三种白蚁的头胸部及消化道结构特征比较

【目的】白蚁具有降解木质纤维素的能力,主要场所在消化道内,因此消化道形态结构的研究具有重要意义。【方法】本文对采自河南省的3种白蚁(平额散白蚁Reticulitermes planifrons Li et Ping、湖南散白蚁Reticulitermes hunanensis Tsai et Peng和双工土白蚁Odontotermes dimorphus Li et Xiao的兵蚁和工蚁头胸部特征及其消化道结构进行了比较。【结果】两种散白蚁兵蚁的头壳、后颏和上颚没有明显差异,但与双工土白蚁差异显著。3种白蚁工蚁的前肠、后肠及消化道平均长度均大于兵蚁,且差异显著(P<0.05),而中肠的差异不显著;双工土白蚁的肠道总长度工蚁/兵蚁比值(1.4倍)大于两种散白蚁(1.1倍);除了嗉囊和囊形胃外,两种散白蚁工蚁的其余消化道各段差异显著(P<0.05),与双工土白蚁相比差异也显著。【结论】工蚁的消化道比兵蚁发达,散白蚁的消化道结构与土白蚁有较大不同,这些结果与工蚁和兵蚁的社会分工有关,同时也表明高等白蚁与低等白蚁相比不仅是采食习惯的差别,自身消化系统结构与分区也有较大不同,并为白蚁的生物防治和生物质资源利用提供了资料。     

两种鼻白蚁之间的入侵关系

设计了5个实验以研究台湾乳白蚁(Coptotermes formosanus)和黄肢散白蚁(Reticulitermes flavipes)的种间斗争关系。所试白蚁来自阿拉巴马州。实验1将两组白蚁的兵蚁和工蚁以自然比例混合(台湾乳白蚁工蚁：兵蚁＝20：5,黄肢散白蚁工蚁：兵蚁＝24：1);实验2观察了双方同数量(各25头)工蚁的种间斗争能力;实验3观察两者同数量(各5头)兵蚁的种间斗争关系;实验4、5分别研究双方工蚁与兵蚁的斗争关系。因两种工蚁形态相似,所以在实验前用0．1％Nile blue A染过的滤纸饲喂一方白蚁24h使虫体着色以便观察。所有实验在22℃,65％RH荧光灯下进行。每个实验将两方白蚁同时放入一培养皿中,观察至一方100％受伤,然后把培养皿移到培养箱内,24小时后统计死亡率。结果表明,实验一开始,双方的工蚁和兵蚁都立即进入残酷的厮杀斗争,台湾乳白蚁兵蚁和工蚁的斗争能力都明显强于黄肢散白蚁;兵蚁的斗争行为不受工蚁的影响,工蚁的斗争能力也不受兵蚁的制约。实验中台湾乳白蚁的工蚁与黄肢散白蚁的兵蚁攻击能力相当,但黄肢散白蚁的工蚁却远弱于台湾乳白蚁的兵蚁。上述结果说明,自然界中不可能出现此两种白蚁群体的偶然性融合;外来的台湾乳白蚁很可能在其群体建立的地区占据优势[动物学报49(3)：295—302,2003]。     

尖唇散白蚁兵蚁和工蚁卵巢和卵子形态特征的比较

兵蚁和工蚁是白蚁中的非生殖品级,兵蚁是由工蚁分化产生。为了探讨兵蚁品级性腺不育的原因以及兵蚁和工蚁性腺发育的差异,采用组织学染色观察与测量方法对尖唇散白蚁Reticulitermes aculabialis Tsai et Hwang的兵蚁和工蚁的卵子发生各阶段进行比较和分析。结果显示:两者性腺发育大小呈极显著性差异(P<0.01),兵蚁与工蚁卵巢横切面面积之比约为1∶7;兵蚁的卵子发生与工蚁相比,仅有卵母细胞的分化期,没有生长期;兵蚁的卵母细胞比工蚁的小,两者的分化期卵母细胞体积之比约为1∶16。该结果表明工蚁向兵蚁转化过程中性腺进一步退化,兵蚁性腺极度退化使其丧失了成为补充繁殖蚁的可能性。该研究结果为兵蚁不能生殖提供组织学上的证据,同时又揭示了兵蚁和工蚁在潜在生殖能力差异方面的组织学基础。     

