed cone and ovule ontogeny in Metasequoia, Sequoia and Sequoiadendron (Taxodiaceae–Coniferales)

TAKASO, T. & TOMUNSON, P. B., 1992. Seed cone and ovule ontogeny in Metasequoia, Sequoia and Sequoiadendron (Taxodiaceae–Coniferales). Structural features of seed cones, up to the initiation of ovules, are developed in the three genera in the summer and late fall prior to pollination in the following spring, when cones renew their further development. Bracts are initiated in a decussate manner in Metasequoia but spirally in Sequoia and Sequoiadendron. No ovuliferous scale is initiated, at most there is a shallow mound of tissue on the adaxial surface of the bract from which the ovules are developed. Metasequoia produces a single series of up to eight ovules in acropetal (centripetal order), Sequoia and Sequoiadendron produce a double series of six to nine ovules, also in acropetal order, since a second series of ovules appear distal to and alternate with the first series. Common features that unite the genera are the somewhat peltate configuration of the cone scales due to late intercalary expansion, the derivation of the vascular supply to the bract-ovule complex from a single bundle and the usual inversion of the seed during late development, to which can be added developmental features. The absence of tooth-like structures, present in some other Taxodiaceae, is discussed in relation to cone evolution in the family being determined by changes in developmental timing. Emphasis is made on the way in which features of morphogenesis, determine cone organization in the three genera independent of an interpretation that relies solely on hypothetical ancestral forms.     

Heterostyly promotes compatible pollination in buckwheats: Comparisons of intraflower,intraplant, and interplant pollen flow in distylous and homostylous Fagopyrum

Premise of the Study

Heterostyly, the reciprocal positioning of stigmas and anthers in different floral morphs, has long been thought to promote intermorph pollination. However, extensive intramorph pollination occurs commonly in heterostylous species, leading to recurrent questions about the functional and evolutionary significance of heterostyly.

Methods

To identify the sources of stigmatic pollen (autogamous [intraflower], geitonogamous [intraplant], vs. interplant), we emasculated either one flower or entire plants in experimental populations of the two closely related buckwheat species, distylous Fagopyrum esculentum and homostylous F. tataricum. Differences in pollen size allowed unambiguous identification of pollen on stigmas.

Results

Only 2.4% of F. tataricum pollen and 1.5% of F. esculentum pollen arrived successfully on compatible stigmas of other plants. In the former (homostylous) species, 71.3% of the pollen load on stigmas was autogamous, 10.8% was geitonogamous, and 17.9% was interplant. In the latter (distylous) species, 37.45% of the pollen on stigmas was autogamous, 13.8% was geitonogamous, 17.0% was intramorph, and 31.75% was intermorph. The amount of incompatible pollen arriving on stigmas was greatly decreased by both one‐flower and whole‐plant emasculations, and thus, the proportion of compatible pollen deposited increased with one‐flower emasculation and increased even more with whole‐plant emasculation.

Conclusions

Our quantification of pollen‐donor sources in these two species indicated that heterostyly in Fagopyrum esculentum provided a nearly 2‐fold fitness advantage (in terms of compatible pollination) over expected (random) pollen transfers between morphs. Because of reduced herkogamy, the homostylous F. tataricum was highly autogamous.     

Incompatibility and pollen competition in Alnus glutinosa: Evidence from pollination experiments

Different types of incompatibility systems were found to operate simultaneously in alnus glutinosa in the course of numerous pollination experiments, including self-pollination and pollination with controlled pollen mixtures. Isozyme genetic markers were used to identify the pollen parent of each offspring from the mixed pollination experiments, thus allowing specification of the fertilization success of each pollen parent. In a first step, these results were compared with observations on in vitro pollen germination experiments. This comparison allows for exploration of the explanatory value of different germination media as models of germination conditions on stigmas. In most cases, the data suggest that the in vitro germination conditions resemble the fertilization conditions in vivo, at least in the sense that they favor the same pollen parents. By providing a generic and operable definition of the two basic types of incompatibility, eliminating (inability to fertilize ovules) and cryptic (resulting in lowered fertilization success of a pollen parent under competition), evidence was detected for the existence of both types of incompatibility in alnus glutinosa, where eliminating incompatibility occurred as self-incompatibility only. However, since this incompatibility seems to act primarily via pollen elimination, seed production is not likely to be negatively affected in natural populations, even for comparatively large amounts of self-pollination. This revised version was published online in July 2006 with corrections to the Cover Date.     

