榕树的性别进化

阐述了榕树性别进化的三种理论:气候选择压力说、非传粉小蜂的选择压说和有限榕树种群维持榕小蜂种群说.并对未来有关榕树性别进化研究进行了展望.     

榕树(Ficus)在中国的分布及其在协同进化研究上的意义

中国有丰富的榕属资源(共99种),世界上6个亚属的榕树在中国均有不同程度的分布。因此,中国榕属植物的资料对从大地理尺度研究种和种间协同进化具有重要的参考价值。介绍了榕树在世界的分布规律,着重阐述了榕树在中国的分布及中国榕树分布种的研究现状与存在的问题,探讨了中国榕树的花序特点、开花物候及在协同进化中的意义,指出了中国榕树资料在世界研究中的地位和作用。     

西双版纳热带雨林中榕树动物群落结构与多样性研究

榕树是西双版纳热带雨林生态系统中的一个关键植物类群,在该类植物中,形成了以榕树果实、叶片、枝杆、枯枝落叶、土壤为食物、栖息、繁殖环境和帮助榕树授粉、传播种子的动物群落;榕树和动物之间形成了互相依赖、协同共存、互保发展的紧密关系。     

6种食用榕树叶营养成分及作为木本蔬菜的评价

榕树(Ficus)是西双版纳热带雨林中的关键种,它的许多种类是当地重要的木本野生蔬菜。对西双版纳少数民族食用的6种榕树嫩枝叶中营养成分的含量进行了分析,并与香椿(Toonasinensis)嫩枝叶中营养成分的含量进行了比较。结果表明:6种榕树嫩枝叶中VE含量高于香椿;5种榕树嫩枝叶中VB2含量为6.53～49.04mg.kg-1(干重);突脉榕嫩枝叶中VB1含量达29.48mg.kg-1(干重)。6种榕树嫩枝叶中矿物质、微量元素含量高于香椿嫩枝叶中的含量;5种榕树嫩枝叶中钙含量为0.31%～1.19%(干重);木瓜榕嫩枝叶中硒含量0.838mg.kg-1(干重),是富硒种类。6种榕树嫩枝叶中蛋白质含量为23.21%～27.78%(干重),高于香椿嫩枝叶中含量(21.33%);按氨基酸比值系数法对6种榕树和香椿嫩枝叶中必需氨基酸与WHO/FAO模式比较,氨基酸比值系数分为69.16～95.98,其中5种榕树嫩枝叶的比值系数分高于80,4种高于香椿(86.1)。食用的榕树嫩枝叶是优质的木本蔬菜。     

两种生态型榕树的叶绿素含量、荧光特性和叶片气体交换日变化的比较研究

比较盆栽 生榕树和两栖型树的形态差异、叶片叶绿素含量、叶绿素荧光特性和气体交换的日变化。两栖型榕树具有较发达的气生根和水生不定根,叶片比陆生榕树宽,并有向中生性 倾向,陆生榕树的叶绿素含量比两栖榕树高,净光合速率略高于水培两栖型榕树,但明显高于土培两栖型榕树,蒸腾速率以水培两栖型树最高,陆生榕树次之,土培两栖型榕树最低,线性回归分析表明,三者的叶片气孔导度与净光合速率变化均呈正相关,气孔导度的变化     

榕树-榕小蜂互惠合作系统中的非对称性博弈

榕树及其传粉榕小蜂是自然界中目前所知道的关系最为紧密的互利共生系统之一。随着研究的深入, 越来越多的证据发现榕树-传粉榕小蜂之间互惠合作的过程中存在着复杂的竞争和对抗关系, 例如榕树与传粉榕小蜂之间对公共资源的竞争、传粉欺骗与宿主对传粉者的惩罚、榕树与传粉小蜂之间的“军备竞赛”等。在相互竞争或者对抗关系中, 双方表现出非对称性相互作用。其非对称性关系主要表现出如下3个特征: (1)收益不对称, 即榕树(宿主)与传粉榕小蜂(共生体)之间在资源利用等方面的实力不对称; (2)榕树与传粉榕小蜂之间的信息不对称; (3)进化速率不对称。这些非对称的相互作用可能导致种群的波动、榕树与传粉榕小蜂相互适应和进化策略的变化。因此, 理解榕树与传粉榕小蜂之间的非对称交互作用有助于理解为什么合作和冲突在互利共生关系中经常能同时存在, 也将有助于解释榕树-传粉榕小蜂种间相互关系和物种的多样性。     

