Reproductive patterns within racemes in protandrous Aconitum gymnandrum (Ranunculaceae): potential mechanism and among-family variation

The adaptive significance and mechanism of patterns in floral sex allocation and female success within inflorescences has attracted attention recently, whereas few studies have examined genetic variation of intra-inflorescence pattern. The purpose of this study is to investigate patterns of reproduction within racemes in protandrous Aconitum gymnandrum Maxim., and illuminate potential mechanisms and genetic variation of such patterns. Data from pot experiment on 40 maternal families were collected in field. Anther number, pollen:ovule ratio and seed germination rate increased from bottom to top flowers within racemes, but other traits, such as gynoecium mass, carpel number, sepal galea height and seed production decreased significantly with flowering sequence. Variation in floral sex allocation within racemes in A. gymnandrum fitted entirely the prediction of protandry, which was not a result of architectural effect. Such selected pattern may result from a variety of factors influencing the mating environment, such as pollinator directionality, display size and flower longevity. Decline of female success within racemes in A. gymnandrum also resulted from male-biased allocation selected by variation in the mating environment, not resource competition or pollen limitation. Moreover, there was genetic variation for most reproductive traits and the position effect, as evinced by significant variation among families.     

Sexual interference of the floral kind

Floral hermaphroditism results in conflicts and compromise in the parental roles of plants during pollination and mating. A potential cost of hermaphroditism is sexual interference between maternal and paternal functions resulting in gamete wastage and reduced fitness. Sexual interference may or may not be associated with self-pollination. In cases where self-pollination occurs, ovule or pollen discounting may reduce mating opportunities. Here I describe forms of sexual interference in flowering plants, distinguishing whether physical or biochemical interactions are involved and whether fitness costs associated with gamete wastage arise from intra-floral versus inter-floral processes. I review the limited experimental evidence for interference between sex functions and evaluate the hypothesis that some floral adaptations usually interpreted as anti-selfing mechanisms may serve an alternative function in reducing mating costs arising from this form of sexual conflict.     

Influence of northern limit range on genetic diversity and structure in a widespread North American tree,sugar maple (Acer saccharum Marshall)

Due to climate change, the ranges of many North American tree species are expected to shift northward. Sugar maple (Acer saccharum Marshall) reaches its northern continuous distributional limit in northeastern North America at the transition between boreal mixed‐wood and temperate deciduous forests. We hypothesized that marginal fragmented northern populations from the boreal mixed wood would have a distinct pattern of genetic structure and diversity. We analyzed variation at 18 microsatellite loci from 23 populations distributed along three latitudinal transects (west, central, and east) that encompass the continuous–discontinuous species range. Each transect was divided into two zones, continuous (temperate deciduous) and discontinuous (boreal mixed wood), based on sugar maple stand abundance. Respective positive and negative relationships were found between the distance of each population to the northern limit (D_north), and allelic richness (A_R) and population differentiation (F_ST). These relations were tested for each transect separately; the pattern (discontinuous–continuous) remained significant only for the western transect. structure analysis revealed the presence of four clusters. The most northern populations of each transect were assigned to a distinct group. Asymmetrical gene flow occurred from the southern into the four northernmost populations. Southern populations in Québec may have originated from two different postglacial migration routes. No evidence was found to validate the hypothesis that northern populations were remnants of a larger population that had migrated further north of the species range after the retreat of the ice sheet. The northernmost sugar maple populations possibly originated from long‐distance dispersal.     

Multi-cycle synchronous protandry in raceme-like inflorescences of a bumblebee-pollinated herb Aconitum grossedentatum

Multi-cycle synchronous dichogamy is expected to be a mechanism for reducing self-pollination and sexual interference. It is often found in plants with umbellate inflorescences where pollinator movement is unpredictable, but not in plants with raceme inflorescences that are pollinated by bumblebees. Plants with raceme inflorescences often acropetally open flowers, resulting in an arrangement of females at lower level and males at upper level. This is good enough to preclude geitonogamy because bees tend to move upwardly within the inflorescences. Furthermore, although the degree of segregation of sexes varies among species, their intraspecific variations within a population have rarely been examined. Here, we present a synchronous protandry in bee-pollinated Aconitum grossedentatum, which has a raceme-like inflorescence and opens flowers basipetally. To evaluate the functional significance of synchronous dichogamy in mating, we firstly observed the distribution of sex phases of open flowers. Then, we assessed the effect of each phase flower on foraging behavior by pollinators and seed-set success. The inflorescences tended to exhibit either male- or female-phase flowers at any moment early in the flowering season, but the degree of segregation of sexes declined over time within a population. The degree of the segregation did not affect bumblebee visits to flowers, but it decreased seed-set success of female-phase flowers at that time. Our results demonstrated that synchronous protandry was beneficial for pollination success in A. grossedentatum by avoiding geitonogamy. Nevertheless, we also found asynchronous protandry late in the season, suggesting that the benefits by synchronous protandry decreased over the season.

