沙地中植物的传粉对策——毛乌素案例研究

地环境严酷多变，植物的物种多样性较低且常成斑块状分布，传粉昆虫的种类较少且其数量在时间和空间上变化较大。生长在沙地中的植物常采取一些独特的传粉对策来适应其严酷多变的的环境。然而，前人对沙生植物传粉生物学的研究还非常有限，至今未总结出其与沙生环境相适应的传粉生物学特点。本文以生长于毛乌素沙地的角蒿、披针叶黄华和羊柴三种主要蜜源植物为材料，通过大量的野外观察和人工实验，对其开花习性、花生物学特点、传粉和交配系统及有性繁殖成功进行了研究，得到以下主要结论： 1. 由于传粉昆虫的种类较少，传粉者的拜访频率低且时空分布不均，角蒿发育出一种有效的生殖保障机制——通过风驱动花冠脱落实现延迟自花授粉。在花开放后期，角蒿仍然张开且未萎蔫的花冠被风吹落，拖动冠生雄蕊沿花柱向前滑动，使得花药穿过张开的柱头裂片，从而实现自花传粉。然而在柱头裂片闭合后，由于柱头表面被封闭，使得这一自花传粉过程不能实现。该植物不存在雌雄异熟，并且是完全自交亲和的。与自然状况下相比，在风排除帐中花冠脱落的比例显著减小，说明风对角蒿花冠的脱落至关重要。在排除传粉者的情况下，花开放后期花冠的脱落显著增加了柱头上的花粉数，相应也显著增加了花的结实和结籽率，说明风驱动的花冠脱落可通过促进自交来增加果实和种子生产。仅散熊蜂和四条无垫蜂两种蜂为角蒿提供传粉，但在自然状况下样地中仅仅一半的花被拜访，并且去雄花的结实率显著低于未去雄的花，表明昆虫传粉仅对花的结实有部分贡献，并且延迟自花传粉能为角蒿提供繁殖保障。另外，角蒿的柱头较敏感，它的两个张开的柱头裂片在受到传粉者触碰后很快紧紧闭合，但是如果此时柱头表面未收到或仅收到少量花粉，闭合的柱头裂片又会重新张开，这一特性是一种动态的雌雄异位，能减少自交和异交的干扰，并且也是角蒿生殖保障机制的一部分。由于涉及花器官的运动及风和敏感柱头的积极参与，因此角蒿的延迟自花传粉与以前报道的自花传粉机制均不同。此外，角蒿由蜂类和风拖动花冠共同参与的传粉模式也是一种新的风和动物联合传粉机制。 2. 披针叶黄华在早春开花，植株密集，花量较大，由于在此期间开花的虫媒植物很少，此时出现的蜂类在食源有限的情况下频繁地拜访披针叶黄华的花。在传粉者频繁拜访的压力下，披针叶黄华采取了花药序次成熟和开裂的对策来实现花粉的最佳散布。披针叶黄华是自交亲和的，单花期长3-4天，它的十枚完全分离的雄蕊在2-3天内分五对序次地伸长和开裂，这五对花药序次开裂通常彼此间隔3-4小时，但这一行为在夜间停止。五对序次开裂花药的大小、花粉数量和可育花粉的比例无显著差异。有七种蜂为披针叶黄华提供传粉，其中山本氏切叶蜂是主要传粉者，它对披针叶黄华的拜访频率很高，一朵花平均每天受到25次拜访。山本氏切叶蜂在一次拜访中移出一朵花内可获得花粉的70-90%，且一次拜访和多次拜访对花粉的移取率无显著差异;在具有1-5对开裂花药的花间，昆虫对花粉的移取率也无显著差异，说明传粉者对花粉的移取能力很强。其花蜜的分泌模式与花药序次开裂的模式及传粉者的拜访特点相适应。正是传粉者高的拜访频率和强的花粉移取能力使得披针叶黄华采取了序次呈现花粉的策略来增加花粉被散布的机会。 3. 羊柴是毛乌素沙地的一种优势克隆植物，花期较长，开花量大。该种植物是不完全雄性先熟的，P/O比为55 610，花蜜的分泌模式与雌性功能的出现和昆虫的拜访相适应。有14种蜂为其提供传粉，其中意大利蜂、散熊蜂、白脸条蜂、和海切叶蜂是四种主要传粉者。该种植物具有自交异交混和交配系统，但自体受精结实率很低，种子繁殖需依靠传粉者提供授粉，传粉者的拜访行为能导致频繁的同株异花传粉。 4. 羊柴植株具有短时间大量开花的特点，但相邻不同斑块的开花时间差异很大，早、晚开花的斑块较少，大多数斑块一致在开花季节中期开花。这种斑块水平上开花的不同步能影响传粉者的种类组成和拜访频率，在整个开花季节内，传粉者的种类逐渐减少，并且早开花的斑块比那些中期开花的有更高的传粉者拜访频率,说明大量中期开花的斑块间存在传粉者竞争。在早开花的斑块中，传粉者的拜访频率与花展示的大小成显著正相关，说明大量开花能增加花展示对传粉者的吸引。但在中期开花的斑块中，随着开花量的持续增加，传粉者拜访减少。与中期开花的斑块相比，早、晚开花的斑块有更高的结实率。斑块水平上植株的结实率与羊柴的盖度、植株密度和重要值等种群统计学参数无显著相关，并且植株的大小和花展示的大小对开花时间与结实率的关系没有影响。由此可知，羊柴在斑块水平上大量开花和不同步开花共存的开花模式可能是对吸引传粉者和减少植株间对传粉者竞争的一种折衷。 5. 生长于毛乌素沙地的这三种主要蜜源植物的花期几乎不重叠，均具有泛化传粉系统，在花期内出现的大多数蜂类都可为披针叶黄华和羊柴提供传粉，比较特殊的是角蒿具有昆虫和风联合传粉的泛化传粉系统。三种植物都是自交亲和的，具有混合交配系统。这可能与沙地环境中蜂类的物种多样性较低，且其数量在时间和空间上变化很大有关。

