From descent with modification to the origins of novelty.

Descent with modification is the foundational framework of all of evolution. Yet evolutionary novelties are defined as lacking affinities to structures that already existed in the ancestral state, i.e. to somehow emerge in the absence of homology. We posit that reconciling both perspectives necessitates the existence of a type of innovation gradient that allows descent with modification to seed the initiation of a novel trait, which once in existence can then diversify into its variant forms. Recent work on diverse, textbook examples of morphological novelties illustrate the value of the innovation gradient concept. Innovations as profound and diverse as insect wings, beetle horns, and treehopper helmets derive from homologous source tissues instructed in their development by homologous gene regulatory networks. Yet rather than rendering these traits no longer novel, we posit that discoveries such as these call for a reassessment of the usefulness of defining evolutionary novelty as necessitating the absence of homology. Instead, we need to redirect our attention to how ancestral homologies scaffold and bias the innovation gradient to facilitate hotspots of innovation in some places, and deep conservation elsewhere.

Cuticular Permeability, Percent Body Water Loss, and Relative Humidity Equilibria Comparisons of Four Termite Species.

This study compared percentage of total body water (%TBW), water loss rate under desiccative conditions, and cuticular permeability (CP) of four termite species (Order: Blattodea, Infraorder: Isoptera) from different habitats, including one subterranean termite, Coptotermes formosanus Shiraki (Blattodea: Rhinotermitidae), one dampwood termite, Neotermes jouteli (Banks) (Blattodea: Kalotermitidae), one 'wetwood' termite, Cryptotermes cavifrons Banks (Blattodea: Kalotermitidae), and one drywood termite, Cryptotermes brevis Walker (Blattodea: Kalotermitidae). There was no consistent pattern associated with the habitat relative humidity (RH) levels and the %TBW among the four termite species tested. Because C. formosanus forage for food, its lowest %TBW may be due to its ability to access water sources, and its need to be more mobile than kalotermitids that remain in wood. Body water loss rate and CP were lower for xeric species such as Cr. brevis than hydric species such as N. jouteli. An RH equilibrium represents the capacity of termite water retention mechanisms that may include the physical structure of cuticular layers, behavioral manipulation of natural openings, and physiological means. As with water loss rate and CP data, RH equilibria were lower for xeric species such as Cr. brevis than hydric species such as N. jouteli. Moreover, RH equilibria for dead individuals of the four termite species were significantly higher than those of live individuals, indicating dead termites lost more water after losing their physiological and behavioral means of retaining water. RH equilibria represent termites' capability in water retention, and there is a strong linear relationship between RH equilibria and CP values. Hence, RH equilibria may offer simpler alternatives to CP estimates.

BgFas1: A fatty acid synthase gene required for both hydrocarbon and cuticular fatty acid biosynthesis in the German cockroach, Blattella germanica (L.).

Insect cuticular hydrocarbons (CHCs), the evolutionary products of aquatic hexapod ancestors expanding to terrestrial environment, are deposited on the surface of insect integument and originally functioned primarily as waterproofing agents. CHCs are derived from the conserved fatty acid synthesis pathway in insects. However, the pivotal fatty acid synthase (FAS) involved in hydrocarbon (HC) biosynthesis remains unknown in many insect orders including the primitive Blattodea. Here, we investigated functional FAS genes that modulate cuticular lipid biogenesis in the German cockroach, Blattella germanica (L.). Based on our full-length transcriptomic data and the available genomic data, seven FAS genes (BgFas1-7) were identified from B. germanica. Tissue-specific expression analysis revealed that BgFas1, BgFas3, BgFas4 and BgFas7 were highly expressed in the integument, whereas BgFas2 was dominantly expressed in the fat body. BgFas5/6 mRNA was almost negligible in the tested tissues. Systemic RNAi screen was performed against BgFas1-7, we found that only RNAi knockdown of BgFas1 caused a dramatic reduction of methyl-branched HCs (mbHCs) and a slight decrease of straight-chain HCs (scHCs) for both internal and external HCs. Significant reduction of cuticular free fatty acids (cFFAs) was also detected within BgFas1-repressed cockroaches, while repression of CYP4G19 resulted in dramatic increase of cFFAs. Moreover, we found that BgFas1 mRNA levels were correlated with insect molting cycles, and could be induced by long-term mild dryness treatment. Furthermore, desiccation assay revealed that BgFas1 suppression accelerated water loss and led to early death of cockroaches under desiccation. Our results indicate that BgFas1 is necessary for both HC and cFFA biosynthesis in B. germanica. In addition, our study also confirms that cuticular lipids, particularly mbCHCs, are critical for desiccation resistance in B. germanica.