     

Estimating the costs of allocation to male and female functions in a monoecious cucurbit,Lagenaria siceraria

Costs of allocation to male versus female functions were determined for the monecious, annual vine Lagenaria siceraria by removing all flower buds of a given gender and using the additional vegetative growth as a measure of the cost of allocation to that gender (following methods of Silvertown 1987). In this cucurbit, we found significant costs associated with male flower production and fruit production, but not with female flower production. These results are not surprising given the strongly male-biased floral ratio (20 male:1 female) and the large gourds of this species. However, our results are in contrast to Silvertown's study of Cucumis sativus which found no cost to male funtion. In addition, our treatments significantly affected floral sex expression, resulting in increased femaleness for treatment plants. This increase is a consequence of increased lateral branch initiation in these plants and the almost exclusive production of female flowers on lateral branches. Fruit production was associated with a shut-down of subsequent flower production. Given that plant architecture leads to pronounced protandry (on average, 17 male flowers bloom before the first female flower) and that male costs are associated with flower production and the bulk of female costs with fruit production, this shutdown effectively separates male and female functions temporally. We agree with Silvertown's conclusion that this form of protandry is an important factor in the maintenance of cosexuality.     

Dragma Hippomanicum VIII. A new species of Astragalus section Strigulosi (Leguminosae) from Oaxaca

Astragalus cenorrhynchus Barneby is related toA. oxyrrhynchus Hemsl., endemic to the Valley of Mexico, but differs in diminished stature, few-flowered racemes of yellowish flowers, and plumper, apically unilocular pod.     

云南草寇花序内不同部位的性分配

两性花植物花序内的性分配常存在差异,资源竞争、结构效应、交配环境(雌雄异熟、传粉者定向访花行为等)或授粉不均匀等几种假说可以解释这种现象。为验证上述假说,该研究以云南草寇两种表型(雄先熟型和雌先熟型)为材料,分析了其花序内不同部位(基部、中部和顶部)的每花花粉数、胚珠数、花粉/胚珠比、结实率和结籽率,花序内传粉者的定向访花行为,以及人工辅助授粉和去花处理对结实率和结籽率的影响。结果表明:两种表型花序内每花花粉数不随部位而变化,每花胚珠数、结实率和结籽率由基部到顶部依次降低,每花花粉/胚珠比由基部到顶部依次增加,表明顶部花存在偏雄的性分配。人工辅助授粉后,结实率、结籽率仍由基部到顶部依次降低,表明授粉不均匀假说不能解释云南草寇花序内不同部位结实率、结籽率的差异。去除基部和中部花后,顶部花人工辅助授粉条件下的结实率、结籽率与基部花人工辅助授粉条件下的结实率、结籽率无差异,表明云南草寇花序内不同部位结实率、结籽率的差异主要由资源竞争引起。雌先熟表型每花花粉数、花粉/胚珠比高于雄先熟表型,表明两种表型存在性分配差异。传粉者主要先访问云南草寇基部的花,然后向顶部移动。云南草寇花序内顶部偏雄的性分配可能是由资源竞争和传粉者定向访花造成的。     

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.     

Inflorescence patterns in the woody Brazilian genus Diplusodon (Lythraceae)

The Brazilian genus Diplusodon is the second largest genus within Lythraceae. Their 85 species occupy diverse habitats within the ‘cerrado’ vegetation, and range from shrubs and treelets to dwarf, xylopodium-bearing subshrubs. A comparative-morphological survey of their inflorescence structures using Trollian typology is here presented, as well as some evolutionary considerations drawn from mapping inflorescence characters onto a preliminary phylogeny. The inflorescences of Diplusodon are mostly polytelic, ranging from single racemes to more or less complex double-, triple-, and multiple-racemes. Frondose, compound racemes are plesiomorphic within the genus. Nevertheless, an array of derived features has been found among their species, including production of lateral cymes, proliferation of the main axis, diverse patterns of internode elongation, reduction of subtending leaves to bracts, development of accessory branches, paedomorphic flowering, and, in three species, reversion to monotely.     