玉米种子脱水耐性与其线粒体完整性及抗氧化系统的关系

本文以正常性玉米种子‘农大108’(Zea Mays L. ‘Nongda 108’) 种胚为实验材料，研究了玉米种子发育过程中脱水耐性的变化规律，细胞匀浆以及线粒体水平上活性氧清除酶活性与种子脱水耐性/敏感性的关系，以及线粒体结构和功能完整性在发育过程中不同阶段对脱水的应答，以期在亚细胞分区水平上，针对活性氧产生的源头位点 (线粒体) 探明种子细胞脱水耐性/敏感性与抗氧化系统运转的关系。结果表明： 玉米种子在发育过程中先获得萌发能力后获得脱水耐性，并且脱水耐性的获得是一个渐进的过程。人工授粉后26天 (Days after pollination, DAP) 之前的种胚不具有脱水耐性，26 DAP时开始获得脱水耐性，到34 DAP后种胚完全获得脱水耐性。 在发育过程中，种胚线粒体的呼吸速率逐渐降低，并且对脱水的敏感性也逐渐下降。脱水会降低脱水敏感性种胚线粒体的结构完整性；脱水同时会降低线粒体功能的完整性，包括线粒体能量产生的速率和效率，以及三羧酸循环关键酶的活性。但当种胚获得脱水耐性后，脱水将不再影响种胚线粒体结构和功能的完整性。 玉米种胚发育过程中脱水耐性的变化与细胞中的抗氧化系统有关。在细胞匀浆水平上，脱水过程中脂质过氧化产物的积累与细胞脱水耐性的关系不明显；但是在线粒体水平上脱水会明显导致脱水敏感性种胚线粒体膜质过氧化程度的升高。脱水导致脱水敏感种胚细胞中几个重要的抗氧化酶活性的下降，但是与细胞匀浆水平相比，在线粒体水平上抗氧化酶系统对脱水更加敏感。 总之，发育早期玉米胚对脱水之所以敏感有两方面的原因，一方面是发育早期线粒体具有较高的代谢速率因而产生过多的活性氧，另一方面是由于脱水导致各抗氧化酶活性的显著降低，失去了抗氧化保护功能。而在发育晚期，早期本来很活跃的许多代谢随之关闭，呼吸速率降到很低，因而产生的活性氧减少，同时由于抗氧化系统对脱水的耐受性，所以脱水不会对线粒体的结构和功能造成伤害。与细胞匀浆水平相比，线粒体水平上抗氧化系统的运转与种胚在发育过程中脱水耐性的获得的关系更加密切。