巴旦杏授粉试验及花粉管生长的荧光显微观察

对巴旦杏4个主栽品种的授粉试验结果表明：自花授粉试验中“鹰嘴”、“双果”坐果率分别为19．74％和4．29％,“纸皮”和“麻壳”坐果率为零;自然授粉状态下4个主栽品种坐果率在9．37％－20．13％之间;人工异花授粉试验中4个品种9个授粉组合坐果率绝大多数都在40％以上,其中混合花粉授粉试验结果最好;人工养蜂条件下的坐果率明显高于无蜂自然授粉的坐果率,昆虫传粉可达到人工混合花粉授粉的效果。授粉试验还表现出巴旦杏雌蕊柱头的有效授粉时间为开花后花期的前4d,授粉越早,越有利于坐果。荧光显微观察表明,巴旦杏花粉管最早约于授粉第7天通过花柱基部到达子房室,第8天前后到达胚珠。     

Influences of Cross Pollination on Pollen Tube Growth and Fruit Set in Zuili Plums （Prunus salicina）

Zuili plum （Prunus salicina L.） trees usually set fruit poorly, although they produce high quality fruit. To elucidate the causes of the poor fruit set, pollen tube growth into pistils and fruit set percentage were investigated after cross-, self- and open-pollination. Ovule development in Zuili pistils was also investigated. Pollen tube penetration into the ovules via the obturator and micropyle was best when Zuili pistils were pollinated by cv. Black Amber （P. domestica） pollen grains, although cross-pollinations with Hongxinli and Miili （P. salicina） pollen were more effective than self- and open-pollination. The fruit set percentage was also highest in pistils pollinated with Black Amber pollen grains. Morphological observation of Zuili pistils revealed that the trees produce ＂double pistils＂, developing two ovaries from a basal pistil, at a rate as high as 28%. In such abnormal pistils, most ovules were lacking an embryo sac or were entirely degenerated. The percentage of normally developed ovules was 24.3% and 8.9% in normal and double pistils, respectively. From these results, we conclude that the main causes of poor fruit set of Zuili plums are a lack of effective cross-pollination and the production of high percentages of double pistils in which normally developed ovules are scarcely formed.     

Embryonal Development of Siberian Pine (Pinus sibirica Du Tour) with the Annual Cycle of Ovulate Cone Development in the Western Sayan Mountains

Morphological and cytoembryological studies of unique forms of Siberian pine (Pinus sibirica Du Tour) with the annual cycle of ovulate cone development were carried out. The morphogenesis of female shoots in these anomalous trees with the annual cycle of development was shown to provide for an accelerated development and very rapid growth of megastrobili and accelerated formation of reproductive structures. Cytoembryological studies of ovules in these trees showed significant shortening of the free-nuclear stage of gametophyte development (up to 3 weeks instead of one year) and ultra-early formation of archegonia and egg cells. However, no fertilization of egg cells and the development of embryos occur in the anomalous forms; the egg-cell nucleus divides in the haploid state. Developing seeds are formed without embryo. The author hypothesizes that the acceleration of embryo development in the anomalous pine trees is caused by enhanced hormonal, carbohydrate, and nitrogen metabolism in ovule tissues.     

Natural levels of pollination intensity and effects of pollen loads on offspring quality in females of Thalictrum pubescens (Ranunculaceae)

Females of Thalictrum pubescens produce stamens containing sterile pollen. Earlier studies have shown the presence of stamens does not increase seed set through increased visitation by insects. Insects may, however, increase pollen deposition on stigmas and increase pollen competition. This paper examines: 1) pollen deposition levels in natural populations and 2) whether larger pollen loads lead to higher quality offspring. The majority of stigmas in two populations received less than 15 grains, but a small proportion had large loads. The latter may represent the occasional insect visit. These visits may provide the opportunity for pollen tube competition. In the greenhouse, flowers receiving heavy-pollen loads had higher seed set than flowers receiving light-pollen loads. Seeds from the two treatments were the same mass, had similar germination times, and seedlings had the same mass. In conclusion, it does not appear that pollen tube competition can account for the maintenance of stamens in females of T. pubescens.     