Cuticular gas exchange by Antarctic sea spiders.

Many marine organisms and life stages lack specialized respiratory structures, like gills, and rely instead on cutaneous respiration, which they facilitate by having thin integuments. This respiratory mode may limit body size, especially if the integument also functions in support or locomotion. Pycnogonids, or sea spiders, are marine arthropods that lack gills and rely on cutaneous respiration but still grow to large sizes. Their cuticle contains pores, which may play a role in gas exchange. Here, we examined alternative paths of gas exchange in sea spiders: (1) oxygen diffuses across pores in the cuticle, a common mechanism in terrestrial eggshells, (2) oxygen diffuses directly across the cuticle, a common mechanism in small aquatic insects, or (3) oxygen diffuses across both pores and cuticle. We examined these possibilities by modeling diffusive oxygen fluxes across all pores in the body of sea spiders and asking whether those fluxes differed from measured metabolic rates. We estimated fluxes across pores using Fick's law parameterized with measurements of pore morphology and oxygen gradients. Modeled oxygen fluxes through pores closely matched oxygen consumption across a range of body sizes, which means the pores facilitate oxygen diffusion. Furthermore, pore volume scaled hypermetrically with body size, which helps larger species facilitate greater diffusive oxygen fluxes across their cuticle. This likely presents a functional trade-off between gas exchange and structural support, in which the cuticle must be thick enough to prevent buckling due to external forces but porous enough to allow sufficient gas exchange.

Mechanical properties of the cuticle of the tick Amblyomma hebraeum (Acari: Ixodidae).

Female ticks of the family Ixodidae increase their mass up to 100-fold during the 7-10 day feeding period. We determined the material properties of the alloscutal cuticle of female Amblyomma hebraeum from the time of moulting through to full engorgement. The material properties of the cuticle were evaluated by a Kelvin-Voigt analysis of compliance determined from the stretch of loops of cuticle under stress. There was a 3-fold increase in cuticle dry mass during the first 3 weeks post-moult, during which the ductility and stiffness of the cuticle increased substantially. Under stress, the cuticle displayed time-dependent stretch, with a plastic (non-recoverable) and viscoelastic (recoverable) component. Plastic deformation was reasonably constant in the range 10-15% over a wide range of induced stress above ∼ 0.6 MPa. The plastic component of tick alloscutal cuticle was about 5-10 times higher than that of unsclerotized insect cuticle. Tick cuticle is far more ductile than unsclerotized insect cuticle. Material properties of the cuticle did not change significantly as a function of cuticular water content over the normal range throughout the feeding cycle (13-37% wet mass). Injected dopamine (DA) reduced one measure of the viscosity of the cuticle by 38%. Plastic deformability of the cuticle was reduced by 70% after an in vitro stretch, but restored in fully engorged ticks, and in in vitro stretched loops by treatment with DA and reduced pH. Thinning of the cuticle by half during the rapid phase of engorgement requires plastic deformation (irreversible strain) in two orthogonal dimensions in excess of 40%. Treatment with DA increased plastic deformation and enabled extensibility (strain at the point of rupture) above 40%.