西双版纳热带雨林聚果榕小蜂的传粉生态学

对聚果榕小蜂 (Ceratosolensp )传粉生态学进行了首次研究。结果表明 ,聚果榕小蜂的雄蜂比雌蜂早羽化数小时 ;雌蜂羽化不能自行打开瘿花和果肉出蜂口 ,两个出蜂口均需雄蜂开凿。而聚果榕的成熟花粉 ,不能自行地从开裂处散发出来 ,必须经榕小蜂的繁殖性雌蜂采集才能散到表面。羽化后的雌蜂在开裂的雄花中不停地用触角柄节、口器上颚和足推动和采集花粉。雌蜂飞出熟榕果寻找嫩隐头花果 ,一般在外飞翔 5～ 80min。雌蜂进入嫩聚果榕的隐头花果内后 ,立即把粘附在足、头、触角和身上的花粉不停地推动到长柱头雌花中 ,授粉行为长达 4～ 9h。然后 ,才把卵产在短柱头雌花中

Diffusive coevolution and mutualism maintenance mechanisms in a fig-fig wasp system

In reciprocal mutualism systems, the exploitation events by exploiters might disrupt the reciprocal mutualism, wherein one exploiter species might even exclude other coexisting exploiter species over an evolutionary time frame. What remains unclear is how such a community is maintained. Niche partitioning, or spatial heterogeneity among the mutualists and exploiters, is generally believed to enable stability within a mutualistic system. However, our examination of a reciprocal mutualism between a fig species (Ficus racemosa) and its pollinator wasp (Ceratosolen fusciceps) shows that spatial niche partitioning does not sufficiently prevent exploiters from overexploiting the common resource (i.e., the female flowers), because of the considerable niche overlap between the mutualists and exploiters. In response to an exploiter, our experiment shows that the fig can (1) abort syconia-containing flowers that have been galled by the exploiter, Apocryptophagus testacea, which oviposits before the pollinators do; and (2) retain syconia-containing flowers galled by Apocryptophagus mayri, which oviposit later than pollinators. However, as a result of (2), there is decreased development of adult non-pollinators or pollinator species in syconia that have not been sufficiently pollinated, but not aborted. Such discriminative abortion of figs or reduction in offspring development of exploiters while rewarding cooperative individuals with higher offspring development by the fig will increase the fitness of cooperative pollinating wasps, but decrease the fitness of exploiters. The fig fig wasp interactions are diffusively coevolved, a case in which fig wasps diversify their genotype, phenotype, or behavior as a result of competition between wasps, while figs diverge their strategies to facilitate the evolution of cooperative fig waps or lessen the detrimental behavior by associated fig wasps. In habitats or syconia that suffer overexploitation, discriminative abortion of figs or reduction in the offspring development of exploiters in syconia that are not or not sufficiently pollinated will decrease exploiter fitness and perhaps even drive the population of exploiters to local extinction, enabling the evolution and maintenance of cooperative pollinators through the movement between habitats or syconia (i.e., the metapopulations).

