聚果榕榕果小蜂种群分布格局及其生境和季节差异比较

采用多个指数对西双版纳地区聚果榕榕果小蜂的种群分布格局进行了检测,并重点运用基于方差／均值比率法的t值测度比较了其生境和季节差异．结果表明,6种聚果榕榕果小蜂的种群分布格局皆呈聚集分布;传粉小蜂聚集强度在生境序列上,原始雨林≈严重干扰地段>>中度干扰地段。在季节序列上,雨季>干热季>雾凉季;非传粉小蜂不同种类不同性别随着生境和季节变化皆表现出明显的聚集强度差异。对各种聚集强度变化的发生机理与其对聚果榕-传粉者互惠共生体进化稳定性进行了讨论,认为保护榕树、保护榕树和传粉昆虫互惠共生体的重点应放在保护其原始森林生境方面。     

聚果榕榕果小蜂种群分布格局及其生境和季节差异比较

采用多个指数对西双版纳地区聚果榕榕果小蜂的种群分布格局进行了检测,并重点运用基于方差/均值比率法的t值测度比较了其生境和季节差异.结果表明,6种聚果榕榕果小蜂的种群分布格局皆呈聚集分布;传粉小蜂聚集强度在生境序列上,原始雨林≈严重干扰地段中度干扰地段,在季节序列上,雨季>干热季>雾凉季;非传粉小蜂不同种类不同性别随着生境和季节变化皆表现出明显的聚集强度差异.对各种聚集强度变化的发生机理与其对聚果榕传粉者互惠共生体进化稳定性进行了讨论,认为保护榕树、保护榕树和传粉昆虫互惠共生体的重点应放在保护其原始森林生境方面.     

季节变化对聚果榕-榕小蜂互利共生系统生长与繁殖的影响

榕属植物及其传粉昆虫榕小蜂是自然界协同进化的经典模型,榕果内雌花资源如何分配一直是备受关注的问题。为验证季节变化对榕树-榕小蜂互利共生系统生长与繁殖的影响,该研究以西双版纳地区的聚果榕(Ficus racemosa)为材料,分析了季节变化对榕果大小、自然进蜂量以及榕树-榕小蜂繁殖的影响,并利用人工控制性放蜂实验和模型拟合,探讨榕果最适进蜂量及不同季节进蜂量对雌花资源分配的影响。结果表明:季节对榕果直径有显著影响,雨季的榕果直径显著小于干热季和雾凉季;不同季节的自然进蜂量也有显著差别,苞片口对调节进蜂数量有重要作用;季节对榕树-榕小蜂繁殖分配也有影响,雾凉季产生的种子数量和榕小蜂数量均最多;同时人工控制实验和二次抛物线模型拟合结果表明,母代雌蜂数量与种子及榕小蜂后代数量均呈抛物线关系,雌蜂数量过多或过少都对榕树-榕小蜂的繁殖不利,自然进蜂量与拟合的最优进蜂量基本一致。研究结果说明榕果进化出了适应西双版纳地区季节变化的繁殖策略。     

同种榕小蜂在木瓜榕两种地理型上的繁殖及传粉特征

榕树与其传粉榕小蜂的共生关系常被认为是专一性的,但该系统中有些榕小蜂可在不同种榕树或者同种榕树的不同亚种、变种和地理型上产卵和传粉。探讨榕小蜂在不同寄主中的繁殖和传粉特征,有利于理解非专性榕蜂系统形成的过程及稳定机制。本研究中,作者分别对比分析了传粉榕小蜂Ceratosolen emarginatus在木瓜榕(Ficus auriculata)的两种地理型auriculata-form和oligodon-form上的产卵和传粉特征。结果显示,进蜂量为1,2,3只时,相同寄主上的榕小蜂后代和种子数量均随进蜂量的增加而增加,且平均单只繁殖雌蜂的后代及种子数量均无差异。这可能是由于进蜂量较低时,两寄主可被利用的繁殖资源较充足,榕小蜂间不存在干扰竞争,可最大化地利用雌花资源。另一方面,进蜂量相同时,同一寄主上产生的种子数量明显多于榕小蜂后代数量,说明榕树的繁殖利益更占优势。榕小蜂在auriculata-form上产生的后代总量和平均单只雌蜂后代数量与oligodon-form均无差异,但后者产生的种子数量明显多于前者,说明当繁殖资源充足时寄主不影响榕小蜂的繁殖,然而寄主差异影响种子产生,即auriculata-form和oligodon-form的繁殖能力已发生分化。     

