黑面神雌花气味传粉前后的变化及其生态意义

该文以与头细蛾传粉相关的叶下珠科植物黑面神为材料,采用动态顶空吸附法分别收集了该物种雌花传粉前后释放的挥发物,利用气相色谱-质谱联用技术分析了其化学成分,探索了三者间花气味的异同,并对其生态意义进行了讨论。结果表明:从该物种传粉前、传粉期和传粉后的雌花气味中共分离出22种挥发物(传粉前12种,传粉期15种和传粉后13种),以脂肪族衍生物和萜类物质为主。3-己烯醛、反-2-己烯醛、顺-3-己烯-1-醇、顺-乙酸-3-己烯酯、反-β-罗勒烯、β-石竹烯、β-蛇麻烯、反,反-α-金合欢烯和长叶烯是其主要挥发物。传粉前后雌花气味的化学种类及组成发生了显著分化,且这种变化主要是由于各时期主要挥发物发生改变而引起的。反-β-罗勒烯在传粉期含量最高,而在传粉后显著下降,3-己烯、反-2-己烯醛、顺-3-己烯-1-醇和顺-乙酸-3-己烯酯在传粉前含量均最高,β-石竹烯含量在传粉后显著升高,且长叶烯仅在传粉后出现。在传粉期和传粉后β-蛇麻烯和反,反-α-金合欢烯的含量均显著上升。同时,传粉期雌花的释放量明显高于传粉前和传粉后的。综上结果显示,黑面神雌花传粉前后花气味发生了质与量的改变。初步推测黑面神雌花...     

叶下珠头细蛾与小果叶下珠互利共生的研究--生物学及系统稳定性

【目的】为了确定小果叶下珠 Phyllanthus microcarpus 与叶下珠头细蛾 Epicephala sp.之间专性互利共生系统稳定性的增强是否主要通过彼此之间相互遏制对方对自身进行过度开采。【方法】通过对小果叶下珠上共生的叶下珠头细蛾形态及生物学的研究,记述了叶下珠头细蛾专性寄生寄主植物时的行为特性及共生双方利益得失。【结果】叶下珠头细蛾在广西一年两代,成虫的羽化数量与寄主植物雌雄花总数峰值变化同步;幼虫对小果叶下珠果实的寄生率为96%,单个果实内幼虫寄生数量＞1,但平均每个果实内被取食的种子为56.9%,叶下珠头细蛾低龄幼虫相对较高的死亡率限制了其对寄主种子的过度取食;果实内种子的适度保留和低龄幼虫的高死亡率是小果叶下珠-叶下珠头细蛾互利共生体系维持稳定的关键因素。【结论】小果叶下珠的表型性状出现了趋异性进化,叶下珠头细蛾与不同表型的小果叶下珠均有互利共生关系。因此,对叶下珠头细蛾形态及生物学进行详细研究有助于深入了解小果叶下珠与头细蛾共生体系物种组成多样性及进化生物学,并为探讨大戟科植物同头细蛾属昆虫协同互利共生多样性形成原因提供依据。     

叶下珠头细蛾与小果叶下珠互利共生的研究——生物学及系统稳定性

【目的】为了确定小果叶下珠Phyllanthus microcarpus与叶下珠头细蛾Epicephala sp.之间专性互利共生系统稳定性的增强是否主要通过彼此之间相互遏制对方对自身进行过度开采。【方法】通过对小果叶下珠上共生的叶下珠头细蛾形态及生物学的研究,记述了叶下珠头细蛾专性寄生寄主植物时的行为特性及共生双方利益得失。【结果】叶下珠头细蛾在广西一年两代,成虫的羽化数量与寄主植物雌雄花总数峰值变化同步;幼虫对小果叶下珠果实的寄生率为96%,单个果实内幼虫寄生数量1,但平均每个果实内被取食的种子为56.9%,叶下珠头细蛾低龄幼虫相对较高的死亡率限制了其对寄主种子的过度取食;果实内种子的适度保留和低龄幼虫的高死亡率是小果叶下珠-叶下珠头细蛾互利共生体系维持稳定的关键因素。【结论】小果叶下珠的表型性状出现了趋异性进化,叶下珠头细蛾与不同表型的小果叶下珠均有互利共生关系。因此,对叶下珠头细蛾形态及生物学进行详细研究有助于深入了解小果叶下珠与头细蛾共生体系物种组成多样性及进化生物学,并为探讨大戟科植物同头细蛾属昆虫协同互利共生多样性形成原因提供依据。     