Pollinator shift and reproductive performance of the qinghai-tibetan plateau endemic and endangered Swertia przewalskii (Gentianaceae)

Reproductive failure results in many plant species becoming endangered. However, little is known of how and to what extent pollinator shifts affect reproductive performance of endangered species as a result of the artificial introduction of alien insects. In this study we examined breeding systems, visitor species, visiting frequency and seed set coefficients of Swertia przewalskii in two years that had different dominant pollinator species (native vs. alien). Flowers of this species were protandrous and herkogamous and insects were needed for the production of seeds. The stigmatic receptivity of this species was shorter than for other gentians. No significant difference in seed set coefficient was found for hand-pollinated plants between the two years, indicating that pollinator shift only had a minor effect on this plant's breeding system. The commonest pollinators in 2002 were native bumblebees, alien honeybees and occasional solitary bees, however, only alien honeybees were observed in 2004. The flower visitation rate in both years was relatively high, although the total visit frequency decreased significantly in 2004. The control flowers without any treatment produced significantly fewer seed sets in 2004 than in 2002. In the past decade the seed production of this species may have partly decreased due to pollination by alien honeybees, however, we suggest that they might have acted as alternative pollinators ensuring seed production of S. przewalskii when native pollinators were unavailable. The main reason that this plant is endangered is probably the result of habitat destruction, but changes in land use, namely intensified agricultural practice and unfavorable animal husbandry have also contributed to its decline. We recommend that in-situ conservation, including the establishment of a protected area, is the best way to preserve this species effectively.

Reproductive ecology of the Qinghai-Tibet Plateau endemic Gentiana straminea (Gentianaceae), a hermaphrodite perennial characterized by herkogamy and dichogamy

The combined occurrence of both herkogamy and dichogamy in a hermaphrodite species has been considered to strongly favour outcrossing. In this study, we investigated in detail the reproductive ecology of Gentiana straminea Maxim. (Gentianaceae), a hermaphrodite perennial endemic to the Qinghai-Tibet Plateau. In a series of observations and experiments over four consecutive years, we examined whether the combination of dichogamy and herkogamy in individual flowers completely prevents geitonogamous pollen transfer in this species. The mode of floral development clearly indicates that autonomous self-pollination is completely avoided through herkogamy and dichogamy in individual flowers. This implication was confirmed by the breeding experiments, since no seed was produced when flowers were isolated. However, this gentian proved to be highly self-compatible when geitonogamous selfing was artificially induced. Many flowers opened simultaneously on individual plants, the ratio of male to female phase flowers was close to 2:1 in each inflorescence, at the full anthesis phase, and they were randomly distributed amongst the upper, middle and lower parts of each stem's inflorescence. On average, Bombus sushikini Skorikov, the most frequent visitor and only legitimate pollinator of G. straminea, visited nearly two flowers per inflorescence, and four flowers per plant. Among the pollinators' foraging bouts, the proportions of geitonogamous visits to inflorescences or flowers within an individual plant were 29% and 37%, respectively. Therefore, despite the strict dichogamous and herkogamous characteristics of the individual flowers, geitonogamous selfing might still prevail in G. straininea because of the size of its floral displays and the continuous visiting behavior of B. sushkini. (C) 2005 Elsevier SAS. All rights reserved.

Molecular evidence for the sister relationship of the eastern Asia-North American intercontinental species pair in the Podophyllum group (Berberidaceae)

The presumed pair relationships of intercontinental vicariad species in the Podophyllum group (Sinopodophyllum hexandrum vs. Podophyllum pelatum and Diphylleia grayi vs. D. cymosa) were recently, considered to be paraphyletic. In the present paper, the trnL-F and ITS gene sequences of the representatives were used to examine the sister relationships of these two vicariad species. A heuristic parsimony analysis based on the trnLF data identified Diphylleia as the basal clade of the other three genera, but provided poor resolution of their inter-relationships. High sequence divergence was found in the ITS data. ITS1 region, more variable but parsimonyuninformative. has no phylogenetic value, Sequence divergence of the ITS2 region provided abundant, phylogenetically informative variable characters. Analysis of ITS2 sequences confirmeda sister relationship between the presumable vicariad species, in spite of a low bootstrap support for Sinopodophyllum hexandrum vs. Podophyllum pelatum. The combined ITS2 and trnL-F data enforced a sister relationship between Sinopodophyllum hexandrum and Podophyllum pelatum with an elevated bootstrap support of 100%. Based on molecular phylogeny, the morphological evolution of this group was discussed. The self-pollination might have evolved from cross-fertilization two times in this group. The different pollination and seed dispersal systems of Sinopodophyllum hexandrum and Podophlyllum pelatum resulted from their adaptations to different ecological habitats. The divergence time of Sinopodophyllum hexandrum-Podophyllum pelatum is estimated to be 6.52+/-1.89 myr based on the ITS divergence. The divergence of this species pair predated or co-occurred with the recent uplift of the Himalayas 4-3 myr during the late Miocene and the formation of the alpine habitats. Sinopodophyllum hexandrum developed a host of specialized characters in its subsequent adaptation to the arid alpine surroundings. The present study confirmed the different patterns of species relationship between Asian-North American disjuncts. The isolation of plant elements between North America and eastern Asia must have been a gradual process, resulting in the different phylogenetic patterns and divergence times of the disjuncts.