高榕雌花期传粉榕小蜂和欺骗性小蜂的繁殖特点

榕树及其专一性传粉榕小蜂组成了动植物界最为经典的协同进化关系,传粉榕小蜂演化出欺骗性是非常罕见的。在雌雄同株的高榕隐头果内,共存着一种传粉榕小蜂Eupristina altissima和一种欺骗性的小蜂Eupristina sp.,两种小蜂在雌花期进入隐头果内繁殖,但有不同的繁殖特点。对比研究了两种小蜂从成虫羽化到产卵和传粉这个阶段的雌蜂个体大小、孕卵量及繁殖差异,结果表明:羽化期两种雌蜂的平均个体小,经飞行小个体的雌蜂易死亡,大个体雌蜂到达接受树,但通过苞片通道,一些个体较大的传粉榕小蜂被夹死导致进入果腔的雌蜂相对小,而欺骗性小蜂易通过苞片以至进入果腔的雌蜂个体较大。两种未产卵雌蜂均表现为个体大者孕卵量较多,但两种雌蜂的平均孕卵量没有差异。即使有充足雌花资源产卵,两种雌蜂均未产完所有卵,产卵后两种雌蜂卵巢中的卵量均显著减少,遗留下的卵量两种小蜂间没有差异。传粉榕小蜂只有部分个体传完所携带花粉,并表现为传粉越成功的雌蜂,产卵越多。存在种内竞争时,两种小蜂的产卵量均减少,传粉榕小蜂的传粉效率也降低。在种间竞争背景下,欺骗性小蜂产卵更成功,传粉榕小蜂的产卵和传粉量均受到极大抑制。研究结果说明雌花期隐头果内传粉榕小蜂只适量利用雌花资源产卵繁殖后代,更有效地传粉繁殖榕树种子,这可能是维持榕-蜂互惠系统稳定共存的重要机制之一;欺骗者稳定存在需降低与传粉者的直接竞争,而欺骗者和传粉者分散在不同果内,甚至是不同的树上繁殖是理想的繁殖策略。     

西双版纳聚果榕隐头果内小蜂群落结构及种间关系

聚果榕Ficus racemosa Linn.是雌雄同株榕树,它是西双版纳热带雨林生态系统中的一个常见种群。聚果榕必须依靠聚果榕小蜂Ceratosolen fusciceps Mayr传粉才能获得有性繁殖,而聚果榕小蜂又必须依靠聚果榕隐头果内短柱花繁衍后代,两者间形成了种间专一的互惠共生体系。同时,在其隐头果内还存在一个复杂的非传粉小蜂功能群,它们主要是榕树种子和传粉榕小蜂的寄生者。在云南省西双版纳自治州勐腊县勐仑镇选取了5个样地,对聚果榕单果内小蜂群落组成和种间相互关系进行研究。在不同时间段采集聚果榕单果242个,共收集小蜂366660头。聚果榕隐头果内有6种小蜂,隶属小蜂总科Chalcidoidae中的榕小蜂科Agaonidae、长尾小蜂科Caliimomidae、金小蜂科Pteromalidae,其中榕小蜂科中的C．fusciceps是聚果榕唯一的传粉者。金小蜂科中的Apocryta westwoodi Grandi和Apocryta sp.两个种是榕小蜂的寄生者,它们的寄生是传粉榕小蜂的种群数量减少因素之一。长尾小蜂科中Platyneura agraensis Joseph,Platyneura mayri Rasplus和Platyneura testacea Motschulsky3个种是寄生榕树种子或与榕小蜂争夺食物(瘿花)资源的小蜂类群,它们的出现与发生,致使聚果榕正在发育成种子的长柱小花形成瘿花,同时一部分种类把卵寄生在已被榕小蜂产卵的短柱小花子房中与榕小蜂争夺食物资源,致使榕小蜂食物资源的匮乏而死亡,对传粉小蜂种群有明显的影响。传粉小蜂从花托口钻入隐头果内,在隐头果内的长柱小花传粉和短柱小花子房中产卵,5种非传粉小蜂从隐头果外部把产卵器刺穿果肉把卵产在小花子房上。通过对5块样地隐头果内小蜂群落的综合分析发现,传粉榕小蜂为优势种群,而Apocryta sp.种的数量最少。在传粉及非传粉小蜂自然群体中各种类性比明显具有偏雌现象。小蜂群落表现异常的是聚果榕孤立株的样地,该样地传粉小蜂个体数量明显下降,非传粉小蜂个体数量则增加,各小蜂种群之间竞争激烈,出现C．fusciceps和P．mayri两个优势种。     