三室算盘子寄主上弯头细蛾的生物学特性及花气味成分分析

【目的】为了从生物学和化学生态学角度探讨弯头细蛾Epicephala ancylopa和寄主三室算盘子Glochidion sp.间专性传粉的互利共生关系稳定性。【方法】本研究在野外观察和室内实验的基础上,对专性传粉育幼互利共生体系中三室算盘子、弯头细蛾生物学特性进行详细研究,探究互利共生双方利益得失;用动态顶空吸附法分别收集三室算盘子雄花和雌花气味物质,运用气相色谱-质谱联用技术(GC-MS)分析并鉴定其有效成分,用峰面积归一化与内标法定性定量;最后通过主成分分析法比较雄花和雌花之间气味化学成分的差异性。【结果】弯头细蛾在云南省普洱太阳河国家森林公园每年有1个世代,成虫和幼虫的活动时间分别在3-4月和8-10月。三室算盘子结实率为44.20%,被蛀食率为69.94%,平均每头幼虫消耗2.55枚种子来满足自身生长发育,寄主植物留有83.06%完好的种子,以维持互利共生关系的稳定。三室算盘子雌雄花气味中共鉴定出24种挥发物,主要以单萜类和倍半萜类物质为主,其中(Z)-罗勒烯和β-榄香烯两种萜类物质含量最高(分别为47.11%和22.72%),推测其是吸引弯头细蛾传粉的主要气味成分;雄花和雌花之间气味化学成分存在明显的差异,具有两性异型性。【结论】弯头细蛾通过以卵越夏和以蛹越冬对策,实现成虫发生期与三室算盘子花期的精准匹配。弯头细蛾成虫白天静伏,傍晚开始活动,三室算盘子花的气味物质也只在晚上才明显释放,且雌雄花气味化学成分的两性异型性有利于弯头细蛾辨别雌雄花,以完成采集花粉与传粉行为。该研究结果为头细蛾属昆虫与算盘子属植物专性传粉互利共生关系稳定性的维持机制提供了新的依据,也为深入开展通过触角电生理检测和生物行为实验来筛选吸引传粉头细蛾的活性物质提供了理论依据。     

小叶黑面神与小叶头细蛾互惠共生系统的种群调控机制(英文)

【目的】探索小叶黑面神Breynia vitis-idaea对小叶头细蛾Epicephala vitisidaea种群数量的调控机制。【方法】跟踪记录小叶黑面神物候及头细蛾的生物学。解剖在小叶黑面神上访花头细蛾的外生殖器,鉴定头细蛾种类。对不同时期小叶黑面神有梗和无梗的果实进行解剖,统计果实内幼虫数量、果实表面孔的数量以及果实表面产卵疤数量,计算头细蛾幼虫存活率。统计不同时期小叶黑面神有梗和无梗的果实的比例。【结果】在福建厦门小叶黑面神每年有5个花果期,相应地,为小叶黑面神传粉的头细蛾每年有5个生活世代。通过解剖,该种头细蛾被鉴定为小叶头细蛾。一头小叶头细蛾幼虫需要消耗2~4粒种子才能发育成熟。小叶黑面神有两种不同形态的果实:有梗和无梗。头细蛾幼虫在无梗果实内的存活率明显高于有梗果实,并且晚秋时期头细蛾幼虫的存活率要高于夏季。小叶黑面神无梗果实的比例在晚秋(82.04%)要高于夏季(31.53%)。【结论】本研究揭示了维持互利共生体系稳定的机制。小叶黑面神能够通过果实基部果梗的有无来调节小叶头细蛾幼虫的存活率。小叶黑面神通过季节性的调节有梗果实的比率,既有效避免了夏季种子被过度消耗的风险,又提高了头细蛾在冬季的存活率。小叶黑面神这种自身调控机制对维持小叶黑面神与小叶头细蛾互惠共生系统的稳定性起到了至关重要的作用。     