白蚁取食行为多型及其机理

对白蚁取食行为的研究涉及5科18属38种白蚁。其取食行为多型可归纳为3类:(1)工蚁主导型:采食活动基本上由工蚁完成,兵蚁不参与或仅极少数个体参与,食物侦查或保护采食工蚁的职能不显著;(2)工蚁-兵蚁协同型:这是白蚁采食组织的常见类型,兵蚁成为采食队伍的固定成员,负责采食活动的安全侦查与防卫、食物源侦查等工作;(3)假工蚁主导型:由于尚未分化形成专职的工蚁,因而大龄幼蚁(广义的假工蚁)成为采食活动的主力,同时兼有巢内的保护职能。在食物加工和分配过程中,低等白蚁的幼蚁在未获得消化纤维素能力前主要通过肛交哺获得营养物质和共生物,且肛交哺是新蜕皮个体重新获得共生鞭毛虫的途径。高等白蚁幼龄工蚁负责食物加工、依赖型品级通过口交哺获得营养物质。白蚁采食活动中的行为多型受踪迹信息素控制,不同品级个体腹板腺有明显分化;不同品级和发育阶段个体食性分化与消化道结构、消化酶活性密切相关;保幼激素影响交哺行为以及消化酶基因表达。本文在评述研究进展的同时,对进一步研究提出了展望。     

黑翅土白蚁触角感器的扫描电镜观察

【目的】通过黑翅土白蚁Odontotermes fomosanus(Shiraki)幼蚁、工蚁、兵蚁触角感器种类、数量及分布的研究,分析不同龄幼蚁、工蚁和兵蚁的触角感器特征,为进一步研究触角感器发育特点及不同个体的行为分化机制提供参考。【方法】通过扫描电镜观察触角形态,分析触角节数、感器种类、数量及分布特征。【结果】工蚁、兵蚁、幼蚁个体触角长度差异显著,触角长度顺序:工蚁>兵蚁>3龄幼蚁>2龄幼蚁>1龄幼蚁;触角感器共有9种类型,分别为短毛形感器、长毛形感器、短刺形感器、长刺形感器、锥状B?hm氏鬃毛、球状钟形感器、刺突状钟形感器、球状钟形感器与刺突状钟形感器中间型、长刺具泡状感器。工蚁、兵蚁缺少长刺具泡状感器。幼蚁触角感器种类和数量随龄数而增加,1龄幼蚁具7种感器,缺少球状钟形感器和球状钟形感器与刺突状钟形感器中间型2种类型;2龄幼蚁具有8种感器,缺球状钟形感器;3龄幼蚁具有全部9种类型感器。【结论】黑翅土白蚁幼蚁、工蚁、兵蚁触角节数和感器显著差异,可作为幼蚁龄数以及非生殖品级判别的形态指标,应用于白蚁非生殖品级行为多型等相关研究。     

保幼激素类似物对白蚁的作用研究进展

本文介绍了保幼激素类似物对白蚁的影响,包括诱导前兵蚁、兵蚁和一些中间品级(形态畸形)的产生、抑制白蚁的取食、减少或消除白蚁的共生原生动物群、对白蚁产生不同水平的急性和慢性毒性、抑制工蚁蜕皮以及繁殖蚁建立新群体等方面的影响,其中最主要影响是诱导前兵、兵蚁和一些中间品级的产生破坏了白蚁群体品级比例的平衡性或完整性.因此,保幼激素类似物可用于白蚁的防治,并在田问防治散白蚁和乳白蚁均取得了较好的防治效果,并探讨了应用保幼激素类似物防治白蚁的潜力与前景.     