聚果榕传粉榕小蜂传粉与产卵之间的权衡

【目的】榕树(Ficus)依赖专性榕小蜂(Agaonidae)传粉,同时为传粉榕小蜂提供繁衍后代的场所,两者形成动植物间经典的协同进化关系。在雌花期果内,榕小蜂需在有限的存活时间内完成传粉和产卵,而传粉榕小蜂如何在传粉与产卵之间进行权衡仍然是悬而未解的问题。本研究旨在明确传粉榕小蜂——一种栉颚榕小蜂Ceratosolen sp.在雌雄同株的聚果榕Ficus racemosa雌花期果内的行为活动及繁殖模式。【方法】借助测微尺测量聚果榕榕果雌花花柱长度与传粉榕小蜂(Ceratosolen sp.)产卵器长度,通过显微视频记录传粉榕小蜂在雌花期果内搜索、传粉及产卵行为;结合单果控制性引蜂试验,测定不同阶段榕小蜂个体大小、孕卵量、携粉量,以及雄花期最终繁殖的榕小蜂后代和榕果种子数量。【结果】聚果榕雌花花柱长度存在树间变异,榕小蜂产卵器长度比绝大多数的雌花花柱长,说明该小蜂可以产卵于大部分的雌花子房里。通常个体大的榕小蜂孕卵量更多,但个体大小与携粉量之间相关性不显著。观察发现,榕小蜂进入雌花期榕果内,前6 h集中产卵,可产下孕卵量的95%,平均搜索用时27 s,产卵用时46 s,此期间传粉行为少见,花粉筐中携带花粉量亦无明显变化;榕小蜂进果后6-24 h,主要执行传粉,其行为主动,连贯高效,单次传粉用时平均为2 s,最终可传完携粉量的80%。控制引蜂试验也证实榕小蜂进入榕果内前6 h主要执行产卵繁殖后代,之后6-24 h主要执行传粉以繁殖榕树种子。【结论】在雌雄同株的聚果榕雌花期榕果内,榕小蜂先产卵、后传粉。本研究首次展示了传粉榕小蜂在聚果榕雌花期榕果内的产卵和传粉行为,并获得与行为相匹配的产卵量和传粉繁殖量,反映了具主动传粉行为的榕小蜂在传粉和产卵之间存在时间和数量上的权衡。     

西双版纳热带雨林对叶榕传粉生物学(英)

研究了西双版纳热带雨林地区雌雄异株植物对叶榕 (FicushispidaL .)的生物学、传粉物候学以及榕小蜂(CeratosolensolmsimarchaliMayr)的传粉和繁殖行为。研究结果表明 :雌雄异株的对叶榕与其他雌雄同株的榕属植物不同 ,它的种子形成与传粉者有着更为复杂的相互关系。对叶榕一年结隐花果 6～ 8次 ,结果高峰期 4～ 5次 ,其中雄性植株仅产生花粉和孕育榕小蜂 ,而雌性植株 (无雄蕊 )则需榕小蜂带花粉进入隐花果内 ,进行传粉授精 ,使之发育成种子。对叶榕的成熟花粉不能从花药开裂处自行散发出来 ,必须由榕小蜂采集才能散落。榕小蜂雌蜂羽化、交配后 ,找到雄花区 ,用足、触角、口器在推动中采集花粉。雌蜂飞出熟榕果找寻雌株或雄株榕树上的幼嫩隐花果 ,一般需 3～ 6 7min ;一部分雌蜂在雄株中寻找幼嫩的隐花果 ,进去产卵繁殖 ,另一部分雌蜂则寻找雌株雌花期嫩隐花果进去传粉。雌蜂在雌株榕树的隐花果内传粉时间为 15～ 2 3h ,在雄株榕树的隐花果内产卵时间为 6～ 9h。对叶榕小蜂在雌株上进入单个隐花果的数量多少关系到雌花结实率 ;观察表明 ,每个隐花果最佳进蜂数为 2头 ;榕小蜂传粉后榕树成熟种子形成率在 5 4 .1%～ 82 .5 %之间 ,平均为 73.8% ;而在雄株上雌蜂进蜂数量则关系到榕小蜂在隐花果内的产卵率 ,     