宽瓣头细蛾形态及生物学特性研究

宽瓣头细蛾Epicephala lativalvaris Li, Wang & Zhang是最近发现在黑面神Breynia fruticosa和喙果黑面神B. rostrata上专性传粉的昆虫。通过野外观察宽瓣头细蛾的访花行为、 室内饲养宽瓣头细蛾幼虫及解剖黑面神和喙果黑面神果实, 首次对其个体发育过程中各虫态的形态特征及生活习性和生活史进行了详细研究。结果表明： 宽瓣头细蛾的生活史因气候的不同而出现差异, 其在福建厦门1年发生4~5代, 有4个羽化主要高峰期, 分别为5月上旬至中旬、 6月中旬至下旬、 8月及9月下旬至10月上旬, 11月中下旬有少数成虫羽化, 12月上旬至次年4月中旬以卵或蛹越冬; 而在海南鹦哥岭1年发生6代, 无越冬现象。成虫分别主动为黑面神或喙果黑面神传粉并将卵产于子房内, 幼虫以寄主植物种子为食, 需要消耗果实内全部6粒种子才能发育成熟, 但每个种群内留有20.37%~77.63%完好的果实, 以维持互利关系的稳定。两种植物的结实率近似, 但宽瓣头细蛾对喙果黑面神的果实危害率是黑面神的2.05倍。结果说明, 在宽瓣头细蛾 黑面神和喙果黑面神专性传粉互利共生关系中, 两种寄主植物所获得的利益近似, 但喙果黑面神为宽瓣头细蛾提供了更多的回报。研究结果为头细蛾属其他种类生物学特性的了解提供了借鉴, 并为探讨宽瓣头细蛾 黑面神和喙果黑面神专性传粉互利共生关系稳定性的维持机制提供了基本依据。     

斜纹夜蛾触角叶神经元及其对植物气味和性信息素的反应

【目的】探索斜纹夜蛾Spodoptera litura触角叶结构及其神经元对植物气味和性信息素的神经识别机制。【方法】利用共聚焦激光技术扫描斜纹夜蛾成虫触角叶结构,同时采用多通道电生理技术(multi-unit recording,MR)记录斜纹夜蛾触角叶对6种寄主植物气味化合物(苯甲醛、苯甲醇、苯乙醛、水杨醛、乙酸叶醇酯和己烯醛)及性信息素顺9反11十四碳二烯乙酸酯和顺9反12十四碳二烯乙酸酯的反应;并在风洞中测定分析斜纹夜蛾对上述化合物的定向行为反应。【结果】共聚焦激光扫描结果显示,雄性和雌性斜纹夜蛾触角叶内分别密集地分布有67和66个神经纤维球;而雌性斜纹夜蛾触角叶内的纤维球总体积和平均体积都高于雄性。负责识别和追踪性信息素的扩大型纤维球复合体(macroglomerular complex,MGC)只在雄性斜纹夜蛾触角叶内发现。MR试验结果显示斜纹夜蛾触角叶内神经元具有3种自发放电模式:稀疏放电(不规则的放电频率)、温和放电(宽而慢的放电频率)和密集放电(暴发性的放电频率)。同时,斜纹夜蛾触角叶神经元对所有刺激的气味表现出3种反应类型:兴奋性、抑制性和无反应。神经元对气味的兴奋性和抑制性反应以及无反应取决于刺激化合物的结构和浓度。雌虫的触角叶神经元对性信息素和单一的植物气味表现出很小的反应,而雄虫对两种性信息素以及苯甲醛、苯甲醇、苯乙醛和水杨醛具有很强的兴奋性反应。斜纹夜蛾风洞试验也显示绝大部分的斜纹夜蛾雄虫都选择停留在性信息素和芳香族化合物上,这与MR的结果一致。【结论】神经元的反应强度和刺激化合物浓度之间的关系根据不同的神经元和刺激化合物而有所不同。在测试的浓度范围内,雄虫触角叶神经元对性信息素的反应强度随着浓度的增加而加强,但是除乙酸叶醇酯外,对其他植物气味的反应强度在测试的浓度范围内没有显著的变化。     