半附生榕树和非附生榕树的细根性状

榕树作为热带雨林生态系统中的一个关键类群,在维持生物多样性方面发挥着重要作用。本研究以中国科学院西双版纳热带植物园内的榕树专类园区的10种榕属树种(5种半附生,5种非附生)为对象,采用根序法对其细根直径、根长、比根长、比表面积、分枝密度、组织密度、氮含量、碳含量和碳氮比等9个细根功能性状进行了研究,分析了半附生榕树和非附生榕树的细根性状差异;同时结合其原始分布生境,从植物的细根形态功能性状角度阐释其生存策略。结果表明:半附生榕的细根根长、细根直径显著高于非附生榕树(P!0.01),而其比根长和分枝密度显著低于非附生榕树,其余性状之间无显著差异(P>0.05); 10种榕树的细根形态特征与养分特征呈微弱或无相关关系,细根的直径与分枝密度、比根长和比表面积呈显著的负相关。研究认为,不同生活型榕树的根系结构和性状特征差异是对环境的适应表现,分布于山脊干旱环境的半附生榕树细根具有直径较粗、根系长、分枝密度和比根长低的特性,使其具有较好的水分吸收能力,并可能与菌根真菌有更紧密的互利共生关系,从而能适应相对干旱的山脊环境。     

榕树-传粉者共生体系的协同进化与系统学研究进展及展望

 榕树-传粉者共生体系是目前植物与昆虫协同进化研究中的典型模式之一。国内外已经开展了大量的相关研究,从不同方面探讨了其特殊的一一对应的共生关系。榕树-传粉者的专一性互惠共生关系中蕴含了与系统发育有关的多因子协同进化的机理,因此,进行系统发育研究将有助于更好地揭示榕树-传粉者的协同进化历史和理解二者的专一性互惠共生关系。本文简单地介绍了目前榕树及其传粉者共生体系的研究状况之后,论述了榕树-传粉者协同进化的系统发育分子生物学研究成果。同时针对国际上在榕属植物的传统的系统与分类研究中存在的一些分歧及榕树传粉者亚科分类不匹配等问题,回顾了榕属的分类研究进展及其与传粉者的关系。最后,结合我国榕树与传粉者共生体系的研究状况对我国榕属的重新分类和系统发育研究作了展望。     

钝叶榕果实内繁殖的两种榕小蜂与寄主榕树间的协同进化

 榕树(Ficus)及其传粉榕小蜂(Agaonidae)构成了高度专一的互惠共生体系。榕树的果实(以下简称榕果)内也寄生着一些非传粉小蜂。 绝大多数非传粉小蜂在榕果外把产卵器刺入果壁产卵到果腔内, 只有极少数种类能够进入果腔内产卵。在西双版纳地区, 钝叶榕(Ficus curtipes)上的杨氏榕树金小蜂(Diaziella yangi)类似于传粉者钝叶榕小蜂(Eupristina sp.), 它也是进入榕果内产卵繁殖后代的, 这就为比 较研究榕果内产卵小蜂与寄主榕树间的关系提供了材料。该文从形态学、行为学和生态学角度比较研究了这两种进入榕果内产卵的小蜂与寄主 钝叶榕之间的作用关系, 研究结果显示: 1)杨氏榕树金小蜂与钝叶榕小蜂的雌蜂头部形状存在趋同进化; 2)两种小蜂的产卵器的平均长度都比 雌花花柱长, 因而能把卵产在子房里; 3)钝叶榕小蜂从瘿花出来需要3~5 h, 交配需要17~19 min, 杨氏榕树金小蜂从瘿花出来只需18~20 min, 交配时间为20~30 s; 4)在自然群落中, 大约90%的雌花期榕果里都只进一只杨氏榕树金小蜂和一只钝叶榕小蜂, 杨氏榕树金小蜂能通过传粉来 增加榕树种子数量, 但对钝叶榕小蜂种群的繁衍造成了极显著的负面影响; 5)两种小蜂于同一时期进入榕果内繁殖, 子代同期成熟羽化, 发育 期与榕树雄花的发育期同步。研究表明: 进入榕果内繁殖的两种小蜂与寄主榕树之间存在着协同进化关系, 杨氏榕树金小蜂为榕树有效地传粉, 这可能是一个由寄生者向互惠方向进化的实例。     