裸子植物雄球花——纤细堆囊穗的修订

研究了采自中国东北晚三叠世羊草沟组的一种裸子植物雄球花——纤细堆囊穗。通过对新材料的研究,发现这种雄球花小孢子叶末端的裂片在形态上差异很大,这是原来没有发现的十分重要的特征。这些保存精美的化石对于纤细堆囊穗特征的阐明和修订以及物种复原很有帮助。将中国的标本与西伯利亚堆囊穗、小堆囊穗、乌尔马堆囊穗和被定为长叶拜拉的雄球花进行了比较,发现它们与纤细堆囊穗在特征上一致,故将它们处理为纤细堆囊穗的异名。修订后的纤细堆囊穗包括上面所提到的所有种。同时,也讨论了堆囊穗属可能的演化意义。它可能是银杏属的远祖,经过小孢子囊数目的减少和小孢子叶长度的缩短而演化到现在的银杏,而产自辽西早白垩世的辽宁银杏可能代表了堆囊穗和现代银杏在形态演化上的一个中间步骤。     

Effects of sequential pollination on the success of "fast" and "slow" pollen donors in Hibiscus moscheutos (Malvaceae)

Competition among pollen grains for the chance to fertilize ovules typically involves two stages: arrival times on stigmas and/or the growth of pollen tubes through styles. In a previous study of Hibiscus moscheutos, we found that individual pollen donors often differed in pollen tube competitive ability. Here we determined whether short delays in pollen arrival time altered the average success of "fast" and "slow" pollen donors when both types of pollen experienced the same delays. Hand-pollination experiments were carried out using four pairs of pollen donors that differed in competitive ability. We allowed delays of 15 or 30 min between the first and second pollen donor and then determined seed paternity using allozyme markers. The second donor typically sired fewer seeds than pollen that arrived earlier, but, contrary to expectation, "faster" pollen did not always sire significantly more seeds than "slower" pollen when each was applied after delays of the same duration. In two of the four pairs of donors, differences that were seen following simultaneous pollinations disappeared when each type of pollen was applied following identical delays of 15 or 30 min. This unexpected response suggests that the dynamics of pollen tube competition are more complex than anticipated.     

Ragweed pollen sampling in Gallarate (North-West of Milan) during four years (1987–1990)

Summary Ambrosia artemisifolia (elatior), an imported pollen, never sampled in the past in the atmosphere of our country, is beconning the most important Composita responsible for late-summer pollinosis in our area.The documented increased pollen concentration confirmed greater sensitization in people coming to our allergological centre.     

The behavior of pollination drop secretion in Ginkgo biloba L.

Pollination drop (PD) secretion plays a critical role in wind pollination in many gymnosperms. We conducted detailed investigations on PD secretion in Ginkgo biloba, and found that PDs could not form when the micropyle was removed, but were able to form after removal of the shoot, leaves, ovular stalk, or ovular collar. The duration and volume of the PD increased under high relative humidity, but addition of salt or sugar did not affect PD secretion, its size, or its duration. Morphological and anatomical observations showed that many secretion cells at the nucellus tip contributed to secreting the PD after the formation of pollen chamber. Under laboratory conditions, the PD persisted for approximately 10 d if not pollinated, and re-formed five times after it was removed, with the total volume of PDs reaching approximately 0.4 μL. These results suggested that PDs can be continuously secreted by the tip of the nucellus cells during the pollination stage to increase the chance of capturing pollen from the air. Importantly, PD secretion is an independent behavior of the ovule and PDs were produced apoplastically.     