The tegumental surface membranes of Schistosoma mansoni are enriched in parasite-specific phospholipid species.

The complex surface structure of adult Schistosoma mansoni, the tegument, is essential for survival of the parasite. This tegument is syncytial and is covered by two closely-apposed lipid bilayers that form the interactive surface with the host. In order to identify parasite-specific phospholipids present in the tegument, the species compositions of the major glycerophospholipid classes, phosphatidylcholine, phosphatidylserine, phosphatidylethanolamine and phosphatidylinositol, including lysophospholipid species, were analysed in adult S. mansoni worms, isolated tegumental membranes and hamster blood cells. It was shown that there are large differences in species composition in all four phospholipid classes between the membranes of S. mansoni and those of the host blood cells. The species compositions of phosphatidylserine and phosphatidylcholine were strikingly different in the tegument compared with the whole worm. The tegumental membranes are especially enriched in lysophospholipids, predominantly eicosenoic acid (20:1)-containing lyso-phosphatidylserine and lyso-phosphatidylethanolamine species. Furthermore, the tegument was strongly enriched in phosphatidylcholine that contained 5-octadecenoic acid, an unusual fatty acid that is not present in the host. As we have shown previously that lysophospholipids from schistosomes affect the parasite-host interaction, excretion of these tegument-specific phospholipid species was examined in vitro and in vivo. Our experiments demonstrated that these lysophospholipids are not significantly secreted during in vitro incubations and are not detectable in peripheral blood of infected hosts. However, these analyses demonstrated a substantial decrease in PI content of blood plasma from schistosome-infected hamsters, which might indicate that schistosomes influence exosome formation by the host.

Soft tissue influence on ex vivo mobility in the hip of Iguana: comparison with in vivo movement and its bearing on joint motion of fossil sprawling tetrapods.

The reconstruction of a joint's maximum range of mobility (ROM) often is a first step when trying to understand the locomotion of fossil tetrapods. But previous studies suggest that the ROM of a joint is restricted by soft tissues surrounding the joint. To expand the limited informative value of ROM studies for the reconstruction of a fossil species' locomotor characteristics, it is moreover necessary to better understand the relationship of ex vivo ROM with the actual in vivo joint movement. To gain insight into the relationship between ex vivo mobility and in vivo movement, we systematically tested for the influence of soft tissues on joint ROM in the hip of the modern lizard Iguana iguana. Then, we compared the ex vivo mobility to in vivo kinematics of the hip joint in the same specimens using X-ray sequences of steady-state treadmill locomotion previously recorded. With stepwise removal of soft tissues and a repeated-measurement protocol, we show that soft tissues surrounding the hip joint considerably limit ROM, highlighting the problems when joint ROM is deduced from bare bones only. We found the integument to have the largest effect on the range of long-axis rotation, pro- and retraction. Importantly, during locomotion the iguana used only a fragment of the ROM that was measured in our least restrictive dissection situation (i.e. pelvis and femur only conjoined by ligaments), demonstrating the discrepancy between hip joint ROM and actual in vivo movement. Our study emphasizes the necessity for caution when attempting to reconstruct joint ROM or even locomotor kinematics from fossil bones only, as actual in vivo movement cannot be deduced directly from any condition of cadaver mobility in Iguana and likely in other tetrapods.

Purification and characterization of coacervate-forming cuticular proteins from Papilio xuthus pupae.

The Papilio xuthus (Lepidoptera: Papilionidae) pupa expresses novel soluble proteins that undergo reversible temperature-dependent coacervate-formation. We purified two coacervate-forming proteins, PX-1 and PX-4, from the wings of pharate adults. PX-1 and PX-4 form coacervates upon warming. Transmission electron microscopy analysis revealed that these proteins assemble ordered bead-like ultrastructures. We cloned and sequenced PX-1 and PX-4 cDNAs. The PX-1 and PX-4 amino acid sequences contain many hydrophobic residues and show homologies to insect cuticular proteins. Moreover, when recombinant PX-1 and PX-4 were overexpressed in Escherichia coli, both recombinant proteins exhibited temperature-dependent coacervation. Furthermore, analyses of truncated mutants of PX-1 suggest that both the Val/Pro-rich region and Gly/lle-rich regions of PX-1 are involved in such coacervation.