尖唇散白蚁工蚁、兵蚁和繁殖蚁精子发生的比较研究

为探讨白蚁非生殖品级和生殖品级生殖细胞发育差异,采用组织学染色技术对尖唇散白蚁Reticulitermes aculabialis Tsaiet Hwang繁殖蚁、工蚁和兵蚁的精巢发育以及精子发生进行了显微观察和比较研究。结果发现3个品级间精巢发育的程度差异很大,三者精巢切面面积相对大小之比为:繁殖蚁∶工蚁=1.7∶1;繁殖蚁∶兵蚁=29.3∶1;工蚁∶兵蚁=17.1∶1。繁殖蚁和工蚁精巢管内有精子的形成,工蚁和繁殖蚁精子的发生都经历了精原细胞、初级精母细胞、次级精母细胞、精细胞和精子时期,但工蚁有大量次级精母细胞呈细胞凋亡状态。兵蚁生殖细胞发育仅有精原细胞、初级精母细胞、次级精母细胞,没有精细胞和精子产生。工蚁的生殖细胞显著小于同一时期繁殖蚁的生殖细胞,兵蚁的各时期生殖细胞均极显著小于繁殖蚁同一时期的生殖细胞。研究表明各品级之间生殖功能分化与生殖细胞发育有直接关系,工蚁有转化为补充繁殖蚁和兵蚁的能力;而兵蚁由于精巢极度退化不能产生精细胞和精子,因此是非生殖品级分化的终极形式,不具有转化成补充繁殖蚁或其它品级的能力。     

黑胸散白蚁幼期不同品级的发育和分化

在黑胸散白蚁Reticulitermes chinensis发育和分化过程中,发现有假工蚁、假若蚁两种虫态。假工蚁由6龄和7龄工蚁转化发育而来,假若蚁由4龄和5龄若蚁转化发育而来。显微测量结果表明,黑胸散白蚁的胚后发育主要包括2个龄期的幼蚁期、6个以上龄期的工蚁期、4个龄期的若蚁期和有翅成虫。在此基础上分析了其他虫态的分化来源,发现兵蚁由3～7龄工蚁分化发育而来,翅鳞型和长翅芽型补充生殖蚁由6龄若蚁转化发育而来,短翅芽型补充生殖蚁由4龄和5龄若蚁转化发育而来,微翅芽型补充生殖蚁既可由4～6龄工蚁转化发育而来,又可由假工蚁和假若蚁转化发育而来,无翅型补充生殖蚁由3～7龄工蚁转化发育而来。提出了黑胸散白蚁群体中不同品级个体的可能分化途径。     

黑胸散白蚁幼期不同品级的发育和分化

在黑胸散白蚁Reticulitermes chinensis发育和分化过程中,发现有假工蚁、假若蚁两种虫态。假工蚁由6龄和7龄工蚁转化发育而来,假若蚁由4龄和5龄若蚁转化发育而来。显微测量结果表明,黑胸散白蚁的胚后发育主要包括2个龄期的幼蚁期、6个以上龄期的工蚁期、4个龄期的若蚁期和有翅成虫。在此基础上分析了其他虫态的分化来源,发现兵蚁由3～7龄工蚁分化发育而来,翅鳞型和长翅芽型补充生殖蚁由6龄若蚁转化发育而来,短翅芽型补充生殖蚁由4龄和5龄若蚁转化发育而来,微翅芽型补充生殖蚁既可由4～6龄工蚁转化发育而来,又可由假工蚁和假若蚁转化发育而来,无翅型补充生殖蚁由3～7龄工蚁转化发育而来。提出了黑胸散白蚁群体中不同品级个体的可能分化途径。     

