海南岛的白蚁垅巢及歪白蚁属一新种(等翅目:白蚁科)

白蚁在地面营造土垅巢的现象,是反映热带景观的一种特色。我国在地面营造土垅巢的白蚁,首先在云南、广西发现,计有三种:土垅大白蚁Macrotermes annandalei (Silvestri);云南土白蚁Odontotermes yunnanensis Tsai et Chen;黄球白蚁Globitermes sulphureus (Haviland)。以上三种白蚁的垅巢,蔡邦华、陈宁生(1964)有详细介绍。而海南岛的白蚁垅巢,一直未报导过。现就海南岛的白蚁垅巢及筑垅巢歪白蚁的一新种,记述如下:     

白蚁巢菌圃真菌多样性研究

培菌性白蚁能在存在于蚁巢或分散在其周围土壤中的菌圃上培养真菌。菌圃在无白蚁存在下培养会生长出炭角菌的子实体。对分别采集自我国西南四川、云南两省的4个土白蚁属菌圃采用原位培养法分离并纯化得到40株炭角菌,划分为13个形态型,ITS1-5.8S-ITS2序列分析确定为两种炭角菌。采用建立ITS基因文库的方法分析了白蚁菌圃真菌群落多样性,结果表明有白蚁存在的菌圃,蚁巢伞为单一优势菌;废弃的蚁巢中的菌圃,木霉、炭角菌等其他真菌成为优势菌。     

培菌白蚁菌圃微生物降解木质纤维素的研究进展

白蚁及其共生微生物协同降解植物细胞壁的机理一直被世界各国科学家所关注。培菌白蚁作为高等白蚁,相比低等食木白蚁具有更多样化的食性,其利用外共生系统“菌圃”,对多种植物材料进行处理。本文综述了菌圃微生物降解木质纤维素的研究进展,以期为深入研究菌圃中木质纤维素降解过程及其机制,并挖掘利用菌圃降解木质纤维素的能力及仿生模拟菌圃开发新的生物质利用系统提供参考。培 菌白蚁在其巢内利用由植物材料修建的多孔海绵状结构——“菌圃”来培养共生真菌鸡枞菌Termitomyces spp.,形成了独特的木质纤维素食物降解和消化策略,使木质纤维素在培菌白蚁及其共生微生物协同作用下被逐步降解。幼年工蚁取食菌圃上的共生真菌菌丝组成的小白球和老年工蚁觅得食物并排出粪便堆积到菌圃上成为上层菌圃。这一过程中,被幼年工蚁取食的共生真菌释放木质素降解酶对包裹在植物多糖外部的木质素屏障进行解聚。菌圃微生物(包括共生真菌)对解聚的木质素基团进一步降解,将多糖长链或主链剪切成短链,使菌圃基质自下而上被逐步降解。最后下层的老熟菌圃被老年工蚁取食,其中肠的内源酶系及后肠微生物将这些短链进一步剪切和利用。因此,蚁巢菌圃及其微生物是培菌白蚁高效转化利用木质纤维素的基础。化学层面的研究表明,菌圃能够实现对植物次生物质解毒和植 物纤维化学结构解构。对共生真菌相关酶系的研究显示可能其在菌圃的植物纤维化学结构和植物次生物质的降解中发挥了作用,但不同属共生真菌间其效率和具体功能不尽相同。而菌圃中的细菌是否发挥了作用和哪些细菌类群发挥了作用等仍有待进一步的研究。相比于低等食木白蚁利用其后肠共生微生物降解木质纤维素,培菌白蚁利用菌圃降解木质纤维素具有非厌氧和能处理多种类型食物两大优势,仿生模拟菌圃降解木质纤维素的机制对林地表面枯枝落叶的资源化利用具有重要意义。     