Effects of experimental manipulation of inflorescence size on pollination and pre-dispersal seed predation in the hummingbird-pollinated plant Ipomopsis aggregata

Large floral displays should theoretically provide advantages to plants through increased pollinator visitation and resulting fruit and seed set. However empirical tests of the response of pollinators to floral display size have been limited by a lack of direct experimentation, and the results of such studies have been equivocal. In addition, other selective agents such as pre-dispersal seed predators might modulate effects of floral display on pollination. By artificially altering flower number, we examined the direct effects of floral display in the monocarpic herb, Ipomopsis aggregata (Polemoniaceae), on visitation rates by broad-tailed and rufous hummingbird pollinators, as well destruction of fruits by a pre-dispersal seed predator (Hylemya: Anthomyiidae). In addition, we quantified the ultimate effects of flower number on female reproductive success. Plants with larger floral displays were most likely to be visited first in any given foraging bout (P < 0.01). As expected, plants with more flowers received more total flower visits. However, we found no gain in the proportion of flowers visited for many- versus few-flowered plants, or the total number of approaches/hour. In fact, a significantly greater percentage of flowers were visited on few-flowered plants. Plants did not compensate for our reduction in flowers by increasing investment in the number or proportion of flowers that set fruit, the number of seeds/fruit, or seed weight. Pre-dispersal seed predation was greater for many- than for few-flowered plants (P < 0.001), but this did not offset the potential fitness gains of producing large displays. Our data support the hypothesis that large floral displays function primarily in long-distance attraction of pollinators, and enhance maternal success. Received: 21 March 1996 / Accepted: 24 October 1996     

The spatial dynamics of pollen beetles in relation to inflorescence growth stage of oilseed rape: implications for trap crop strategies

Semi‐field‐scale arrays of oilseed rape (Brassica napus L.) (Brassicaceae) plants were used to observe the development of distributions of pollen beetles (Meligethes aeneus Fabricius) (Coleoptera: Nitidulidae) in a simulated trap crop system where inflorescence growth stage alone was used to manipulate the pest. Over two successive years, pairs of 1 m spaced square arrays of 100 glasshouse‐grown plants were placed 40 m apart in the field in May, and were subject to natural infestation by pollen beetles. The test plot of each pair had a simulated trap crop, with an outer row of plants at early flowering stage intended to protect more susceptible inner plants at late bud stage, and the control plot had all plants at the late‐bud stage, simulating a standard crop situation. Pollen beetles were counted daily on each plant for 10–13 days. The spatio‐temporal development of plot infestation was analysed in relation to the distribution of racemes in bud and raceme in flowers using Spatial Analysis by Distance IndicEs (SADIE), and tests of edge and centre distribution. Inflorescence growth stage characteristics were shown to be important in determining the spatial distributions of pollen beetles. In control plots, the numbers of racemes in bud and in flower were never edge or centre distributed. In test plots, racemes in flower were always edge distributed, and racemes in bud began edge distributed and became centre distributed. Pollen beetle numbers were usually spatially associated with the abundance of racemes in bud and/or in flower. In control plots, pollen beetles were neither edge nor centre distributed, but in test plots they maintained a significant edge distribution for 7–10 days. At the end of the experiments, females were more centre distributed in the test plots than males, and were more closely associated with racemes with buds, whereas males were more associated with racemes with flowers. In early flowering stage plants, the number of racemes in flowers were a good indicator of the abundance of racemes in buds, but this relationship was lost as flowering progressed. Although flowering racemes provide strong cues for immigrating pollen beetles, the abundance of buds may be a more important determinant of residence time, particularly for females, and is therefore a critical determinant of trap crop effectiveness.     