小叶榕隐头果内加利里金小蜂的发生规律

小叶榕Ficus microcarpa是一种广泛分布于亚洲热带和亚热带地区的雌雄同株榕树,在它的隐头果中,除了专一性传粉的榕小蜂外,还寄生着多种非传粉小蜂,这些非传粉小蜂在榕果发育的不同时期来到果外产卵,对榕树和传粉榕小蜂繁衍后代产生不同的影响。加利里金小蜂Odontofroggatia galili是寄生于小叶榕雌花子房里的主要非传粉小蜂。本文在2012~2013年对该种小蜂的发生规律进行了研究,结果表明:该种小蜂雌雄蜂有翅,雄蜂二型,个体大小有显著差异。加利里金小蜂在榕果的雌花前期产卵,能独立造瘿;在西双版纳地区,干热季和雨季从卵发育到成虫羽化需31~38 d,雾凉季则长达58~72 d。成虫在榕果雄花期羽化出蜂,大多数雄蜂在苞片口等待羽化出果的雌蜂交配,雌雄蜂通常仅交配一次;为争夺交配机会,雄蜂间存在激烈打斗,且个体大者获交配机会多。成虫期雌蜂寿命2 d,雄蜂寿命仅1 d。小叶榕40%的树上隐头果内有加利里金小蜂寄生,但不同地区寄生率和单果内加利里金小蜂雌雄蜂数量树间均存在显著差异。在原生地西双版纳地区加利里金小蜂种群小,传粉榕小蜂种群占主导地位;而在引入地昆明地区,加利里金小蜂发生频率高,种群显著较大;随着纬度北移和海拔升高,在榕果内罕见有传粉榕小蜂传粉和繁殖,而加利里金小蜂种群则不断升高。说明小叶榕在海拔较高和纬度偏北地区栽种,不适宜传粉榕小蜂繁殖,而非传粉的加利里金小蜂则较为适应,获得成功繁殖及种群扩张。     

垂叶榕榕小蜂群落及种间互作网络季节动态

群落中的物种相互作用构成了复杂的生态网络。有关物种的数量和组成的季节性动态变化已有较多的研究, 但是对于生态网络的动态变化知之甚少。揭示生态网络的动态变化对于理解群落的稳定性以及群落的动态变化过程和机理具有重要意义。本研究以垂叶榕(Ficus benjamina)榕小蜂群落为研究对象, 分别在西双版纳的干季和雨季采集了榕小蜂的种类和数量信息。比较了两个季节榕小蜂群落的动态变化以及共存网络的参数(例如网路直径、连接数、嵌套性和群落矩阵温度)变化。结果显示: 雨季榕果内传粉榕小蜂Eupristina koningsbergeri所占比例高于干季, 传粉榕小蜂的种群数量也高于干季, 而在干季非传粉榕小蜂的种类增加(干季15种小蜂, 雨季14种)。从榕树-传粉榕小蜂互利共生系统的适合度来看, 干季非传粉小蜂的增加对传粉榕小蜂和榕树的适合度是不利的。在干季, 共存网络物种间的连接数(干季0.95, 雨季0.47)多于雨季, 群落矩阵温度(干季23.24, 雨季2.64)也显著高于雨季。表明干季榕小蜂群落组成及种间关系较雨季更为复杂而多样, 高的矩阵温度暗示群落受到的干扰更大。     