培菌白蚁菌圃和粪便微生物多样性分析

【背景】培菌白蚁是属于白蚁科的一类与鸡枞菌属真菌共生的高等白蚁,其与体内肠道微生物和体外菌圃微生物形成三维共生体系。【目的】分析培菌白蚁菌圃和粪便的微生物多样性,并与肠道微生物进行比较。【方法】通过Illumina MiSeq高通量测序方法对培菌白蚁菌圃和粪便样品进行细菌16S rRNA基因和真菌ITS测序分析。【结果】高通量测序获得培菌白蚁菌圃和粪便样品细菌和真菌的有效序列和OTU数目。5个样品细菌OTU数目在90-199之间,而真菌OTU在10-58之间,细菌的种类多样性明显大于真菌。不论是细菌还是真菌,粪便样品的OTU数目多于菌圃样品。经物种分类分析,菌圃样品主要优势细菌是变形菌门(Proteobacteria),其相对含量超过82.4%;其次是拟杆菌门(Bacteroidetes)和厚壁菌门(Firmicutes);粪便样品中优势细菌为拟杆菌门,其次是变形菌门,粪便优势菌属为别样杆菌属和营发酵单胞菌属,这与培菌白蚁肠道菌多样性组成一致。培菌白蚁菌圃和粪便样品共生真菌主要为担子菌门(Basidiomycota)和子囊菌门(Ascomycota)。菌圃优势真菌为鸡枞菌属(Termi...     

白蚁菌圃真菌多样性研究进展

白蚁菌圃存在于白蚁巢中,具有硬而脆的多孔结构,是特殊的真菌生存环境。当有白蚁在白蚁巢内活动时,蚁巢伞Termitomyces是菌圃上的优势菌;当白蚁巢被废弃,炭角菌Xylaria成为菌圃上的优势真菌。菌圃中还存在其他微生物如无性型真菌（anamorphic fungi）和酵母等。菌圃中的真菌很多具有潜在药用价值或其他经济价值。从蚁巢伞、炭角菌等主要真菌类群出发,结合分子生态学研究菌圃真菌多样性的方法,综述了白蚁菌圃真菌多样性的研究进展,揭示了目前的研究热点及存在的问题,并针对这些问题提出可能的发展方向。     

培菌白蚁与鸡枞菌研究进展

培菌白蚁起源于非洲,蚁巢内具有复杂的社会分工.培菌白蚁依靠独特的蚁巢结构维持内部稳态和气体循环.菌圃是白蚁培育鸡枞菌的场所.鸡枞菌隶属于担子菌亚门,但其传播方式和生活史具有区别于其它担子菌的特点.鸡枞菌协助白蚁进行植物纤维的消化,白蚁则为鸡枞菌提供合适的生长环境,并控制鸡枞菌的遗传结构.培菌白蚁和鸡枞菌形成紧密的共生关...     

黑翅土白蚁菌圃对采食工蚁的引诱活性

黑翅土白蚁是一种重要的培菌白蚁,采食工蚁取食菌圃物质,采集树皮等植物材料作为菌圃的培养基。本文研究了菌圃、林间采集材料(杂交鹅掌楸树干外表皮)对采食工蚁的引诱活性,并与常用的白蚁饵料基质密粘褶菌松木粉培养物进行了比较。利用Y形嗅觉仪分别测定上、中、下层主巢菌圃、副巢菌圃、杂交鹅掌楸树干外表皮、密粘褶菌松木粉培养物对采食工蚁的引诱活性;比较了主巢下层菌圃与杂交鹅掌楸树干外表皮、主巢下层菌圃与密粘褶菌松木粉培养物、副巢菌圃与杂交鹅掌楸树干外表皮、副巢菌圃与密粘褶菌松木粉培养物、杂交鹅掌楸树干外表皮与密粘褶菌松木粉培养物的引诱活性。结果显示,(1)空白对照条件下,不同测定材料趋性反应率大小顺序:密粘褶菌松木粉培养物副巢菌圃主巢下层菌圃主巢中层菌圃主巢上层菌圃杂交鹅掌楸树干外表皮。不同材料的趋性反应率显著性检验:密粘褶菌松木粉培养物与副巢菌圃、副巢菌圃与主巢下层菌圃、主巢下层菌圃与主巢中层菌圃、主巢中层菌圃与主巢上层菌圃、主巢上层菌圃与杂交鹅掌楸树干外表皮之间差异不显著,其余各组差异显著或极显著。(2)不同测定材料趋势反应率两两比较测定:密粘褶菌松木粉培养物高于杂交鹅掌楸树干外表皮(P0.01);副巢菌圃高于杂交鹅掌楸树干外表皮(P0.01);主巢下层菌圃高于杂交鹅掌楸树干外表皮(P0.05);副巢菌圃与密粘褶菌松木粉培养物、主巢下层菌圃与密粘褶菌松木粉培养物差异不显著。(3)黑翅土白蚁共生菌圃对采食工蚁具有明显的引诱活性,引诱活性强度与菌圃的类型、部位有关。(4)主巢下层菌圃、副巢菌圃有较高的引诱活性,引诱活性强度与密粘褶菌松木粉培养物相近。相比之下,主巢中层菌圃、主巢上层菌圃以及杂交鹅掌楸树干外表皮引诱活性较低。菌圃物质对采食工蚁的引诱活性比林间采集材料的引诱活性更高。共生菌圃中引诱活性组分的理化性质、组成特点、来源等尚需进一步研究。     

