Adaptive function of duodichogamy: Why do chestnut trees have two pollen emission phases?

Premise

Intersexual mating facilitation in flowering plants has been largely underexplored. Duodichogamy is a rare flowering system in which individual plants flower in the sequence male-female-male. We studied the adaptive advantages of this flowering system using chestnuts (Castanea spp., Fagaceae) as models. These insect-pollinated trees produce many unisexual male catkins responsible for a first staminate phase and a few bisexual catkins responsible for a second staminate phase. We hypothesized that duodichogamy increases female mating success by facilitating pollen deposition on stigmas of the rewardless female flowers through their proximity with attractive male flowers responsible for the minor staminate phase.

Methods

We monitored insect visits to 11 chestnut trees during the entire flowering period and explored reproductive traits of all known duodichogamous species using published evidence.

Results

In chestnuts, insects visited trees more frequently during the first staminate phase but visited female flowers more frequently during the second staminate phase. All 21 animal-pollinated duodichogamous species identified are mass-flowering woody plants at high risk of self-pollination. In 20 of 21 cases, gynoecia (female flower parts) are located close to androecia (male flower parts), typically those responsible for the second minor staminate phase, whereas androecia are often distant from gynoecia.

Conclusions

Our results suggest that duodichogamy increases female mating success by facilitating pollen deposition on stigmas by means of the attractiveness of the associated male flowers while effectively limiting self-pollination.

     

雌雄异熟植物露蕊乌头开花时间对雌雄功能期及表型性别的影响

开花时间决定了植物雌雄功能的交配机会, 最终影响繁殖成功。交配环境假说认为雌雄异熟植物开花时间的差异能引起植物表型性别的变异, 改变种群内的交配环境, 影响植物对雌雄功能的最佳性分配。为了研究开花时间对雌雄异熟植物的雌雄性别时期及表型性别的影响, 本文以毛茛科雄性先熟植物露蕊乌头(Aconitum gymnandrum)为实验材料, 记录了雄性和雌性功能期, 分析了植株开花时间、花的雌雄功能期和表型性别的关系。结果表明: 在植物同一花序内, 较晚开放的花有更长的雄性期和更短的雌性期, 性分配在时间上偏雄。雌雄功能期在时间上的相对分配随植物开花时间的变化表现出相似的趋势： 较晚开的花或较晚开花的个体, 花的雄性功能期相对于雌性功能期更长, 在时间上更偏向雄性功能。而且, 开花时间的差异影响种群内花的性比和植物个体的表型性别动态。随着开花时间由早到晚的变化, 种群内早期以雄花为主,末期以雌花为主, 种群内性别环境由偏雄向偏雌变化, 因此植株个体的平均表型性别则从偏雌转向偏雄。本文结果支持交配环境假说, 雄性先熟的露蕊乌头开花早期, 种群内花的性别比偏雄, 种群表型性别环境偏雄, 因而植物个体平均表型性别偏雌, 性别分配(即时间分配)偏向雌性功能, 而晚开花个体的平均性别偏雄, 更偏向雄性功能的分配。     

Fly pollination and duodichogamy in Bridelia stipularis and Cleistanthus sumatranus (Phyllanthaceae)

The globally distributed family Phyllanthaceae comprises 1745 species, all of which produce unisexual flowers, and most of the species occur in the tropics. The few species in which the pollination system has been studied are pollinated by flies, wasps, or specialized moths, and at least two species are duodichogamous. Here we report on the flowering phenology and pollination of the Chinese species Bridelia stipularis and Cleistanthus sumatranus, which were studied on the tropical Hainan Island. Both species are duodichogamous and indicated to be fly‐pollinated. The calliphorid species Chrysomya megacephala, Isomyia isomyia, Pierretia sp., Hemipyrellia sp., and Achoetandrus tufifacies were the most frequent flower visitors to B. stipularis; some of these species together with syrphids (Meliscaeva cinctella, Dideopsis sp., and Eristalis sp.) were the most abundant visitors to Cl. sumatranus flowers. Fruit set in hand‐pollinated flowers was higher than in open‐pollinated flowers, but because of natural fruit abortion the difference was not significant, indicating that fly pollinators do not limit reproduction of these Phyllanthaceae species. This study provides an addition to the limited Asian literature on pollination by flies and the even more limited literature on duodichogamy. It could help to broaden our understanding for analysis of the evolution of duodichogamy in future research, but the specific ecological factors behind the evolution of this rare sexual system are still difficult to generalize.     