Cold hardiness in various organs and tissues of Rhododendron species and the supercooling ability of flower buds as the most susceptible organ

The freezing resistance of various organs and tissues was determined in 24 Rhododendron species (mainly Subgenus Tsutsutsi) having different ecological distributions. The order of hardiness for organ or tissue is as follows: leaf bud > wood ≧ bark > flower bud, and the flower bud is characterized as the most cold-susceptible organ. The relationship of killing temperature (KT) to northern distribution was the most significant in leaf buds compared to other organs and tissues. KTs of leaf buds for the most hardy species were ?45 °C (or below) and those for the most tender species were about ?23 °C, while KTs of flower buds were about ?28 °C for the former and ?16 °C for the latter. Although KTs of flower buds native to southwestern Japan were well correlated with the exothermic temperature distribution (ETD) of florets, those in the more northern species were generally lower than ETDs. The supercooling ability of flower buds appears to be sufficient to avoid the freezing stress since the extreme minimum temperature (EMT) at the northern limit of natural distribution for each tree species examined was not lower than the KT and ETD of the flower buds.     

When to estimate sex ratio in natural populations of insects? A study on sex ratio variations of gall midges within a generation

Precise estimation of arthropods' sex ratio is an important issue in a wide range of ecological studies and biological control programs. Although, in many cases changes in arthropods' sex ratio may be under the control of parents or some symbiotic microorganisms, biased sex ratios in some other species are caused by some extrinsic factors, neglect of which may lead to under/overestimation of true sex ratio. In this paper, we pursued those factors that cause false estimation of sex ratio in insects' species. We studied the predatory gall midge, Aphidoletes aphidimyza Rondani (Diptera: Cecidomyiidae), an important biological control agent of aphids, that shows protandry (i.e. early male emergence), differential lifespan of sexes, and differential distribution of sexes across habitat. Ten populations of A. aphidimyza were released separately in transparent cages and their sex ratio variations were recorded every 12 hours. The primary sex ratio in this species seems to be slightly male‐biased (52.41% males), however early emergence of males biases the sex ratio up to 72% males in a few hours after emergence. Shortly after the emergence of females, the sex ratio reaches its primary situation, but as a result of male‐biased mortality after mating, the proportion of females increases gradually to 97% by the fourth and fifth days after emergence. These results explicitly suggest that direct estimation of sex ratio in natural populations may be affected by some secondary factors such as differential mortality of sexes, protandry, and differential distribution of males and females over time and/or across habitat.     

上海大金山岛植被功能组成对草本植物、土壤动物和细菌多样性的影响

揭示植物群落对土壤动物和微生物多样性的上行调控效应有助于理解不同营养级生物多样性的维持机制。以往的研究主要集中在植物物种多样性对土壤动物和微生物多样性的影响,而关于植被功能组成自下而上的影响研究较少。以上海大金山岛13个植物群落为对象,在分析落叶木本植物占比与树木和草本物种多样性,以及土壤动物和细菌多样性的关系基础上,利用结构方程模型区分了落叶木本植物占比对土壤动物和细菌多样性的直接与间接影响效应。结果显示：落叶木本植物占比不仅分别对草本物种多样性和土壤动物多样性分别产生直接正效应和负效应（P<0.01;P<0.05）,也会通过草本物种的级联效应间接的降低地下土壤细菌多样性（P<0.10）。然而,木本植物多样性仅与草本物种多样性显著正关联（P<0.10）,与土壤动物和细菌多样性无显著关联（P>0.10）。该研究结果表明,相较于木本植物多样性,落叶植物占比在中亚热带北缘森林生态系统不同营养级生物多样性维持格局中扮演着更为重要的角色。     

Pattern of Flower and Fruit Production in Stryphnodendron adstringens, an Andromonoecious Legume Tree of Central Brazil

Abstract: Patterns of flower and fruit production in racemes of Stryphnodendron adstringens, an andromonoecious Brazilian savanna tree species, were studied in two natural areas near Uberlândia-MG. Racemes were divided in three parts: apex, centre, and base. Number of flowers, gender, and nectar and pollen production were analyzed for each section. Frequency of visitors to each part of the inflorescence was also quantified. Hand self- and cross-pollinations were performed in complete racemes and fruit set used to determine breeding system. The racemes produced a mean of 329 flowers, more densely packed in the central portion. Hermaphrodite and male flowers occur along the inflorescence but hermaphrodite flowers are more common in the centre. Fruit set was markedly low but does not seem to be limited by pollination service, since free open-pollinated racemes and hand cross-pollinated ones do not differ in fruit production rates. Fruits resulted mostly from cross-pollinated flowers and fruit production was biased to the central portion of the raceme. Nectar yield was higher in the central portion of the raceme and visitors arrived more commonly on this portion of the inflorescence. However, most flowers did not produce nectar. The pattern of fruit production seems to be a consequence of the hermaphrodite flower distribution in the raceme and it is not constrained by pollen flow or flower opening sequence.     