土垅大白蚁Macrotermes annandalei(Silvestri)采食品级的踪迹信息素

【目的】解析土垅大白蚁Macrotermes annandalei(Silvestri)采食行为中的踪迹信息交流机制。【方法】通过对其踪迹信息素进行化学和行为活性分析。【结果】土垅大白蚁兵蚁腹板腺源踪迹信息素的主要成分为顺-3-十二碳单烯醇-1((3Z)-dodec-3-en-l-ol,DOE),与工蚁的踪迹信息素化学结构完全相同。大兵蚁腺体当中DOE的含量最高,分别是大、小工蚁品级和小兵蚁品级DOE含量的3.88倍和3.78倍,与其在采食行为中的功能相适应。尽管小兵蚁的位置处于采食队伍的先锋,但小兵蚁的踪迹信息素含量与工蚁相当,不能起到召募采食的作用。行为观察和跟踪信息素行为活性表明大兵蚁具有通过分泌大量踪迹信息素召募工蚁品级的能力。【结论】土垅大白蚁依靠大兵蚁分泌大量踪迹信息素单一组分及其在采食队伍中的位置的变化来实现采食行为中的方向信息的快速传递交流。     

我国白蚁资源利用的现状

白蚁是世界性害虫,但白蚁资源有着很高的利用价值。文章简要论述我国白蚁资源利用的研究现状及研究进展,主要包括白蚁本身、蚁巢及菌圃、蚁巢共生物--鸡榕菌Termitomyces albuminosus(Berk.)Heim、乌灵参Xylaria nigripes等直接利用价值及白蚁活动加速自然界物质循环、改善土壤理化性质、探矿等间接利用价值。同时还对白蚁资源研究利用的前景进行了展望。     

谈谈白蚁与人类的密切关系

白蚁对人类的危害众所周知 ,白蚁对人类的有益方面 ,人们知之甚少。该文重点论述白蚁对人类有益的方面。白蚁在自然界中 ,是一个名符其实的“清道夫” ,它将林地中的枯枝、落叶等转化为有益的肥料 ,回归自然 ,再被植物吸收 ;白蚁本身可加工成营养丰富的美味佳肴食品 ,也可精制成防病治病的良药和效果明显的保健补品 ;白蚁的附生物 ,如鸡菌更是美味佳肴 ,菌圃和炭棒菌也能精制成治疗多种疾病的良药。     