Influence of asymmetrical mating patterns and male reproductive success on the maintenance of sexual polymorphism in Acer pictum subsp. mono (Aceraceae)

Populations of Acer species often contain more than three sex phenotypes with complex sexual polymorphism including duodichogamy, protandry and protogyny. We identified the mechanisms that maintain sexual polymorphism in Acer pictum subsp. mono, a temperate tree from northern China, by investigating maternal mating patterns and male reproductive success. We used paternity analyses to estimate rates of outcrossing and disassortative mating, as well as male outcrossed siring success, in a population of A. pictum subsp. mono with uneven sex phenotype ratios (duodichogamous 69.1%, protandrous 19.6%, protogynous 11.3%). We used a pollen‐transfer model to investigate whether the unequal ratios of sex phenotypes could be explained by the observed patterns of mating. Most progeny resulted from outcrossing, particularly disassortative among the sex phenotypes. Although the duodichogamous phenotype showed a significant amount of intraphenotypic mating, the frequency did not exceed that of disassortative mating. We detected no significant differences in male outcrossed siring success among the sex phenotypes. The pollen‐transfer model demonstrated that sex phenotype ratios could be maintained by the observed mating pattern in the population. Our results indicate that disassortative mating among the sex phenotypes can maintain sexual polymorphism in A. pictum subsp. mono and that ratios biased towards duodichogamy can result from frequent intraphenotypic mating in this phenotype.     

Relationship between floral longevity and sex allocation among flowers within inflorescences in Aquilegia buergeriana var. oxysepala (Ranunculaceae)

Understanding the fitness of plants with inflorescences requires examining variation in sex allocation among flowers within inflorescences. We examined whether differences in the duration of the male and female phases of flowering lead to variation in sex allocation and reproductive success among flowers within inflorescences. In 2002 and 2003, we quantified floral longevity, floral sex allocation, and reproductive success between the first and the second flowers within inflorescences in a protandrous species, Aquilegia buergeriana var. oxysepala. Floral longevity was greater in the first flowers than in the second ones in both years. The male phase lasted longer, and the initial number of pollen grains and the number of pollen grains removed were greater in the first flowers than in the second ones in both years. Within first flowers, the number of pollen grains removed was greater in flowers that had longer male phases, thus duration of the male phase may positively affect male reproductive success in the first flowers. The female phase lasted longer and the number of ovules was greater in the first flowers than in the second only in 2002. However, seed production per flower and female phase duration in both years were not significantly related. The variation in the number of pollen grains among flowers in this species may be caused by the variation in male phase duration.     

FLOWERING,SEX RATIOS,POLLEN-OVULE RATIOS,FRUIT SET,AND REPRODUCTIVE EFFORT OF A DIOECIOUS TREE,MYRISTICA INSIPIDA (MYRISTICACEAE), IN TWO DIFFERENT RAIN FOREST COMMUNITIES