囟土白蚁蚁后卵巢管数量、蚁巢体积与蚁群中各蚁型数量的相关性

囟土白蚁Odontotermes fontanellus Kemner是安徽为害林木、水库堤坝的主要蚁种。研究结果表明:1.在一定的巢龄范围内,随着蚁后身体增长,卵巢管数量增多,一侧的卵巢管最多可达3726条。2.蚁后体长与工蚁数、兵蚁数、幼蚁数、整巢蚁数之间成幂函数关系。3.卵巢管数与蚁后体长、工蚁数、兵蚁数、幼蚁数、整巢蚁数之间成幂函数关系,4,蚁巢体积(y)与蚁后体长(X_2)、兵蚁数(X_4)、的关系式为:y=6.687e~(0.0566)X_2+0.6741X_4     

Association between caste and genotype in the termite Mastotermes darwiniensis Froggatt (Isoptera: Mastotermitidae)

Abstract Termite workers and soldiers differ markedly in their morphology and behaviour. We sought evidence for genetic influences on caste determination in the giant northern termite, Mastotermes darwiniensis , by investigating if workers and soldiers from the same colony differed genetically. The genotypes of 795 termites from 11 distinct colonies were assayed at six polymorphic microsatellite loci. We found that the multilocus genotypes of workers and soldiers from 8 of the 11 colonies did not differ significantly. Thus, the majority of the data provided no evidence for a genetic association with caste in workers and soldiers of M. darwiniensis . However, the genotype frequencies of workers and soldiers from three colonies differed, suggesting that genotype is occasionally associated with caste in this species. The genetic differentiation of castes within these colonies could reflect differences in the propensities of termites with distinct genotypes to develop into particular castes and provide a selective advantage to colonies headed by multiple reproductives.     

Comparison of the gut microbiota from soldier and worker castes of the termite Reticulitermes grassei

The bacterial microbiota from the whole gut of soldier and worker castes of the termite Reticulitermes grassei was isolated and studied. In addition, the 16S rDNA bacterial genes from gut DNA were PCR-amplified using Bacteria-selective primers, and the 16S rDNA amplicons subsequently cloned into Escherichia coli. Sequences of the cloned inserts were then used to determine closest relatives by comparison with published sequences and with sequences from our previous work. The clones were found to be affiliated with the phyla Spirochaetes, Proteobacteria, Firmicutes, Bacteroidetes, Actinobacteria, Synergistetes, Verrucomicrobia, and candidate phyla Termite Group 1 (TG1) and Termite Group 2 (TG2). No significant differences were observed with respect to the relative bacterial abundances between soldier and worker phylotypes. The phylotypes obtained in this study were compared with reported sequences from other termites, especially those of phylotypes related to Spirochaetes, Wolbachia (an Alphaproteobacteria), Actinobacteria, and TG1. Many of the clone phylotypes detected in soldiers grouped with those of workers. Moreover, clones CRgS91 (soldiers) and CRgW68 (workers), both affiliated with 'Endomicrobia', were the same phylotype. Soldiers and workers also seemed to have similar relative protist abundances. Heterotrophic, poly-β-hydroxyalkanoate-accumulating bacteria were isolated from the gut of soldiers and shown to be affiliated with Actinobacteria and Gammaproteobacteria. We noted that Wolbachia was detected in soldiers but not in workers. Overall, the maintenance by soldiers and workers of comparable axial and radial redox gradients in the gut is consistent with the similarities in the prokaryotes and protists comprising their microbiota.     

