Termites are model social organisms characterized by a polyphenic caste system. Subterranean termites (Rhinotermitidae) are ecologically and economically important species, including acting as destructive pests. Rhinotermitidae occupies an important evolutionary position within the clade representing a transitional taxon between the higher (Termitidae) and lower (other families) termites. Here, we report the genome, transcriptome, and methylome of the Japanese subterranean termite Reticulitermes speratus. Our analyses highlight the significance of gene duplication in social evolution in this termite. Gene duplication associated with caste-biased gene expression was prevalent in the R. speratus genome. The duplicated genes comprised diverse categories related to social functions, including lipocalins (chemical communication), cellulases (wood digestion and social interaction), lysozymes (social immunity), geranylgeranyl diphosphate synthase (social defense), and a novel class of termite lineage–specific genes with unknown functions. Paralogous genes were often observed in tandem in the genome, but their expression patterns were highly variable, exhibiting caste biases. Some of the assayed duplicated genes were expressed in caste-specific organs, such as the accessory glands of the queen ovary and the frontal glands of soldier heads. We propose that gene duplication facilitates social evolution through regulatory diversification, leading to caste-biased expression and subfunctionalization and/or neofunctionalization conferring caste-specialized functions.
We demonstrate time-domain sampling of mid-infrared electric field transients and their conjugate counterparts exploiting the dynamical Pockels effect. To this end, the complete polarization change of few-femtosecond probe pulses is studied. An intuitive picture based on a phasor representation is established before gaining quantitative understanding in experiment and theory. In the standard version of electro-optic sampling, the electric field is determined by analyzing the change of ellipticity of the probe polarization. Beyond this, we find that a temporal gradient of the input electric field manifests itself in a rotation of the polarization ellipsoid of the probe. The relative contribution of sum-and difference-frequency mixing processes and their spectral distribution over the near-infrared probe bandwidth are identified as key aspects. If one of these processes dominates, detecting ellipticity changes and polarization rotation as a function of time delay results in two wave forms which are Hilbert transforms of each other. Such conditions may be achieved by angle phase matching in birefringent materials or spectral filtering of the probe after the nonlinear interaction. In this case, a static phase introduced by birefringence or reflection at metallic mirrors results in a specific phase shift of both time traces with respect to the input electric field. Contributions from sum-and difference-frequency generation are found to be equivalent when using electro-optic sensors with isotropic refractive index. Polarization rotations in the lowand high-frequency parts of the probe then tend to cancel out. In this limit, spurious additional phase shifts do not change the phase of the detected transients. This fact leads to a robust recovery of the carrier-envelope phase of the input wave form. Clarifying the role of imperfections of superachromatic phase retarders completes our survey on proper determination of the electric field and its conjugate variable.
Insects represent the biodiversity of the terrestrial ecosystem, and their prosperity is attributable to their association with symbiotic microorganisms. By sequestering microbial functionality into their bodies, organs, tissues, or cells, diverse insects have successfully exploited otherwise inaccessible ecological niches and resources, including herbivory enabled by utilization of indigestible plant cell wall components. In leaf beetles of the subfamily Cassininae, an ancient symbiont lineage,
Stammera
, whose genome is extremely reduced and specialized for encoding pectin-degrading enzymes, is hosted in gut-associated symbiotic organs and contributes to the host’s food plant digestion.
Microorganisms often live in symbiosis with their hosts, and some are considered mutualists, where all species involved benefit from the interaction. How free-living microorganisms have evolved to become mutualists is unclear. Here we report an experimental system in which non-symbiotic Escherichia coli evolves into an insect mutualist. The stinkbug Plautia stali is typically associated with its essential gut symbiont, Pantoea sp., which colonizes a specialized symbiotic organ. When sterilized newborn nymphs were infected with E. coli rather than Pantoea sp., only a few insects survived, in which E. coli exhibited specific localization to the symbiotic organ and vertical transmission to the offspring. Through transgenerational maintenance with P. stali, several hypermutating E. coli lines independently evolved to support the host’s high adult emergence and improved body colour; these were called ‘mutualistic’ E. coli. These mutants exhibited slower bacterial growth, smaller size, loss of flagellar motility and lack of an extracellular matrix. Transcriptomic and genomic analyses of ‘mutualistic’ E. coli lines revealed independent mutations that disrupted the carbon catabolite repression global transcriptional regulator system. Each mutation reproduced the mutualistic phenotypes when introduced into wild-type E. coli, confirming that single carbon catabolite repression mutations can make E. coli an insect mutualist. These findings provide an experimental system for future work on host–microbe symbioses and may explain why microbial mutualisms are omnipresent in nature.
We present a method of developing a high-speed terahertz time-domain polarimeter based on an electro-optic sampling setup, which consists of a rapid polarization analysis of the probe optical beam within 1 ms with an angular resolution of ∼100 mrad using an electro-optic modulator (EOM). A fast optical delay line enabled us to obtain the vector waveform of the terahertz pulse within 100 ms. The setup does not utilize a manual/mechanical rotational stage, but the use of the polarization modulation technique employing the EOM ensures a high-speed and high-accuracy measurement of the state of polarization of terahertz waves.
In social insects, interactions among colony members trigger caste differentiation with morphological modifications. In termite soldier differentiation, the mandible size considerably increases through two moltings (via the presoldier stage) under the control of juvenile hormone (JH). Regulatory genes are predicted to provide patterning information that induces the mandible-specific cell proliferation. To identify factors responsible for the mandibular enlargement, expression analyses of 18 candidate genes were carried out in the termite Hodotermopsis sjostedti. Among those, dachshund (dac), which identifies the intermediate domain along the proximodistal appendage axis, showed mandible-specific upregulation prior to the molt into presoldiers, which can explain the pattern of cell proliferation for the mandibular elongation. Knockdown of dac by RNAi reduced the mandibular length and distorted its morphology. Furthermore, the epistatic relationships among Methoprene tolerant, Insulin receptor, Deformed (Dfd) and dac were revealed by combined RNAi and qRT-PCR analyses, suggesting that dac is regulated by Dfd, downstream of the JH and insulin signaling pathways. Thus, caste-specific morphogenesis is controlled by interactions between the factors that provide spatial information and physiological status.
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