The flowering of Myristica insipida R. Br. was studied in two rain forest communities in northern Queensland. This dioecious, subcanopy tree had a male-biased sex ratio at both study sites. In the lowland population the male-bias could be attributed to males (trees producing staminate flowers) starting to flower at a smaller average size than females (trees producing pistillate flowers). There were no intersexual differences in spacing or distribution within the study sites. Males trees flowered earlier, flowered longer, and produced over twice as many flowers as females during the study season. Although the onset of flowering was rather variable, 18–22 days following heavy rains, most trees had a synchronous period of maximum flowering. Pollination manipulations determined that there was no fruit development without pollination, and that increasing pollen loads resulted in increased fruit set with diminishing effect. Taking into account the sex-ratios and intersexual differences in flower production, the pollen-ovule ratio was calculated to be 16,000–19,000. Male trees were found to expend more energy on flowering than female trees. Open-pollination resulted in 1.0% of female flowers setting fruit. The much greater cost of fruit production resulted in females expending 421% more energy on reproduction than males. Fruit and seed production were judged to be pollination-limited. Nonetheless, this species exhibited several characteristics that are predicted if dioecy evolved by means of sexual selection.     

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.     

How New Guinea rainforest flower resources vary in time and space: Implications for nectarivorous birds

Abstract Variability in spatial and temporal patterning of flowering by populations of rainforest trees fed upon by honeyeaters and flower-visiting parrots was studied for 2 years in lowland tropical hill forest in Papua New Guinea. All 2200 trees in a 3 ha plot were tagged, identified, mapped and monitored monthly. Of 274 tree species present, 86 flowered during the course of the study; during any given month, approximately 20% of the species flowering that month were visited by nectarivorous birds. Results showed that overall flower resources (total number of species, and number of bird-pollinated species, individuals and flowers) fluctuated during the year, decreased during the dry season and increased during the wet season. In addition, there was a wide range of temporal variation within and among tree species in length and timing of flowering period, percentage of each conspecific population flowering from year to year, and degree of synchrony among flowering conspecifics. Spatial dispersion of tree populations also varied, from clumps to scattered single individuals. Resident bird species were correlated with synchronously flowering trees, whereas nomadic bird species were correlated with asynchronously flowering trees. Resident birds were also associated with smaller blooming displays per tree, whereas nomadic birds were associated with trees that bloomed massively. There was no correlation between avian nomadism and spatial dispersion of tree populations. Thus nomadic birds seem to range in search of rich but unpredictable resources; resident birds may rely more on predictable, but smaller resources.     

A test of pollinator specificity and morphological convergence between nectarivorous birds and rainforest tree flowers in New Guinea

Interactions between flowering trees in a representative sample of vegetation, and the birds that fed at their flowers, were studied for 2 years in lowland tropical hill forest in New Guinea. All 2,200 trees in a 3-ha plot were tagged, identified, mapped, and monitored monthly. Approximately 60% of all individual trees flowered during the study; all species that these flowering individuals belonged to were evaluated for bird visitation. Approximately 13% of the 164 resident species of New Guinea avifauna at the study site, especially honeyeaters and parrots, visited flowers. In the forest inventory plot, approximately 15–22% of all 86 tree species that flowered during the study were visited by birds; most of these tree species were canopy species. Results showed that there was no statistically significant correlation between bird species grouped by bill morphology and flower species grouped as morphotypes and ranked by nectar accessibility, although strong but unexpected bird/plant associations were evident. These associations may be related to variables such as body mass or perch size. These results are discussed in comparison with results from the Neotropics and Australia, and in terms of morphological convergence and pollinator specificity in pollination systems.     

Constitutive expression of Arabidopsis LEAFY or APETALA1 genes in citrus reduces their generation time

Citrus trees have a long juvenile phase that delays their reproductive development by between 6 and 20 years, depending on the species. With the aim of accelerating their flowering time, we transformed juvenile citrus seedlings to constitutively express the Arabidopsis LEAFY (LFY) or APETALA1 (AP1) genes, which promote flower initiation in Arabidopsis. Both types of transgenic citrus produced fertile flowers and fruits as early as the first year, notably through a mechanism involving an appreciable shortening of their juvenile phase. Furthermore, expression of AP1 was as efficient as LFY in the initiation of flowers, and did not produce any severe developmental abnormality. Both types of transgenic trees flowered in consecutive years, and their flowering response was under environmental control. In addition, zygotic and nucellar derived transgenic seedlings had a very short juvenile phase and flowered in their first spring, demonstrating the stability and inheritance of this trait. These results open new possibilities for domestication, genetic improvement, and experimental research in citrus and other woody species.     