Colony-dependent sex differences in protozoan communities of the lower termite <Emphasis Type="Italic">Reticulitermes speratus</Emphasis> (Isoptera: Rhinotermitidae)

In many animals, sex differences in hormones, behavior, and immunity lead to differences in their gut microbial communities. One of the best-known examples of mutualistic symbiosis is that between lower termites and their intestinal protozoa. Although differences in the protozoan communities of different castes have been studied in lower termites, nothing is known about the sex differences in protozoan communities in neuter castes. Here, we show that termite workers have different protozoan communities according to sex depending on the colony. We investigated the communities of symbiotic protozoa living in lower termites, Reticulitermes speratus, and how they are affected by sex and caste. Workers had the largest numbers of protozoa, followed by soldiers, whereas reproductives (primary kings and secondary queens) had no protozoa. Workers showed colony-dependent sex differences in the total abundance of protozoa, whereas soldiers showed no such sex differences. There were significant sex effect and/or interaction effect between colony and sex in abundances of five species of protozoa in workers. Workers also showed significant sex differences and/or colony-dependent sex differences in proportion of six species of protozoa. These may result in sex differences in the host–symbiont interaction due to physiological or behavioral sex differences in workers that have not been recognized previously. This study has an important implication: although workers are not engaged in reproduction, their potential sex difference may affect various aspects of social interactions.     

Intracolonial differences in gut bacterial community between worker and soldier castes of Coptotermes formosanus

Abstract The establishment of symbiotic relationships with intestinal microorganisms enables termites to thrive on recalcitrant substrates such as cellulose and wood. A termite colony is composed of several different castes which have distinct feeding habits. The soldiers, for example, cannot feed by themselves and depend on workers, who feed them with digested or semi‐digested foods. To investigate the influence of feeding habits on the bacterial symbionts, a comparative study of gut bacteria between worker and soldier castes of the termite Coptotermes formosanus was conducted. The bacterial communities of both castes were investigated using denaturing gradient gel electrophoresis (DGGE) and clonal analysis of 16S ribosomal DNA (rDNA). Both methods indicated Bacteroidetes was the common predominant group; the common dominant phylotype was affiliated with a reported uncultured Bacteroidetes phylotype (BCf1–03). There were significant differences in Bacteroidetes and Spirochaetes between two castes. Compared to the gut bacteria of workers, those of soldiers were lower in abundance and diversity of Bacteroidetes and slightly higher in Spirochaetes. Two phylotypes (W8, W11) affiliated to Bacteroidetes and two (W26, W29) affiliated to Spirochaetes were exclusively found in the DGGE profile of the worker caste. Bacteroidetes are assumed to be involved in fermentation of sugars and nitrogenous compounds as well as degradation of uric acid. Spirochaetes are supposed to aid in the functions of acetogenesis and N₂‐fixation. The different feeding habits between workers and soldiers of C. formosanus may explain the observed differences in the gut bacterial community.     

Hunting strategy of a generalist ant species proposed as a biological control agent against termites

We studied the hunting behaviour of Myrmicaria opaciventris (Hymenoptera: Formicidae) in order to evaluate if it can be used as a biological control agent against the termites that damage sugarcane plantations. Hunting workers foraged in groups and recruited nestmates at short-range when they encountered large termite soldiers or groups of small termite workers. Differences in prey capture concerned the: (1) means of detection (from a distance or by contact); (2) termite body part seized (small termites seized by the body; large termites by an appendage); (3) percentages of prey abandoned; and (4) use of venom. The sting of the workers is spatulated implying a topical application of the venom on the prey. Large termites were stretched by several workers whose adherence to the substrate is facilitated by well-developed arolia and claws on the legs while others spread venom on the body and carved it up. An adaptation to termite capture was noted with a distribution of tasks between the workers which subdued prey, and those which transported it. In the former case, the workers easily eliminated termite soldiers, successively attacked several termite workers and even captured new individuals while holding the first ones captured between their mandibles before retrieving them all at once. The remaining individuals were retrieved by the transporting workers. Given this particularly effective predatory strategy, we concluded that, under certain conditions, M. opaciventris can be used as a biological control agent against termites.     