Active pollination favours sexual dimorphism in floral scent

Zoophilous flowers often transmit olfactory signals to attract pollinators. In plants with unisexual flowers, such signals are usually similar between the sexes because attraction of the same animal to both male and female flowers is essential for conspecific pollen transfer. Here, we present a remarkable example of sexual dimorphism in floral signal observed in reproductively highly specialized clades of the tribe Phyllantheae (Phyllanthaceae). These plants are pollinated by species-specific, seed-parasitic Epicephala moths (Gracillariidae) that actively collect pollen from male flowers and pollinate the female flowers in which they oviposit; by doing so, they ensure seeds for their offspring. We found that Epicephala-pollinated Phyllanthaceae plants consistently exhibit major qualitative differences in scent between male and female flowers, often involving compounds derived from different biosynthetic pathways. In a choice test, mated female Epicephala moths preferred the scent of male flowers over that of female flowers, suggesting that male floral scent elicits pollen-collecting behaviour. Epicephala pollination evolved multiple times in Phyllantheae, at least thrice accompanied by transition from sexual monomorphism to dimorphism in floral scent. This is the first example in which sexually dimorphic floral scent has evolved to signal an alternative reward provided by each sex, provoking the pollinator''s legitimate altruistic behaviour.     

Nocturnal emission and post-pollination change of floral scent in the leafflower tree,Glochidion rubrum,exclusively pollinated by seed-parasitic leafflower moths

Many insect-pollinated plants use floral scent signals to attract and guide the effective pollinators, and temporal patterns of their floral scent emission may be tuned to respond to the pollinator's activity and pollination status. In the intimate nursery pollination mutualism between monoecious Glochidion trees (Phyllanthaceae) and Epicephala moths (Gracillariidae), floral scent signals mediate species-specific interactions and influence the moth's efficient pollen-collecting and pollen-depositing behaviors on male and female flowers, respectively. We tested the hypotheses that both sexes of flowers of Epicephala-pollinated Glochidion rubrum exhibit a diel pattern of scent emission matching the activity period of the nocturnally active pollinator, and that female flowers change the chemical signal after pollination to reduce further visits and oviposition by the pollinator. We investigated the diel change of floral scent emissions during two consecutive days and the influence of pollination on the floral scent by conducting hand-pollinations in the field. The total scent emission of male and female flowers was higher at night than in the day, which would be expected from the nocturnal visitations of Epicephala moths. Some compounds exhibited a clear nocturnal emission rhythm. Hand-pollination experiments revealed that emission of two compounds, nerolidol and eugenol, drastically decreased in pollinated flowers, suggesting that these compounds may function as key attractants for the pollinator; however, the total scent emission of the female flower was not influenced by hand-pollination. The pattern of the floral scent emission of G. rubrum may be optimized to attract nocturnal pollinators and reduce oviposition.     

Sniffing out patterns of sexual dimorphism in floral scent

1.   A major transition in flowering plants has been the evolution of separate sexes (dioecy) from combined sexes (hermaphroditism). This transition is often, but not always, accompanied by the evolution of sexual dimorphism in attractive traits, and floral scent is no exception.
2.   In this review I aim to improve our understanding of variation in sexual dimorphism in floral scent characteristics by first explicating the relevant hypotheses, and then deriving explicit predictions for the pattern of floral scent from each.
3.   Next, I synthesize and qualitatively review published data on floral volatile emission rate and composition in 33 gender dimorphic species to identify emerging patterns, and evaluate these in light of predictions derived from the hypotheses.
4.   Although conclusions must be viewed as preliminary, a handful of strong patterns were revealed: (1) in the majority of the species studied males emit more volatiles per flower than females, (2) in over half of the species studied the sexes differed in at least one aspect of scent composition, (3) sexual dimorphism in scent composition was less common in species with rewardless females and/or brood site pollination than those that offered nectar and/or pollen, (4) a one-to-one correspondence between sex differences in overall scent composition and male organ-specific scent production is largely not found.
5.   This review has highlighted gaps in our understanding of the genesis of patterns of sexual dimorphism in floral scent, and makes clear that to move the field forward we need to shift our focus from pattern to process, and this will be best achieved by simultaneously testing alternative hypotheses at the same level of analysis.     

Floral sexual dimorphism and flower choice by pollinators in a nectarless monoecious vine Akebia quinata (Lardizabalaceae)

We investigated the pollination system and movement patterns of pollinators among flowers of the nectarless, monoecious vine Akebia quinata in natural populations and experimental floral arrays. Female flowers did not offer any rewards for pollinators and were larger than male flowers. Pollinators of A.quinata , such as small solitary bees and hoverflies, clearly discriminated between male and female flowers. Hoverflies always visited male flowers and rarely visited female flowers. In contrast, solitary bees tended to visit female flowers first when entering the array, but then switched to male flowers within the same foraging bout. This tendency disappeared when the sepal size of female flowers was experimentally reduced to the size of male flowers. Thus, observed non-randomness in flower choice by solitary bees may be caused by female>male sexual dimorphism and may increase the chance of cross-pollination and lower the probability of geitonogamous pollination in a single visit to a plant. Therefore, floral sexual dimorphism in A.quinata is considered to be an adaptation for deceptive pollination associated with discriminating pollinators.     