Gender variation of sequential inflorescences in a monoecious plant Sagittaria trifolia (Alismataceae)

In protogynous plants, female flowers of early blooming plants are at a reproductive disadvantage because they cannot set fruit due to the lack of available pollen. To study this phenomenon, gender expression of the monoecious herb Sagittaria trifolia was investigated over the entire flowering season in two field and two cultivated populations in Hubei and Hunan Provinces, China. In racemes of S. trifolia, flowers open sequentially from bottom to top, with female flowers opening first followed by male flowers. This creates a temporal separation of sexes in the species. Under field conditions small plants are often male, with production of both male and female flowers increasing with plant size. Femaleness increased among sequential inflorescences since female flower production increased whereas male flower production did not. Seed production was greater in large inflorescences because they contain more female flowers, and the number of ovules increased in female flowers at basal positions within the raceme. A consistent pattern of high seed set was observed in flowers from both field and cultivated populations. About 1 % of unfertilized ovules resulted from no pollination and 2 % of the seeds produced were only partly developed due to resource limitation. In the first inflorescence of the six experimental populations, 6.7-40.0 % of individuals produced only male flowers, and female flowers of 1.9-6.5 % individuals were aborted. The occurrence of male flowers in early blooming inflorescences could be an adaptive strategy to conserve resources and enhance pollination of female flowers in protogynous S. trifolia.     

The roles of female and hermaphroditic flowers in the gynodioecious–gynomonoecious Silene littorea: insights into the phenology of sex expression

Some gynodioecious species have intermediate individuals that bear both female and hermaphroditic flowers. This phenomenon is known as a gynodioecious–gynomonoecious sexual system. Gender expression in such species has received little attention in the past, and the phenologies of male and female functions have also yet to be explored. In this study, we examined variations in gender patterns, their effects on female reproductive success and sex expression in depth throughout the flowering period in two populations. The studied populations of Silene littorea contained mostly gynomonoecious plants and the number of pure females was very low. The gynomonoecious plants showed high variability in the total proportion of female flowers. In addition, the proportion of female flowers in each plant varied widely across the flowering season. Although there was a trend towards maleness, our measures of functional gender suggested that most plants transmit their genes via both pollen and ovules. Fruit set and seed set were not significantly different among populations; in contrast, flower production significantly varied between the two populations – and among plants – with consequent variation in total seed production. Conversely, gender and sex expression were similar in both populations. Plants with higher phenotypic femaleness did not have higher fruit set, seed set or total female fecundity. The mating environment fluctuated little across the flowering period, but fluctuations were higher in the population with low flower production. We therefore conclude that the high proportion of gynomonoecious individuals in our studied populations of S. littorea may be advantageous for the species, providing the benefits of both hermaphroditic and female flowers.     