Recomendacoes para a frutificacao da abobora hibrida tipo Tetsukabuto: uso de polinizadores e reguladores de crescimento de plantas.

1999

Fitness of hybrids between rapeseed (Brassica napus) and wild Brassica rapa in natural habitats

J. Allainguillaume, M. Alexander, J. M. Bullock, M. Saunders, C. J. Allender, G. King, C. S. Ford, M. J. Wilkinson. (2006). Fitness of hybrids between rapeseed Brassica napus and wild Brassica rapa in natural habitats. Molecular Ecology, 15 (4) 1175-1184. RAE2008

Invertebrate diversity in Irish and British forests

Ireland and Britain were once covered in natural forest, but extensive anthropogenic deforestation reduced forest cover to less than 1% and 5 %, respectively, by the beginning of the 20th century. Large-scale afforestation has since increased the level of forest cover to 11% in Ireland and 12% in Britain, with the majority of planted forests comprising small monoculture plantations, many of which are of non - native conifer tree species. At present the forest cover of Ireland and Britain generally consists of small areas of remnant semi-natural woodland and pockets of these plantation forests within a predominantly agricultural landscape. Invertebrates comprise a large proportion of the biodiversity found within forested habitats. In particular, spiders and carabid beetles play an important role in food webs as both predators and prey and respond to small-scale changes in habitat structure, meaning they are particularly sensitive to forest management. Hoverflies play an important role in control and pollination and have been successfully used as indicators of habitat disturbance and quality. This research addressed a number of topics pertinent to the forest types present in the contemporary Irish and British landscapes and aimed to investigate the invertebrate diversity of these forests. Spiders and carabid beetles were sampled using pitfall trapping and hoverflies were sampled using Malaise net trapping. Topics included the impacts of afforestation, the importance of open space, the choice of tree species, and the use of indicators for biodiversity assessment, as well as rare native woodlands and the effect of grazing on invertebrate diversity. The results are discussed and evidence-based recommendations are made for forest policy and management to protect and enhance invertebrate biodiversity in order to promote sustainable forest management in Ireland and Britain.

OPERA IN AMERICA: MUSIC OF, BY, AND FOR THE PEOPLE­: A PERFORMANCE DISSERTATION

Opera in America: Music of, by, and for the people is a study of the relationship between American popular culture and opera in the United States. Four performance projects demonstrate the on-going exchange between the operatic community-including its composer, singers, and patrons-and the country's popular entertainment industry with its broad audience base. Numerous examples of artistic cross pollination between lowbrow and highbrow music will illustrate the artistic and social consequences created by this artistic amalgamation. Program #1, By George! By Ira! By Gershwin!, is a retrospective of Gershwin's vocal music representing a blending of popular and serious music in both style and form. The concert includes selections from Porgy and Bess, a work considered by many musicologists as the first American opera. Program #2, Shadowboxer, is a premiere performance of an opera by Frank Proto and John Chenault. For this newly commissioned work, I serve as Assistant Director to Leon Major. Shadowboxer provides a clear example of opera utilizing popular culture both musically and dramatically to tell the true story of American hero and legendary boxer, Joe Louis. Program #3, Just a Song at Twilight, is an original theatrical music piece featuring music, letters, diaries, and journals of the Gilded Age, an era when opera was synonymous with popular entertainment. Special attention is focused on tum-of­ the-century singers who performed in both opera and vaudeville. Program #4 is a presentation of Dominick Argento's Miss Manners on Music and illustrates the strong relationship that can exist between opera and American popular entertainment. Originally conceived as a song cycle, I have staged the work as a one-act opera sung and acted by soprano Carmen Balthrop. This piece is based on the writings of pop icon and newspaper columnist Judith Martin, otherwise known as Miss Manners. All four performances are recorded in audio and video formats.