热带雨林三种榕树隐头果昆虫群落结构与功能群生态特性

鸡嗉果榕Ficus semicordata、苹果榕Ficus oligodon、对叶榕Ficus hispida是西双版纳热带雨林和次生林中最常见的树种,它们是热带雨林生态系统中的一类先锋种。在每一种榕树隐头果内外,形成了给榕树传粉功能群、寄生子房制造瘿花功能群、复寄生其它昆虫功能群、蛀食和取食隐头果功能群、捕食性天敌功能群、刺吸隐头果汁液功能群、取食熟果加速果实腐烂分解功能群等昆虫组成的群体。本工作在西双版纳勐仑国家自然保护区、植物园沟谷雨林、植物园榕树园等样地内对鸡嗉果榕、苹果榕、对叶榕3种榕树隐头果内外的昆虫种群结构和功能群生态学特性开展了研究。结果表明：样地内采集到141037号昆虫标本,隶属6个目,18个科,42属,61种。从种类上比较,捕食功能群、蛀食及取食功能群最丰富,占榕树隐头果昆虫总种数的57．4％。从个体数量上来看,传粉类群的数量最大,占总数的68．6％。每一种榕树与传粉功能群(榕小蜂)之间构成了种内专一的互惠共生关系,这种关系是由于榕树必须依赖榕小蜂的传粉而获得有性繁殖,而榕小蜂也必须依赖榕树隐头果内部分小花子房作为繁殖与生存场所才能正常繁衍后代形成的,它们相互之间在生物界中形成了最密切的协同进化关系。传粉昆虫功能群、捕食昆虫功能群对榕树隐头果的生长发育起着促进的作用,而其它功能群或多或少均对榕树隐头果的生长发育起着损害性作用,特别是制造瘿花的非传粉小蜂功能群破坏性最大,次之是蛀食和取食功能群、刺吸果汁功能群、分解功能群对隐头果也起到危害作用,而复寄生昆虫功能群则对传粉功能群造成种群下降,从而致使传粉功能降低,对榕树隐头果起着间接的负作用。     

A comparison of growth and reproduction, under laboratory conditions, of males and females of a dioecious fig tree

The interaction between Ficus spp. (Moraceae) and their pollinating wasps (Chalcidoidae: Agaonidae) is a highly co-evolved mutualism. Approximately half of all fig species are monoecious and produce a mixture of wasps and seeds within the same fig. In functionally dioecious fig trees male and female functions are separate. Figs on male trees produce wasps and pollen, whereas figs borne on female plants produce only seeds. Dioecious fig phenology provides an excellent opportunity to investigate the effect of sexual specialization on the obligate fig?Cfig wasp interaction and the non-pollinators associated with the system. Here we describe laboratory studies of phenological variation between two sexes in terms of vegetative growth and fig production in a dioecious fig tree Ficus montana. We also describe reproductive output in terms of wasp production in males and seeds in females. Intrasexual asynchrony was observed for the plants, with synchrony between the sexes with year-round production of figs. Male plants grew more rapidly, but leaf phenology was very similar. Crop sizes and development times were the same for males and females. Seasonal effects were strong for leaf phenology and fig initiation, but had a very limited effect on fig composition. The results show that the phenological differences described for other dioecious figs do not apply to all species.     

Variation in inflorescence size in a dioecious fig tree and its consequences for the plant and its pollinator fig wasp

The host-specific relationship between fig trees (Ficus) and their pollinator wasps (Agaonidae) is a classic case of obligate mutualism. Pollinators reproduce within highly specialised inflorescences (figs) of fig trees that depend on the pollinator offspring for the dispersal of their pollen. About half of all fig trees are functionally dioecious, with separate male and female plants responsible for separate sexual functions. Pollen and the fig wasps that disperse it are produced within male figs, whereas female figs produce only seeds. Figs vary greatly in size between different species, with female flower numbers varying from tens to many thousands. Within species, the number of female flowers present in each fig is potentially a major determinant of the numbers of pollinator offspring and seeds produced. We recorded variation in female flower numbers within male and female figs of the dioecious Ficus montana growing under controlled conditions, and assessed the sources and consequences of inflorescence size variation for the reproductive success of the plants and their pollinator (Kradibia tentacularis). Female flower numbers varied greatly within and between plants, as did the reproductive success of the plants, and their pollinators. The numbers of pollinator offspring in male figs and seeds in female figs were positively correlated with female flower numbers, but the numbers of male flowers and a parasitoid of the pollinator were not. The significant variation in flower number among figs produced by different individuals growing under uniform conditions indicates that there is a genetic influence on inflorescence size and that this character may be subject to selection.     