The evolution of dioecy, heterodichogamy, and labile sex expression in Acer

The northern hemisphere tree genus Acer comprises 124 species, most of them monoecious, but 13 dioecious. The monoecious species flower dichogamously, duodichogamously (male, female, male), or in some species heterodichogamously (two morphs that each produce male and female flowers but at reciprocal times). Dioecious species cannot engage in these temporal strategies. Using a phylogeny for 66 species and subspecies obtained from 6600 nucleotides of chloroplast introns, spacers, and a protein-coding gene, we address the hypothesis (Pannell and Verdú, Evolution 60: 660-673. 2006) that dioecy evolved from heterodichogamy. This hypothesis was based on phylogenetic analyses (Gleiser and Verdú, New Phytol. 165: 633-640. 2005) that included 29-39 species of Acer coded for five sexual strategies (duodichogamous monoecy, heterodichogamous androdioecy, heterodichogamous trioecy, dichogamous subdioecy, and dioecy) treated as ordered states or as a single continuous variable. When reviewing the basis for these scorings, we found errors that together with the small taxon sample, cast doubt on the earlier inferences. Based on published studies, we coded 56 species of Acer for four sexual strategies, dioecy, monoecy with dichogamous or duodichogamous flowering, monoecy with heterodichogamous flowering, or labile sex expression, in which individuals reverse their sex allocation depending on environment-phenotype interactions. Using Bayesian character mapping, we infer an average of 15 transformations, a third of them involving changes from monoecy-cum-duodichogamy to dioecy; less frequent were changes from this strategy to heterodichogamy; dioecy rarely reverts to other sexual systems. Contra the earlier inferences, we found no switches between heterodichogamy and dioecy. Unexpectedly, most of the species with labile sex expression are grouped together, suggesting that phenotypic plasticity in Acer may be a heritable sexual strategy. Because of the complex flowering phenologies, however, a concern remains that monoecy in Acer might not always be distinguishable from labile sex expression, which needs to be addressed by long-term monitoring of monoecious trees. The 13 dioecious species occur in phylogenetically disparate clades that date back to the Late Eocene and Oligocene, judging from a fossil-calibrated relaxed molecular clock.     

Morphological variation in the flowers of Jacaratia mexicana A. DC. (Caricaceae), a subdioecious tree

The Caricaceae is a small family of tropical trees and herbs in which most species are dioecious. In the present study, we extend our previous work on dioecy in the Caricaceae, characterising the morphological variation in sexual expression in flowers of the dioecious tree Jacaratia mexicana . We found that, in J. mexicana , female plants produce only pistillate flowers, while male plants are sexually variable and can bear three different types of flowers: staminate, pistillate and perfect. To characterise the distinct types of flowers, we measured 26 morphological variables. Our results indicate that: (i) pistillate flowers from male trees carry healthy-looking ovules and are morphologically similar, although smaller than, pistillate flowers on female plants; (ii) staminate flowers have a rudimentary, non-functional pistil and are the only flowers capable of producing nectar; and (iii) perfect flowers produce healthy-looking ovules and pollen, but have smaller ovaries than pistillate flowers and fewer anthers than staminate flowers, and do not produce nectar. The restriction of sexual variation to male trees is consistent with the evolutionary path of dioecy from hermaphrodite ancestors through the initial invasion of male-sterile plants and a subsequent gradual reduction in female fertility in cosexual individuals (gynodioecy pathway), but further work is needed to confirm this hypothesis.     

Flowering phenology, display size, and fruit set in an understory dioecious shrub, Aucuba japonica (Cornaceae)

I investigated the effects of display size and flowering phenology on fruit set in Aucuba japonica, an understory dioecious shrub pollinated by opportunistic insects. Natural variations in display size, flowering phenology, and fruit set were monitored in 1997. A hand-pollination experiment was also conducted to check whether pollen limitation was a factor in fruit set in the field. Increases in floral display size did not affect fruit set; the proportion of flowers that set fruit was almost constant irrespective of the total number of flowers per inflorescence, the total number of inflorescences per plant, and the total number of flowers per plant. The hand-pollination experiment showed that fruit set was not pollen limited despite the low mating probability that resulted from the combination of dioecism and the species' dependence on opportunistic pollinators. This was due, in part, to the fact that female flowers did not have a predetermined period of receptivity, but instead remained receptive until they received pollen. In contrast, flowering phenology did affect fruit set. Fruit set was most abundant when male and female flowering was most abundant. This suggests there was some degree of pollen limitation during the part of the flowering season when male flowers were scarce.     