Fungal nutrition allocation enhances mutualism with fungus-growing termite

Fungal nodules and aged fungus gardens are products of termite fungiculture systems, and are the diets of termites. To understand the nutrition flow in fungiculture, we quantified the number and mass of fungal nodules produced along with fungus garden maturation and analysed the α-amino acid and fatty acid compositions of fungal nodules, fungus gardens, and termite tissues of a fungus-growing termite, Odontotermes formosanus. 1 g of fungus garden produced 5,148 fungal nodules (∼68.0 mg). Approximately 7.0% of α-amino acids were allocated to the fungal nodules and the rest (∼93.0%) remained in the fungus gardens. The compositions of α-amino acids or fatty acids in aged fungus gardens and fungal nodules were more similar to that of termite tissues than fresh fungus gardens, which supports the idea that termites nutritionally depend on the fungal products. Among the 18 α-amino acids, tryptophan was an essential amino acid and was the only one missing from fresh and aged fungus gardens, but found in fungal nodules at significantly higher concentrations. Hence, termites must consume fungal nodules to obtain tryptophan for survival. Furthermore, the fungus spores incorporated in nodules, were transferred when nodules were ingested by termites. We propose that allocating tryptophan in fungal nodules is crucial to enhance the mutualism between the fungus and termite.     

Metagenomic and metaproteomic insights into bacterial communities in leaf-cutter ant fungus gardens

Herbivores gain access to nutrients stored in plant biomass largely by harnessing the metabolic activities of microbes. Leaf-cutter ants of the genus Atta are a hallmark example; these dominant neotropical herbivores cultivate symbiotic fungus gardens on large quantities of fresh plant forage. As the external digestive system of the ants, fungus gardens facilitate the production and sustenance of millions of workers. Using metagenomic and metaproteomic techniques, we characterize the bacterial diversity and physiological potential of fungus gardens from two species of Atta. Our analysis of over 1.2 Gbp of community metagenomic sequence and three 16S pyrotag libraries reveals that in addition to harboring the dominant fungal crop, these ecosystems contain abundant populations of Enterobacteriaceae, including the genera Enterobacter, Pantoea, Klebsiella, Citrobacter and Escherichia. We show that these bacterial communities possess genes associated with lignocellulose degradation and diverse biosynthetic pathways, suggesting that they play a role in nutrient cycling by converting the nitrogen-poor forage of the ants into B-vitamins, amino acids and other cellular components. Our metaproteomic analysis confirms that bacterial glycosyl hydrolases and proteins with putative biosynthetic functions are produced in both field-collected and laboratory-reared colonies. These results are consistent with the hypothesis that fungus gardens are specialized fungus–bacteria communities that convert plant material into energy for their ant hosts. Together with recent investigations into the microbial symbionts of vertebrates, our work underscores the importance of microbial communities in the ecology and evolution of herbivorous metazoans.     

Farming termites determine the genetic population structure of Termitomyces fungal symbionts

Symbiotic interactions between macrotermitine termites and their fungal symbionts have a moderate degree of specificity. Consistent with horizontal symbiont transmission, host switching has been frequent over evolutionary time so that single termite species can often be associated with several fungal symbionts. However, even in the few termite lineages that secondarily adopted vertical symbiont transmission, the fungal symbionts are not monophyletic. We addressed this paradox by studying differential transmission of fungal symbionts by alate male and female reproductives, and the genetic population structure of Termitomyces fungus gardens across 74 colonies of Macrotermes bellicosus in four west and central African countries. We confirm earlier, more limited, studies showing that the Termitomyces symbionts of M. bellicosus are normally transmitted vertically and clonally by dispersing males. We also document that the symbionts associated with this termite species belong to three main lineages that do not constitute a monophyletic group. The most common lineage occurs over the entire geographical region that we studied, including west, central and southern Africa, where it is also associated with the alternative termite hosts Macrotermes subhyalinus and Macrotermes natalensis. While Termitomyces associated with these alternative hosts are horizontally transmitted and recombine freely, the genetic population structure of the same Termitomyces associated with M. bellicosus is consistent with predominantly clonal reproduction and only occasional recombination. This implies that the genetic population structure of Termitomyces is controlled by the termite host and not by the Termitomyces symbiont.     