Living on the edge: Fig tree phenology at the northern range limit of monoecious Ficus in China

Fig trees (Ficus) are a species-rich group of mainly tropical and subtropical plants that are of ecological importance because of the large numbers of vertebrates that utilise their figs for food. Factors limiting their distributions to warmer regions are still poorly understood, but are likely to include factors linked to their specialised pollination biology, because each Ficus species is dependent on one or a small number of host-specific fig wasps (Agaonidae) for pollination. Adult fig wasps are short-lived, but some species are capable of dispersing extremely long distances to pollinate their hosts. Close to its northern range limit we investigated the phenology of Ficus virens, the monoecious fig tree that reaches furthest north in China. Relatively few trees produced any figs, and very few retained figs throughout the winter. Despite this, new crops produced in spring were pollinated, with seasonally migrant pollinators from plants growing further south the most likely pollen vectors. An inability to initiate new crops at low temperatures may limit the distribution of monoecious fig trees to warmer areas.     

介质中不同Ca~(2+)浓度对五种榕树幼苗钙含量的影响

选择高榕(Ficusaltissima)、木瓜榕(F.auriculata)、聚果榕(F.racemosa)、鸡嗉子榕(F.semicorda ta)以及对叶榕(F.hispida)5种榕树,对其幼苗进行不同Ca2+(CaCl2)浓度下的水培实验,结果表明,5种幼苗钙含量随介质Ca2+浓度的增加而增加,增加趋势大致相同。高Ca2+浓度下(10mmol/L)幼苗地上部的钙含量为低Ca2+浓度下(0.05mmol/L)的1.51～1.63倍。5种榕树幼苗钙含量随培养液Ca2+浓度变化的Yadj值的多重比较结果显示,对叶榕钙含量显著高于其他四种(P<0 01),高榕则显著低于其他四种(P<0 01),木瓜榕、聚果榕、鸡嗉子榕之间的差异不显著。不同榕树幼苗地上部分钙含量受介质钙浓度影响,但可能主要决定于基因型的差异,表明榕属植物富钙基因的存在,这为高钙榕树植物资源开发提供初步的理论依据。     

First detection of “Candidatus Phytoplasma asteris” - and “Candidatus Phytoplasma solani”-related strains in fig trees

In July 2017, a survey was conducted in a fig collection plot at Locorotondo (south of Italy) to investigate the possible presence of phytoplasmas in plants showing yellowing, deformed leaves, short internodes, mottling and mosaic. Samples were collected from symptomatic plants and tested by nested PCR assays using universal and specific primers to amplify the 16S rDNA of these prokaryotes. PCR results detected the presence of phytoplasma sequences in twenty plant samples that resulted clustering two phylogenetically distinct phytoplasmas, i.e., “Candidatus Phytoplasma asteris” and “Candidatus Phytoplasma solani” affiliated to 16SrI and 16SrXII ribosomal groups, respectively. The presence of phytoplasmas belonging to both ribosomal groups was confirmed with group specific quantitative PCR and RFLP assays on 16S ribosomal amplicons. Results of this study indicate for the first time the occurrence of phytoplasmas in fig; however, more work should be carried out to verify their association with the symptoms observed on diseased fig plants.     

Sexual differences in the attractiveness of figs to pollinators: females stay attractive for longer

1. Figs on male dioecious fig trees (Ficus, Moraceae) are breeding sites for pollinator fig wasps (Hymenoptera, Agaonidae), but figs on female plants are traps that produce only seeds. As the short‐lived fig wasps cannot reproduce in female figs, natural selection should favour individuals that avoid them. Several studies have failed to detect such discrimination, a result attributed to inter‐sexual mimicry and ‘selection to rush’ in the wasps, but their experiments failed to explicitly take into account fig age (how long they had been waiting to be pollinated). 2. We compared the relative attraction of male and female figs of known ages of the South East Asian Ficus montana Burm. f. to its pollina tor Liporrhopalum tentacularis Grandi and examined how the reproductive success of the plant and its pollinator change with the age of the figs. 3. Mean retention time for un‐pollinated figs on female plants was 16 days whereas in male figs it was 12 days. Female figs remained attractive for up to 2 weeks, although the wasps were less willing to enter older figs. After pollinator entry, receptivity continued for several days, lasting longer in figs entered by a single wasp. Consistent with abortion rates, attractiveness persisted longer in female figs. Older figs produced fewer fig wasp offspring, but similar numbers of seeds. 4. The sexual differences in floral longevity in F. montana may represent part of a previously un‐recognised reproductive strategy in some fig trees that allows male plants to ‘export’ pollinators while also maintaining a resident fig wasp population.     