Moving your sons to safety: galls containing male fig wasps expand into the centre of figs, away from enemies

Figs are the inflorescences of fig trees (Ficus spp., Moraceae). They are shaped like a hollow ball, lined on their inner surface by numerous tiny female flowers. Pollination is carried out by host-specific fig wasps (Agaonidae). Female pollinators enter the figs through a narrow entrance gate and once inside can walk around on a platform generated by the stigmas of the flowers. They lay their eggs into the ovules, via the stigmas and styles, and also gall the flowers, causing the ovules to expand and their pedicels to elongate. A single pollinator larva develops in each galled ovule. Numerous species of non-pollinating fig wasps (NPFW, belonging to other families of Chalcidoidea) also make use of galled ovules in the figs. Some initiate galls, others make use of pollinator-generated galls, killing pollinator larvae. Most NPFW oviposit from the outside of figs, making peripherally-located pollinator larvae more prone to attack. Style length variation is high among monoecious Ficus spp. and pollinators mainly oviposit into more centrally-located ovules, with shorter styles. Style length variation is lower in male (wasp-producing) figs of dioecious Ficus spp., making ovules equally vulnerable to attack by NPFW at the time that pollinators oviposit. We recorded the spatial distributions of galled ovules in mature male figs of the dioecious Ficus hirta in Southern China. The galls contained pollinators and three NPFW that kill them. Pollinators were concentrated in galls located towards the centre of the figs, NPFW towards the periphery. Due to greater pedicel elongation by male galls, male pollinators became located in more central galls than their females, and so were less likely to be attacked. This helps ensure that sufficient males survive, despite strongly female-biased sex ratios, and may be a consequence of the pollinator females laying mostly male eggs at the start of oviposition sequences.     

Seasonality of weather and tree phenology in a tropical evergreen mountain rain forest

Flowering and fruiting as phenological events of 12 tree species in an evergreen tropical mountain rain forest in southern Ecuador were examined over a period of 3–4 years. Leaf shedding of two species was observed for 12 months. Parallel to the phenological recordings, meteorological parameters were monitored in detail and related to the flowering and fruiting activity of the trees. In spite of the perhumid climate of that area, a high degree of intra- and inter-specific synchronisation of phenological traits was apparent. With the exception of one species that flowered more or less continuously, two groups of trees could be observed, one of which flowered during the less humid months (September to October) while the second group started to initiate flowers towards the end of that phase and flowered during the heavy rains (April to July). As reflected by correlation coefficients, the all-time series of meteorological parameters showed a distinct seasonality of 8–12 months, apparently following the quasi-periodic oscillation of precipitation and related cloudiness. As revealed by power spectrum analysis and Markov persistence, rainfall and minimum temperature appear to be the only parameters with a periodicity free of long-term variations. The phenological events of most of the plant species showed a similar periodicity of 8–12 months, which followed the annual oscillation of relatively less and more humid periods and thus was in phase or in counter-phase with the oscillations of the meteorological parameters. Periods of unusual cold or dryness, presumably resulting from underlying longer-term trends or oscillations (such as ENSO), affected the homogeneity of quasi-12-month flowering events, fruit maturation and also the production of germinable seeds. Some species show underlying quasi-2-year-oscillations, for example that synchronise with the development of air temperature; others reveal an underlying decrease or increase in flowering activity over the observation period, influenced for instance by solar irradiance. As Ecuador suffers the highest rate of deforestation in South America, there is an urgent need for indigenous plant material for reforestation. A detailed knowledge of the biology of reproduction in relation to governing external factors (mainly climate) is thus required.     

外来入侵植物意大利苍耳的传粉生态学特性

从开花物候、花粉活力、柱头可受性、传粉媒介、花粉散布距离、雄性、雌性功能以及繁育系统等方面系统地研究了意大利苍耳的传粉生态学特性,旨在为揭示该物种成功入侵的机理提供科学依据。研究结果显示:意大利苍耳种群的花期较长,雌花花期可达40d,雄花花期可达30d。其雄性和雌性功能都很强,意大利苍耳单株雄花序和花粉量分别高达3847个/株和37903037粒/株,单株雌花序和单株胚珠数高达3847个/株和7694枚/株,种群平均结实率高达80.59%,即平均每株个体能形成6200枚成熟的果实。花粉活力的日变化呈单峰型曲线,早晨和傍晚的花粉活力最低,14:00时的活力高达99%。雌蕊柱头一经伸出总苞即具有可受性,可受期长达7d,开花后的3—4d柱头的可受性最强。意大利苍耳为风媒传粉植物,在4.5km/h的微风条件下,其花粉的散布距离可达45m,大量的花粉集中分布在距花粉源0—20m的范围内,这对于往往高密度连片分布的意大利苍耳种群来说无疑是一种高效的传粉策略。意大利苍耳的交配系统灵活多样,盛花期自然结实率达到100%,套袋试验结果表明该植物自交亲和,自株自然授粉的结实率高达93%。表明较长的花期、大量的雌雄花序及花粉数量、较高的花粉活力、较长的柱头可授期、较远的花粉风媒散布距离、混合交配系统、以及较高的结实率是意大利苍耳繁殖成功的重要保障,也是其成功入侵的重要原因。     