端黑萤和边褐端黑萤两性异形和雌性个体生育力

检测了端黑萤(Abscondita chinensis)和边褐端黑萤(Abscondita terminalis)繁殖期个体大小和形态特征的两性异形以及雌性繁殖输出。两因素方差分析表明,端黑萤的全长显著小于边褐端黑萤的全长,雌性个体显著大于雄性个体。以全长为协变量的两因素协方差分析显示,端黑萤的前胸背板宽、鞘翅长、复眼宽、胸长、腹宽、发光器面积及体重均显著小于边褐端黑萤,而腹长显著大于边褐端黑萤,触角长物种间差异不显著;雌性的前胸背板宽、腹长、腹宽及体重均显著大于雄性,而鞘翅长、复眼宽、胸长、触角长、发光器面积均显著小于雄性;物种和性别的交互作用对前胸背板宽、复眼宽、腹长、触角长及发光器面积影响显著,对其余的鞘翅长、胸长、腹宽及体重影响不显著。端黑萤和边褐端黑萤的两性异形指数分别为0.144和0.091。9个形态特征变量的主成分分析(Eigenvalue ≥ 1)发现,前2个主成分共解释56.8%的变异;复眼宽和发光器面积在第一主成分有较高的负负载系数,腹长有较高的正负载系数(解释31.3%变异);前胸背板宽和鞘翅长在第二主成分有较高的负负载系数(解释25.5%变异)。端黑萤和边褐端黑萤的平均窝卵数分别为28.9粒和18粒。线性回归显示,端黑萤和边褐端黑萤的窝卵数均与母体的全长及体重呈显著的正相关。单因素方差分析表明,特定全长的端黑萤的窝卵数显著大于边褐端黑萤。端黑萤和边褐端黑萤均属于雌性偏大的两性异形,是生育力选择、能量分配和运动综合影响的结果。雌萤腹腔容量等关键局部特征的增大是对生育力选择的适应,且生育力大小与两性异形差异程度成正相关。     

Sexual and temporal variations in floral scent in the subdioecious shrub Eurya japonica Thunb

In many flowering plants, floral scents are a significant trait for visitors, playing an important role in attracting pollinators and/or detracting herbivores. The evolution of flowering plants from hermaphroditism to dioecy is often accompanied by sexual dimorphism in floral scent. In this study, floral scents emitted by different sexual morphs of the subdioecious shrub Eurya japonica Thunb. were collected using a dynamic headspace method, and sexual and temporal variations were evaluated by gas chromatography–mass spectrometry (GC–MS). Two volatiles, α‐pinene and linalool, were identified as the major components of floral scents in females, hermaphrodites, and males. The males emit higher amounts of floral scents, particularly α‐pinene, compared to females or hermaphrodites. Floral scents emitted by males generally decrease as flowers enter senescence, whereas those from females or hermaphrodites do not significantly differ. Intraspecific variations in floral scents of subdioecious species provided by this study would contribute to better understanding of sexual dimorphism in floral scent.     

余甘子天然居群果实形态变异研究

为了从数量性状上分析余甘子天然居群果实表型性状在居群间和居群内的变异,在云南省选取了6个余甘子天然居群,分别测量了其8个果实表型性状并作比较分析。结果显示:(1)余甘子果实表型性状在居群间和居群内存在着较丰富的变异,8个果实性状在居群间的F值为20.17～35.21,部分差异达到显著水平;居群内的F值为1.27～3.07,差异均达到极显著水平。(2)余甘子居群内的平均表型变异接近90%,是余甘子果实表型遗传变异的主要来源。(3)余甘子表型性状的变化与生态和地理因子显著相关;主成分分析结果显示,单果重、果实横径、外果皮重、果核宽、果核重等5个表型性状指标是反映滇西北余甘子果实表型差异的主要因素。