Sur le nid et la biologie deMacrotermes gilvus Holmgr. Dans les rizières du Cambodge

Résumé Macrotermes gilvus est fréquent dans les rizières de la plaine du Mékong, inondées plusieurs mois chaque année. L'architecture des nids est décrite et comparée à celle d'autres régions.Le problème de l'alimentation des Termites pendant les hautes eaux est examiné; il semble que les meules à champignons jouent le rôle de réserves alimentaires, permettant la survie des sociétés.

---

Summary Macrotermes gilvus is common in the paddy-fields of the Mekong plain which are inondated for several months of the year. The architecture of the nests is described and compared with that of nests from the other regions.The problem of the food supply of these Termites during floods is examined. The fungus gardens appear to be food reserves that enable the survival of the colonies.

     

Ocurrence of the antibiotic producing bacterium Burkholderia sp. in colonies of the leaf-cutting ant Atta sexdens rubropilosa

Fungus garden material from recently established Atta sexdens rubropilosa colonies (6-12 months old) was sampled to detect antibiotic producing microorganisms that inhibited the growth of pathogens of insects and of the fungus gardens but did not affect their mutualistic fungus. A bacterium with activity against the entomopathogenic fungus Beauveria bassiana was isolated from 56% of the gardens tested (n=57) and identified from its biochemical profile and from 16S and 23S ribosomal DNA sequences as a member of the genus Burkholderia. The ant-associated Burkholderia isolates secreted a potent, anti-fungal agent that inhibited germination of conidia of the entomopathogenic fungi B. bassiana, Metarhizium anisopliae, of the saprophytic Verticillium lecanii, and also of a specialist fungus garden Escovopsis weberi. Growth of the ant's mutualist fungus was unaffected.     

Microbial Community Structure of Leaf-Cutter Ant Fungus Gardens and Refuse Dumps

Background

Leaf-cutter ants use fresh plant material to grow a mutualistic fungus that serves as the ants'' primary food source. Within fungus gardens, various plant compounds are metabolized and transformed into nutrients suitable for ant consumption. This symbiotic association produces a large amount of refuse consisting primarily of partly degraded plant material. A leaf-cutter ant colony is thus divided into two spatially and chemically distinct environments that together represent a plant biomass degradation gradient. Little is known about the microbial community structure in gardens and dumps or variation between lab and field colonies.

Methodology/Principal Findings

Using microbial membrane lipid analysis and a variety of community metrics, we assessed and compared the microbiota of fungus gardens and refuse dumps from both laboratory-maintained and field-collected colonies. We found that gardens contained a diverse and consistent community of microbes, dominated by Gram-negative bacteria, particularly γ-Proteobacteria and Bacteroidetes. These findings were consistent across lab and field gardens, as well as host ant taxa. In contrast, dumps were enriched for Gram-positive and anaerobic bacteria. Broad-scale clustering analyses revealed that community relatedness between samples reflected system component (gardens/dumps) rather than colony source (lab/field). At finer scales samples clustered according to colony source.

Conclusions/Significance

Here we report the first comparative analysis of the microbiota from leaf-cutter ant colonies. Our work reveals the presence of two distinct communities: one in the fungus garden and the other in the refuse dump. Though we find some effect of colony source on community structure, our data indicate the presence of consistently associated microbes within gardens and dumps. Substrate composition and system component appear to be the most important factor in structuring the microbial communities. These results thus suggest that resident communities are shaped by the plant degradation gradient created by ant behavior, specifically their fungiculture and waste management.     