Worker aggression of ant Lasius japonicus enhanced by termite soldier–specific secretion as an alarm pheromone of Reticulitermes speratus

The subterranean termite Reticulitermes speratus usually nests in rotten wood trunks, which may also be occupied by the Japanese garden ant Lasius japonicus. Few battles were observed between them under ordinary circumstances because they inhabit separate nesting sites. However, once the termite nesting sites were artificially broken, the ant workers invaded and hunted the termites, although the termite soldiers fought against the ants. This study aims to confirm intra‐ and inter‐specific chemical interactions between the termite and ant. Solid phase microextraction–gas chromatograph (SPME‐GC) analyses revealed that R. speratus soldiers secreted caste‐specific sesquiterpene hydrocarbon when they were irritated. Both the hexane extract of the soldiers and its hydrocarbon fraction, as well as the crushed soldier bodies, attracted the soldiers but dispersed the workers when presented on the trails. We also confirmed that the soldier chemicals enhanced aggressiveness of L. japonicus, which rushed around the odor sources and hunted any termites that were present. These findings suggest that: (i) the soldier–specific secretion might serve as an alarm pheromone in termite chemical communication, in which components recruit soldiers and also warn the other colony members away; and (ii) termite communication is eavesdropped on by L. japonicus workers to locate and hunt the termites.     

Sexual difference in juvenile-hormone titer in workers leads to sex-biased soldier differentiation in termites

In termites, the soldier caste, with its specialized defensive morphology, is one of the most important characteristics for sociality. Most of the basal termite species have both male and female soldiers, and the soldier sex ratio is almost equal or only slightly biased. However, in the apical lineages (especially family Termitidae), there are many species that have soldiers with strongly biased sex ratio. Generally in termites, since high juvenile hormone (JH) titer is required for soldier differentiation from a worker via a presoldier stage, it was hypothesized that the biased soldier-sex ratio was caused by differences in JH sensitivity and/or JH titer between male and female workers. Therefore, we focused on the presoldier differentiation and the worker JH titer in species with only male soldiers (Nasutitermes takasagoensis) and with both male and female soldiers (Reticulitermes speratus) in natural conditions. In the former species, there are four types of workers; male minor, male medium, female medium and female major workers, and presoldiers differentiate from male minor workers. First, we tried to artificially induce presoldiers from male and female workers. In N. takasagoensis, the presoldier differentiation rate and mortality was significantly higher in male minor workers. Morphological analyses showed that both male and female induced presoldiers possessed normal soldier-specific morphologies. It was suggested that female workers, from which soldiers do not differentiate under natural conditions, also maintained the physiological and developmental potential for soldier differentiation. In R. speratus, however, no differences were observed in solder differentiation rate and mortality between male and female workers. Second, the JH titers of each sex/type of workers were quantified by high performance liquid chromatography–mass spectrometry in two different seasons (April and December). The results showed that, in N. takasagoensis, JH titer in male minor workers was consistently higher than those in other worker types. In R. speratus, in contrast, there were no significant differences in JH titers between male and female workers. These results suggested that, in N. takasagoensis, male minor workers maintain JH titers at a high level throughout a year, and this may cause the male-biased presoldier differentiation.     

Regulation of juvenile hormone titers by soldiers in the Formosan subterranean termite, Coptotermes formosanus

In field collections of the Formosan subterranean termite Coptotermes formosanus, soldiers averaged less than 10%. The proportion of soldiers increased to about 25% or higher when termites were kept in the laboratory, as did the juvenile hormone III (JH III) titers for both workers and soldiers. In laboratory experiments with a proportion of soldiers 25% or higher, very few new pre-soldiers were formed and the JH titer in existing soldiers remained constant. On the contrary, workers from groups containing less than 25% soldiers formed more pre-soldiers. In such cases, both workers and soldiers showed higher JH titers. Newly formed soldiers also had higher JH titers than older soldiers. It is speculated that populations containing higher soldier proportions inhibit further soldier differentiation, eventually holding the JH titers under the threshold or suppressing the activity of corpora allata resulting in lower JH titers in workers.