桃红颈天牛成虫的交配行为及对不同寄主植物的栖落和产卵选择性

【目的】桃红颈天牛Aromia bungii是一种钻蛀性害虫,主要以蔷薇科果树和观赏树木为寄主。本研究旨在搞清楚桃红颈天牛成虫的交配产卵行为及其对不同寄主植物的栖落和产卵选择性。【方法】选择苹果Malus pumila、山樱花Prunus serrulata var. spontanea、桃Prunus persica、西府海棠Malus micromalus、毛樱桃Prunus tomentosa、秋子梨Pyrus ussuriensis、杏Prunus armeniaca和红叶李Prunus cerasifera var. atropurea 8种寄主植物木段,构建为成虫的活动场所,观察桃红颈天牛成虫的交配和产卵行为,并测定桃红颈天牛成虫在不同寄主木段上的栖落数量和产卵量。【结果】桃红颈天牛成虫的交配行为分为避让、打斗和交配3种行为。交配过程可分为相遇抱对、插入输精、交配后陪护3个阶段。桃红颈天牛雌虫产卵时无刻槽行为,喜欢将卵产于较为狭窄的缝隙中。雌虫偏好在桃树木段上栖落,雄虫则偏好在毛樱桃木段上栖落。李亚科李属的桃是桃红颈天牛最喜欢的产卵寄主木段,而雌虫在苹果亚科的苹果、海棠和秋子梨木段上产卵极少。相关性分析结果表明雌虫在寄主木段上的栖落数量与产卵量占比间存在正相关性,雄虫在寄主木段上的栖落数量与雌虫在寄主木段上的栖落数量之间也存在正相关性。【结论】桃红颈天牛雌成虫最喜欢桃树木段,雄虫对寄主植物的选择性与雌虫并不完全相同。     

DEMOGRAPHIC ASPECTS OF FLOWER AND FRUIT PRODUCTION IN BEAN PLANTS,PHASEOLUS VULGARIS L.

In this paper we assess the relative contributions to total pod yield of cohorts of flowers of known age. Bean plant (Phaseolus vulgaris L.) growth was monitored and 'births' of individual flowers were recorded and their fates followed. For every cohort of flowers an array of yield measures was calculated, including the mean and total cohort dry weight of fruits, and the numbers of viable and aborted seeds and completely undeveloped ovules. When mature, pods were removed from half of the plants. Plants whose mature pods were removed produced significantly greater numbers of flowers, ovules and viable seeds and a greater total weight of pods than did untreated plants. However, the proportion of ovules giving rise to viable seeds and to aborted seeds and undeveloped ovules was the same in plants of both groups. Approximately 50% of ovules of marketable pods, in all plants, yielded viable seeds. The 12 cohorts of flowers contributed markedly different amounts to plant yield. Early and later cohorts contributed much less than did middle cohorts in terms of marketable pod dry weight and numbers of fully developed seeds, aborted seeds, and undeveloped ovules. Peak productivity was therefore obtained from the middle phase of flowering in these annual bean plants. The lower pod yield of later cohorts is interpreted as a result of competition for limited resources between maturing pods and new flowers. We consider possible causes for the various fates of flowers and ovules and discuss flowers as plant modules suitable for demographic examination.