Young but not defenceless: antifungal activity during embryonic development of a social insect

Termites live in environments heavily colonized by diverse microorganisms, including pathogens. Eggs laid within the nest are likely to experience similar pathogenic pressures as those experienced by older nest-mates. Consequently, eggs may be under selective pressures to be immune-competent. Through in vitro experiments using developing embryos of the dampwood termite, Zootermopsis angusticollis , we tested the ontogeny, location and strength of their antifungal activity against the fungus, Metarhizium brunneum . Exterior washes of the chorion (extra-chorionic) and components within the chorion (intra-chorionic) were incubated with fungal conidia, which were then scored for viability. The fungistatic activity was location and developmental stage dependent. Extra-chorionic washes had relatively weak antifungal activity. Intra-chorionic homogenates were highly antifungal, exhibiting increased potency through development. The positive correlation between intra-chorionic fungistasis and developmental stage is probably due to the expression of endogenous proteins during embryogenesis. Boiling of both the extra-chorionic washes and the intra-chorionic contents rescued conidia viability, indicating the antifungal agent(s) is (are) heat-sensitive and probably proteinaceous. This study is the first to address embryonic antifungal activity in a hemimetabolous, eusocial taxon. Our results support the hypothesis that microbes have been significant agents of selection in termites, fostering the evolution of antifungal properties even in the most immature stage of development.

Who goes there? Social surveillance as a response to intergroup conflict in a primitive termite

Intergroup conflict has been suggested as a major force shaping the evolution of social behaviour in animal groups. A long-standing hypothesis is that groups at risk of attack by rivals should become more socially cohesive, to increase resilience or protect against future attack. However, it is usually unclear how cohesive behaviours (such as grooming or social contacts) function in intergroup conflict. We performed an experiment in which we exposed young colonies of the dampwood termite, Zootermopsis angusticollis , to a rival colony while preventing physical combat with a permeable barrier. We measured social contacts, allogrooming and trophallaxis before, during and after exposure. Termites showed elevated rates of social contacts during exposure to a rival compared to the pre-exposure phase, but rates returned to pre-exposure levels after colonies were separated for 9 days. There was evidence of a delayed effect of conflict on worker trophallaxis. We suggest that social contacts during intergroup conflict function as a form of social surveillance, to check individual identity and assess colony resource holding potential. Intergroup conflict may increase social cohesion in both the short and the long term, improving the effectiveness of groups in competition.

Relish as a Candidate Marker for Transgenerational Immune Priming in a Dampwood Termite (Blattodae: Archeotermopsidae)

Natural selection should favor the transfer of immune competence from one generation to the next in a context-dependent manner. Transgenerational immune priming (TGIP) is expected to evolve when species exploit pathogen-rich environments and exhibit extended overlap of parent–offspring generations. Dampwood termites are hemimetabolous, eusocial insects (Blattodea: Archeotermopsidae) that possess both of these traits. We predict that offspring of pathogen-exposed queens of Zootermopsis angusticollis will show evidence of a primed immune system relative to the offspring of unexposed controls. We found that Relish transcripts, one of two immune marker loci tested, were enhanced in two-day-old embryos when laid by Serratia-injected queens. These data implicate the immune deficiency (IMD) signaling pathway in TGIP. Although an independent antibacterial assay revealed that embryos do express antibacterial properties, these do not vary as a function of parental treatment. Taken together, Z. angusticollis shows transcriptional but not translational evidence for TGIP. This apparent incongruence between the transcriptional and antimicrobial response from termites suggests that effectors are either absent in two-day-old embryos or their activity is too subtle to detect with our antibacterial assay. In total, we provide the first suggestive evidence of transgenerational immune priming in a termite.

Global elevational, latitudinal, and climatic limits for termites and the redescription of Rugitermes laticollis Snyder (Isoptera: Kalotermitidae) from the Andean Highlands

We compile, map, and discuss global elevational, latitudinal, thermal, and rainfall extremes of termite localities from literature sources and unpublished records. Rugitermes laticollis from Ecuador and Bolivia occurs at higher elevation (2700-3600 m) than any other termite species. Termites span the globe from 54.3°N (Zootermopsis angusticollis in British Columbia (B.C.), Canada) to 48.9°S (Porotermes quadricollis in Magdalena, Chile). The coldest locality supporting termites (Reticulitermes sp.) is at Churn Creek, B.C., where the mean annual temperature is 4°C. Lake Havasu City, Arizona, where Heterotermes aureus and Gnathamitermes perplexus occur, has the highest recorded temperature maximum (52°C) for a termite locality. Cryptotermes brevis and Neotermes chilensis are endemic to the Pacific Coast of Peru and Chile where rain is essentially absent. We further provide locality extremes for six termite families from six zoogeographical regions. In addition, the winged imago of Ru. laticollis is redescribed and the soldier is described for the first time.