N:P stoichiometry in Ficus racemosa and its mutualistic pollinator

Aims Nitrogen (N) and phosphorus (P) are limiting nutrients to life across a variety of ecosystems. N:P stoichiometry, concerning the balance of these two elements, has recently received great attention. However, little is known about the nature of N:P stoichiometry in obligate mutualism.Methods N:P stoichiometry of Ficus racemosa and its pollinating wasp Ceratosolen fusciceps, an example of coevolving obligate mutualism, was investigated, and the N:P stoichiometric traits of male versus female wasps were compared.Important findings Nutrient concentrations in C. fusciceps were much higher than in its host. N enrichment in fig wasp was evidently stronger than phosphorus. N concentrations of male fig wasps were significantly higher than those of females, while P concentrations of female fig wasps were remarkably higher than those of male ones. Therefore, N:P ratios in male fig wasps were significantly greater than in female fig wasps. N:P ratio in fig-pollinating wasp displayed linear functions to fig N contents, suggesting that N limitation in fig wasps may dominate the nutritional relationship between fig pollinator and its host. Fig wasp population size had significant influences on N concentrations in host fig and female wasp per se. Driven by the nutritional stress of pollinating and parasite insects, fig fruit preferred increasing its diameter first but not nutrient richness. Values for N and P contents of fig pollinators showed seasonal differences with greater N:P ratios in dry season than in rainy season. The observations suggest that tropical climate change would result in more severe N limitation to fig-pollinating wasp and may further influence the stability of fig–fig wasp mutualism.     

Variation in trophic cascade strength is triggered by top–down process in an ant–wasp‐fig system

Changes in the strength of trophic cascades over time have been associated with dramatic shifts in community structure and function. However, the pattern, process, and potential underlying mechanism of temporal variation in trophic cascades remains relatively unexplored. A top–down trophic cascade has been documented for the effects of predacious weaver ants Oecophylla smaragdina on the success of fig tree Ficus racemosa seed production. Ants cause high mortality of non‐pollinating fig wasps Sycophaga mayri that parasitize fruits, leading to greater success for the pollinating fig wasp–fig tree mutualists. Here, using a design in which pairs of branches were selected on a tree, and ants were excluded from one of each pair, we quantified the magnitude of the trophic cascade in the cool–dry, hot–dry and rainy (hot–wet) seasons in Xishuangbanna, southwest China. We also recorded the daily behavioral dynamics of ants and fig wasps in different seasons and analyzed the correlation between behavioral, activity and trophic cascade strength. We found that the strength of the trophic cascade was strong in the hot–dry season, diminished in the rainy season and disappeared in the cool–dry season in this system. The strength of species interactions between ants and non‐pollinating fig wasps, is positively correlated with trophic cascade strength, indicating that trophic cascade strength is determined by a top–down process when the community is well established. Moreover, because pollinating fig wasps, Ceratosolen fusciceps, play a central role in the establishment of fig wasp communities, when C. fusciceps wasps are absent, the community quickly disassembles as is the case in the cool–dry season. In summary, the strength of the trophic cascade is triggered by top–down processes, however, the occurrence of the trophic cascade is determined by a keystone species that plays a central role in assembly of the community.     

Egg deposition patterns of fig pollinating wasps: implications for studies on the stability of the mutualism

1. Fig pollinating wasps (Agaonidae) enter Ficus inflorescences (figs), oviposit in some of the flowers, and pollinate in the process. Each larva completes its development within a single flower. In most cases, an inflorescence entered by a wasp will represent its only egg‐laying site. The mechanisms that prevent pollinating wasps from exploiting all the flowers inside a fig are not understood. In this study, hypotheses about flower use by pollinating fig wasps were tested by investigating egg deposition patterns in three species. 2. Either one or three wasps were introduced into figs. The figs were then harvested. Serial sections allowed assessment of the presence or absence of a wasp egg in a sample of flowers in each fig. The overall proportion of flowers with eggs and the spatial distribution of eggs were then compared in single wasp figs and three foundress figs. 3. In all species, the proportion of flowers with a wasp egg increased with foundress number but less than three‐fold. 4. In all species, at least in single foundress figs, flowers near the fig cavity were more likely to receive a wasp egg than were flowers near the fig wall. 5. In two species, when the number of foundresses was multiplied by three, there was an increase in the use of flowers near the fig wall, while in the third species, the increase was spread evenly among flowers. 6. Factors affecting wasp egg deposition patterns and the potential of investigating such patterns for studying the stability of the mutualism are discussed.     