The dynamics of plant cell-wall polysaccharide decomposition in leaf-cutting ant fungus gardens

The degradation of live plant biomass in fungus gardens of leaf-cutting ants is poorly characterised but fundamental for understanding the mutual advantages and efficiency of this obligate nutritional symbiosis. Controversies about the extent to which the garden-symbiont Leucocoprinus gongylophorus degrades cellulose have hampered our understanding of the selection forces that induced large scale herbivory and of the ensuing ecological footprint of these ants. Here we use a recently established technique, based on polysaccharide microarrays probed with antibodies and carbohydrate binding modules, to map the occurrence of cell wall polymers in consecutive sections of the fungus garden of the leaf-cutting ant Acromyrmex echinatior. We show that pectin, xyloglucan and some xylan epitopes are degraded, whereas more highly substituted xylan and cellulose epitopes remain as residuals in the waste material that the ants remove from their fungus garden. These results demonstrate that biomass entering leaf-cutting ant fungus gardens is only partially utilized and explain why disproportionally large amounts of plant material are needed to sustain colony growth. They also explain why substantial communities of microbial and invertebrate symbionts have evolved associations with the dump material from leaf-cutting ant nests, to exploit decomposition niches that the ant garden-fungus does not utilize. Our approach thus provides detailed insight into the nutritional benefits and shortcomings associated with fungus-farming in ants.     

Leucoagaricus gongylophorus Produces Diverse Enzymes for the Degradation of Recalcitrant Plant Polymers in Leaf-Cutter Ant Fungus Gardens

Plants represent a large reservoir of organic carbon comprised primarily of recalcitrant polymers that most metazoans are unable to deconstruct. Many herbivores gain access to nutrients in this material indirectly by associating with microbial symbionts, and leaf-cutter ants are a paradigmatic example. These ants use fresh foliar biomass as manure to cultivate gardens composed primarily of Leucoagaricus gongylophorus, a basidiomycetous fungus that produces specialized hyphal swellings that serve as a food source for the host ant colony. Although leaf-cutter ants are conspicuous herbivores that contribute substantially to carbon turnover in Neotropical ecosystems, the process through which plant biomass is degraded in their fungus gardens is not well understood. Here we present the first draft genome of L. gongylophorus, and, using genomic and metaproteomic tools, we investigate its role in lignocellulose degradation in the gardens of both Atta cephalotes and Acromyrmex echinatior leaf-cutter ants. We show that L. gongylophorus produces a diversity of lignocellulases in ant gardens and is likely the primary driver of plant biomass degradation in these ecosystems. We also show that this fungus produces distinct sets of lignocellulases throughout the different stages of biomass degradation, including numerous cellulases and laccases that likely play an important role in lignocellulose degradation. Our study provides a detailed analysis of plant biomass degradation in leaf-cutter ant fungus gardens and insight into the enzymes underlying the symbiosis between these dominant herbivores and their obligate fungal cultivar.     

Symbiotic fungi produce laccases potentially involved in phenol degradation in fungus combs of fungus-growing termites in Thailand

Fungus-growing termites efficiently decompose plant litter through their symbiotic relationship with basidiomycete fungi of the genus Termitomyces. Here, we investigated phenol-oxidizing enzymes in symbiotic fungi and fungus combs (a substrate used to cultivate symbiotic fungi) from termites belonging to the genera Macrotermes, Odontotermes, and Microtermes in Thailand, because these enzymes are potentially involved in the degradation of phenolic compounds during fungus comb aging. Laccase activity was detected in all the fungus combs examined as well as in the culture supernatants of isolated symbiotic fungi. Conversely, no peroxidase activity was detected in any of the fungus combs or the symbiotic fungal cultures. The laccase cDNA fragments were amplified directly from RNA extracted from fungus combs of five termite species and a fungal isolate using degenerate primers targeting conserved copper binding domains of basidiomycete laccases, resulting in a total of 13 putative laccase cDNA sequences being identified. The full-length sequences of the laccase cDNA and the corresponding gene, lcc1-2, were identified from the fungus comb of Macrotermes gilvus and a Termitomyces strain isolated from the same fungus comb, respectively. Partial purification of laccase from the fungus comb showed that the lcc1-2 gene product was a dominant laccase in the fungus comb. These findings indicate that the symbiotic fungus secretes laccase to the fungus comb. In addition to laccase, we report novel genes that showed a significant similarity with fungal laccases, but the gene product lacked laccase activity. Interestingly, these genes were highly expressed in symbiotic fungi of all the termite hosts examined.