四川省福建柏林的初步研究

本文报道了福建柏(Fokienia hodginsii)的地理分布和以它为优势种的森林植被在四川省的生态条件、群落结构及其经营措施。福建柏属我国第一批二类重点保护植物。四川是它分布的最北边界。一般散生于亚热带常绿阔叶林之中,以它为优势种成林者较少。四川省江津县四面山福建柏林为一典型代表。笔者对它进行了初步的研究并建议加强保护和发展。     

Sex‐related variation in migration phenology in relation to sexual dimorphism: a test of competing hypotheses for the evolution of protandry

Timing of arrival/emergence to the breeding grounds is under contrasting natural and sexual selection pressures. Because of differences in sex roles and physiology, the balance between these pressures on either sex may differ, leading to earlier male (protandry) or female (protogyny) arrival. We test several competing hypotheses for the evolution of protandry using migration data for 22 bird species, including for the first time several monochromatic ones where sexual selection is supposedly less intense. Across species, protandry positively covaried with sexual size dimorphism but not with dichromatism. Within species, there was weak evidence that males migrate earlier because, being larger, they are less susceptible to adverse conditions. Our results do not support the ‘rank advantage’ and the ‘differential susceptibility’ hypotheses, nor the ‘mate opportunity’ hypothesis, which predicts covariation of protandry with dichromatism. Conversely, they are compatible with ‘mate choice’ arguments, whereby females use condition‐dependent arrival date to assess mate quality.     

Spectral polymorphisms in angiosperm flowers determined by differential ultraviolet reflectance

The ultraviolet reflectance pattern of flowers from 54 species representing 22 families, many native or introduced to Japan, were surveyed. Their differential ultraviolet patterns, i.e. spectral polymorphisms, were recorded by paired photographs of fresh flowers: one taken in full sunlight and the other through a visible spectrum absorbing filter. The different types of polymorphisms observed varied from total absorption by the whole flower to limited absorption by selected floral parts, such as corolla bases (nectar guides) or anthers. Some visible spectrum polymorphisms, e.g. inAnemone andHepatica, were not associated with different ultraviolet patterns. In three species ofIris, a distinct ultraviolet absorbing landing spot and nectar, tunnel guide as well as reflecting anthers are reported. The downward exerted anthers ofSolanum nigrum which shed pollen through bee wing vibration were ultraviolet absorbent. The implications of these ultraviolet patterns for biochemical (flavonoid), taxonomic, pollination and eco-evolutionary research are discussed. Attention is drawn to the seasonal distribution of ultraviolet radiation in the northern hemisphere and its selective role on northern floras.     

Factors controlling fruit set in hermaphroditic plants: Studies with the Australian proteaceae

The number of fruits produced by many hermaphroditic plants is usually far fewer than the number of flowers available for fertilization. There are various possible explanations for the low fruit:flower ratio, some proximate and others ultimate. Recent studies, especially in northern hemisphere systems, have used field experiments to test some of them, but there are potential difficulties with the methodology of some experiments and with the testing of ultimate hypotheses. It is important to examine the possible explanations for low fruit: flower ratios with a range of different systems. This article reviews studies on Australian species of woody, perennial shrubs in the family Proteaceae; this evolutionarily distinct group of plants and pollinators has several unusual and interesting characteristics, and provides a valuable addition to the better-known northern hemisphere studies.     

克钦楼梯草, 缅甸荨麻科一新种

该文描述了自缅甸北部发现的荨麻科楼梯草属一新种,克钦楼梯草。此种在体态上与骤尖楼梯草甚为相似,与后者的区别在于本种的每一茎节具正常叶和一退化叶,托叶狭披针状条形和无脉,雌总苞苞片无角状突起,雌小苞片较大,呈楔状长圆形,雌花具一小花被片,以及雌蕊具一宽倒卵球形柱头。