Pollination effectiveness and pollen dispersal in a Rhododendron ferrugineum (Ericaceae) population

Many alpine plants are predominantly outcrossing, thus plant reproductive success is highly dependent on effectiveness of pollinators. How pollinators transfer pollen from one flower to another is of great interest in understanding the genetic structure in plant populations. We studied (1) the role and effectiveness of insect visitors for pollination, and (2) their contribution as pollen vectors for gene dispersal in a Rhododendron ferrugineum population. Various insect visitors were recorded, including Hymenoptera, Diptera, Coleoptera, and Lepidoptera. The most frequent and effective insects were honey bees and bumblebees. Muscid flies were considered as important pollinators, particularly due to their relatively high visitation rate. Syrphid flies, Formicidae, and Coleoptera were ineffective in transporting pollen, while the effectiveness of Lepidoptera and Empididae was negligible. A fluorescence labelling experiment revealed that pollen dispersal was restricted (0 - 2 m) in a dense R. ferrugineum stand and decreased in a leptokurtic fashion. This might lead to geitonogamous self-pollination that could explain the close relationship between individuals found in genetic studies of R. ferrugineum. However, some pollen grains may travel 40 - 45 m, which implies the occurrence of cross-pollination through the foraging activities of bumblebees and honey bees.     

Beyond pollen:ovule ratios: Evolutionary consequences of pollinator dependence and pollination efficiency for pollen and ovule production in angiosperms

Premise

The relative per-flower production of ovules and pollen varies broadly with angiosperm mating systems, with outcrossing types commonly producing more pollen grains per ovule than selfing types. The evolutionary causes of this variation are contentious, especially the relevance of pollination risk. Resolution of this debate may have been hampered by its focus on pollen:ovule (P:O) ratios rather than on the evolution of pollen and ovule numbers per se.

Methods

Using published mean ovule and pollen counts, we analyzed associations with the proportion of removed pollen that reaches stigmas (pollen-transfer efficiency) and differences between pollinator-dependent and autogamous forms within and among species. Analyses involved Bayesian methods that simultaneously considered variation in pollen and ovule numbers and accounted for phylogenetic relatedness. We also assessed the utility of P:O ratios as mating-system proxies and their association with female outcrossing rates.

Results

Median pollen number declined consistently with pollen-transfer efficiency among species, whereas median ovule number did not. Similarly, in both intraspecific and interspecific analyses, pollinator-dependent plants produced more pollen than autogamous plants, whereas ovule production did not differ statistically. Distributions of P:O ratios overlapped extensively for self-incompatible and self-compatible species and for different mating-system classes, and P:O ratios correlated weakly with outcrossing rate.

Conclusions

Our findings demonstrate that pollinator dependence and pollination efficiency commonly influence the evolution of pollen number per flower but have more limited effects on ovule number. P:O ratios provide ambiguous, possibly misleading, information about mating systems, especially when compared among clades.

     

Dioecy and the evolution of pollination systems inSchiedea and Alsinidendron (Caryophyllaceae:Alsinoideae) in the Hawaiian Islands

The transition from biotic to wind pollination and the consequencesof this transition for the evolution of dioecious breeding systems wereinvestigated in Schiedea and Alsinidendron(Caryophyllaceae: Alsinoideae), genera endemic to the Hawaiian Islands. The potential for wind pollination was studied for five species ofSchiedea using a wind tunnel. Morphological correlates of windpollination for these species were then used to infer the presence orabsence of wind pollination in the remaining Schiedea species. Hermaphroditic Alsinidendron and Schiedea species,which occur in mesic to wet forests, or less commonly in dry habitats,show little or no indication of wind pollination. These species had lowpollen:ovule ratios, large relative pollen size, diffuse inflorescences,substantial nectar production in several cases, and appear to bebiotically pollinated or autogamous. Sexually dimorphic species, whichall occur in dry habitats, are wind pollinated, based on wind tunnelresults or morphological adaptations indicating the potential for windpollination. These adaptations include high pollen:ovule ratios, smallpollen size, moderately to highly condensed inflorescences, and reducednectaries and nectar production. Shifts to wind pollination anddimorphism are strongly correlated in Schiedea, suggesting theclose functional relationship of the twophenomena.