茉莉酸甲酯诱导大戟三萜类代谢的研究

京大戟是多年生草本药用植物,入药部分是其干燥根,但可入药的京大戟资源由于生长缓慢以及环境污染的加剧而越发匮乏,因此解决大戟资源日益紧张的问题是当今药用植物资源开发与利用方向的重要课题。京大戟含有三萜类、二萜类、黄酮类等丰富的活性成分,一些常见药用植物的有效成分是三萜类化合物,其在抗病毒、抗肿瘤、免疫调节等方面具有很好的活性。对植物萜类物质代谢起重要作用的关键酶,如3-羟基,3-甲基戊二酰辅酶A还原酶(hmgr)、鲨烯合酶(sqs)、法尼基焦磷酸合酶(fps)的基因克隆及活性研究取得了进展和突破,但通过调控萜类物质代谢途径中关键酶基因的表达来诱导终产物合成的研究鲜有报道。通过研究大戟萜类物质代谢途径进而利用基因工程手段提升目的物质的产量来解决京大戟药源短缺问题具有重要意义。该研究以大戟愈伤组织为材料,使用茉莉酸甲酯分别按时间梯度和浓度梯度进行诱导,将诱导后的愈伤组织分为两部分:一部分提取其总RNA,以actin为内参基因进行反转录,实时定量RT-PCR分析大戟三萜类代谢途径中hmgr、sqs与fps基因的相对表达差异;另一部分用于提取其总三萜并使用分光光度法进行含量测定。实时定量RT-PCR分析结果表明,茉莉酸甲酯可诱导3个基因的表达,但其表达模式不一样。相应的京大戟愈伤组织中总三萜的含量明显提高,最高可较未处理样品增加27%。研究结果可为茉莉酸甲酯促进药用植物大戟三萜类物质积累的分子机制研究提供参考。     

Putting your sons in the right place: the spatial distribution of fig wasp offspring inside figs

Abstract. 1. Pollinating fig wasps (Hymenoptera, Agaonidae) display sex ratio adjustment, producing less female‐biased combined sex ratios as the number of ovipositing females (foundresses) inside a fig increases. Because males have low mobility, the oviposition sites (galled ovules) chosen by each foundress are likely to have consequences for the mating structure of wasp populations within the figs. 2. In this study, the spatial location of male and female progeny of the pollinating fig wasp Liporrhopalum tentacularis developing within figs of its host plant Ficus montana was examined to investigate two questions: (i) are male and/or female wasp offspring clustered together or interspersed? and (ii) is their distribution affected by whether one or two foundresses are present? Microsatellite markers were used to identify the progeny of different foundresses in dual‐foundress figs. 3. More offspring developed in the central part of the figs, compared with the ostiolar and basal parts, irrespective of foundress number. Neither male nor female wasp offspring were clustered within a fig. 4. The sons of the second foundress to enter a fig were positioned at similar minimum distances to both sibling and non‐sibling females, whereas the sons of the first foundress were closer to their sibling females than to non‐sibling females. If male wasps mate predominantly with females in adjacent galls, then the positioning of sons by the second foundresses is beneficial for them both in terms of reduced sibling mating and because they are provided with ready access to the female progeny of the first foundress.     

非传粉小蜂对榕-蜂共生系统的影响

榕-蜂共生系统是桑科榕属(Ficus)植物与传粉榕小蜂专一互惠形成的生态学关系。但是,也有一些非传粉的小蜂出现在这个系统中,对榕-蜂共生系统可能产生较大的影响。西双版纳的聚果榕(Ficus racemosa)树上主要有5种非传粉小蜂,分别在榕果发育的不同阶段从果外向果内产卵。在传粉榕小蜂进果之前的花前期,Platyneura testace、Apocrypta sp.和P. mayri这3种非传粉小蜂先后到果外产卵繁殖后代,对榕-蜂共生系统造成显著影响,尤其是影响传粉榕小蜂的繁殖。在传粉榕小蜂进果之后的间花期,P. mayri、A. westwoodi和P. agraensis这3种非传粉小蜂相继到果外产卵,它们虽然能减少种子形成和传粉榕小蜂繁殖的数量,但最终没有对榕-蜂共生系统造成显著的影响。造瘿类的P. mayri可在花前期和间花期产卵繁殖,在花前期产卵时它主要是影响传粉榕小蜂的繁殖,而在间花期产卵时它则更多地是影响种子的生产。