Range-edge effects promote clonal growth in peripheral populations of the one-sided wintergreen Orthilia secunda

Existing in suboptimal conditions is a frequent occurrence for species inhabiting the cusp of their ecological range. In range-edge populations of plants, the scarcity of suitable habitat may be reflected in small population sizes which may result in increased self-pollination and/or inbreeding and an increase in the incidence of clonal reproduction. These factors may result in a decrease in levels of genetic diversity and a loss of potential adaptive variation that may compromise species' ability to cope with changes in their environment, an issue that is particularly relevant today with the current concern surrounding global climate change and its effect on species' distributional ranges. In the present study, we have compared the levels of clonal reproduction in the one-sided wintergreen Orthilia secunda (L.) House in (1) populations from its main continuous distribution range, (2) populations occurring on the limits of the continuous range, and (3) peripheral populations outwith the species' continuous distribution range. Range-edge populations in Scotland and Sweden displayed significantly lower genotypic richness and diversity than those from the main area of the species' distribution in these countries. Populations from Ireland, which occur in the temperate zone rather than the boreal conditions that are the preferred habitat for the species, and which represent relict populations left over from cooler periods in the Earth's history, displayed no within-population genetic diversity, suggesting a complete lack of sexual reproduction. Furthermore, the genetic distinctiveness of the Irish populations, which contained alleles not found in either the Scottish or the Swedish populations, highlights the value of 'trailing edge' populations and supports the concept of 'parochial conservation', namely the conservation of species that are locally rare but globally common.

Cryptic species diversity in a widespread bumble bee complex revealed using mitochondrial DNA RFLPs

Cryptic species diversity is thought to be common within the class Insecta, posing problems for basic ecological and population genetic studies and conservation management. Within the temperate bumble bee (Bombus spp.) fauna, members of the subgenus Bombus sensu stricto are amongst the most abundant and widespread. However, their species diversity is controversial due to the extreme difficulty or inability morphologically to identify the majority of individuals to species. Our character-based phylogenetic analyses of partial CO1 (700 bp) from 39 individuals spread across their sympatric European ranges provided unequivocal support for five taxa (3-22 diagnostic DNA base pair sites per species). Inclusion of 20 Irish specimens to the dataset revealed >= 2.3% sequence divergence between taxa and 200 m) whilst B. cryptarum was relatively more abundant at higher altitudes. Bombus magnus was rarely encountered at urban sites. Both B. lucorum and B. terrestris are nowadays reared commercially for pollination and transported globally. Our RFLP approach to identify native fauna can underpin ecological studies of these important cryptic species as well as the impact of commercial bumble bees on them.

Large-Range Movements of Neotropical Orchid Bees Observed via Radio Telemetry

Neotropical orchid bees (Euglossini) are often cited as classic examples of trapline-foragers with potentially extensive foraging ranges. If long-distance movements are habitual, rare plants in widely scattered locations may benefit from euglossine pollination services. Here we report the first successful use of micro radio telemetry to track the movement of an insect pollinator in a complex and forested environment. Our results indicate that individual male orchid bees (Exaerete frontalis) habitually use large rainforest areas (at least 42-115 ha) on a daily basis. Aerial telemetry located individuals up to 5 km away from their core areas, and bees were often stationary, for variable periods, between flights to successive localities. These data suggest a higher degree of site fidelity than what may be expected in a free living male bee, and has implications for our understanding of biological activity patterns and the evolution of forest pollinators.