Seasonality of Leaf and Fig Production in Ficus squamosa,a Fig Tree with Seeds Dispersed by Water

The phenology of plants reflects selection generated by seasonal climatic factors and interactions with other plants and animals, within constraints imposed by their phylogenetic history. Fig trees (Ficus) need to produce figs year-round to support their short-lived fig wasp pollinators, but this requirement is partially de-coupled in dioecious species, where female trees only develop seeds, not pollinator offspring. This allows female trees to concentrate seed production at more favorable times of the year. Ficus squamosa is a riparian species whose dispersal is mainly by water, rather than animals. Seeds can float and travel in long distances. We recorded the leaf and reproductive phenology of 174 individuals for three years in Chiang Mai, Northern Thailand. New leaves were produced throughout the year. Fig production occurred year-round, but with large seasonal variations that correlated with temperature and rainfall. Female and male trees initiated maximal fig crops at different times, with production in female trees confined mainly to the rainy season and male figs concentrating fig production in the preceding months, but also often bearing figs continually. Ficus squamosa concentrates seed production by female plants at times when water levels are high, favouring dispersal by water, and asynchronous flowering within male trees allow fig wasps to cycle there, providing them with potential benefits by maintaining pollinators for times when female figs become available to pollinate.     

Insect responses to host plant provision beyond natural boundaries: latitudinal and altitudinal variation in a Chinese fig wasp community

Many plants are grown outside their natural ranges. Plantings adjacent to native ranges provide an opportunity to monitor community assembly among associated insects and their parasitoids in novel environments, to determine whether gradients in species richness emerge and to examine their consequences for host plant reproductive success. We recorded the fig wasps (Chalcidoidea) associated with a single plant resource (ovules of Ficus microcarpa) along a 1200 km transect in southwest China that extended for 1000 km beyond the tree's natural northern range margin. The fig wasps included the tree's agaonid pollinator and other species that feed on the ovules or are their parasitoids. Phytophagous fig wasps (12 species) were more numerous than parasitoids (nine species). The proportion of figs occupied by fig wasps declined with increasing latitude, as did the proportion of utilized ovules in occupied figs. Species richness, diversity, and abundance of fig wasps also significantly changed along both latitudinal and altitudinal gradients. Parasitoids declined more steeply with latitude than phytophages. Seed production declined beyond the natural northern range margin, and at high elevation, because pollinator fig wasps became rare or absent. This suggests that pollinator climatic tolerances helped limit the tree's natural distribution, although competition with another species may have excluded pollinators at the highest altitude site. Isolation by distance may prevent colonization of northern sites by some fig wasps and act in combination with direct and host‐mediated climatic effects to generate gradients in community composition, with parasitoids inherently more sensitive because of declines in the abundance of potential hosts.     

Genetic characterization of fig tree mutants with molecular markers

The fig (Ficus carica L.) is a fruit tree of great world importance and, therefore, the genetic improvement becomes an important field of research for better crops, being necessary to gather information on this species, mainly regarding its genetic variability so that appropriate propagation projects and management are made. The improvement programs of fig trees using conventional procedures in order to obtain new cultivars are rare in many countries, such as Brazil, especially due to the little genetic variability and to the difficulties in obtaining plants from gamete fusion once the wasp Blastophaga psenes, responsible for the natural pollinating, is not found in Brazil. In this way, the mutagenic genetic improvement becomes a solution of it. For this reason, in an experiment conducted earlier, fig plants formed by cuttings treated with gamma ray were selected based on their agronomic characteristics of interest. We determined the genetic variability in these fig tree selections, using RAPD and AFLP molecular markers, comparing them to each other and to the Roxo-de-Valinhos, used as the standard. For the reactions of DNA amplification, 140 RAPD primers and 12 primer combinations for AFLP analysis were used. The selections did not differ genetically between themselves and between them and the Roxo-de-Valinhos cultivar. Techniques that can detect polymorphism between treatments, such as DNA sequencing, must be tested. The phenotypic variation of plants may be due to epigenetic variation, necessitating the use of techniques with methylation-sensitive restriction enzymes.