Autophagy drives epidermal deterioration in a Drosophila model of tissue aging.

Organismal lifespan has been the primary readout in aging research. However, how longevity genes control tissue-specific aging remains an open question. To examine the crosstalk between longevity programs and specific tissues during aging, biomarkers of organ-specific aging are urgently needed. Since the earliest signs of aging occur in the skin, we sought to examine skin aging in a genetically tractable model. Here we introduce a Drosophila model of skin aging. The epidermis undergoes a dramatic morphological deterioration with age that includes membrane and nuclear loss. These changes were decelerated in a long-lived mutant and accelerated in a short-lived mutant. An increase in autophagy markers correlated with epidermal aging. Finally, the epidermis of Atg7 mutants retained younger characteristics, suggesting that autophagy is a critical driver of epidermal aging. This is surprising given that autophagy is generally viewed as protective during aging. Since Atg7 mutants are short-lived, the deceleration of epidermal aging in this mutant suggests that in the epidermis healthspan can be uncoupled from longevity. Because the aging readout we introduce here has an early onset and is easily visualized, genetic dissection using our model should identify other novel mechanisms by which lifespan genes feed into tissue-specific aging.

The exoskeleton of the female genitalic region in Petrobiellus takunagae (Insecta: Archaeognatha): insect-wide terminology, homologies, and functional interpretations.

The exoskeleton of the female genitalic region (abdominal venters 7-9) in Petrobiellus takunagae (Machilidae-Petrobiellinae) is studied using light microscopy and SEM. Sclerites are distinguished from membrane by the degree of cuticular flexibility. However, the microsculpture of the cuticle is shown to be useful in characterising the heterogeneity of the cuticle and in detecting weak sclerotisations. The morphology of Petrobiellus is compared with that in Trigoniophthalmus alternatus (Machilidae-Machilinae) described previously. While venter 7 is similar, venters 8 and 9 show many differences in the presence/absence or fusion/separation of particular sclerites. This suggests female genitalic morphology to be a valuable character system for phylogenetic and taxonomic work in Archaeognatha. Comparison with other insect orders is aimed at detecting homologous structures and conditions. Important points are: (1) Petrobiellus has a sclerotised genital lobe posteriorly on venter 7, similar to Zygentoma and Dictyoptera; it bears the gonopore. (2) Petrobiellus has a posterior sclerite on venter 9 that is very similar to a sclerite of Odonata. (3) The morphology of the coxal lobes of venter 9 (gonoplacs) suggests their function as a sheath of the ovipositor. From female genitalic morphology we deduce the process of oviposition, describing an external egg transportation tract.

No evidence for behavioral responses to circularly polarized light in four scarab beetle species with circularly polarizing exocuticle.

The strongest known circular polarization of biotic origin is the left-circularly polarized (LCP) light reflected from the metallic shiny exocuticle of certain beetles of the family Scarabaeidae. This phenomenon has been discovered by Michelson in 1911. Although since 1955 it has been known that the human eye perceives a visual illusion when stimulated by circularly polarized (CP) light, it was discovered only recently that a stomatopod shrimp is able to perceive circular polarization. It is pertinent to suppose that scarab beetles reflecting LCP light in an optical environment (vegetation) being deficient in CP signals may also perceive circular polarization and use it to find each other (mate/conspecifics) as until now it has been believed. We tested this hypothesis in six choice experiments with several hundred individuals of four scarab species: Anomala dubia, Anomala vitis (Coleoptera, Scarabaeidae, Rutelinae), and Cetonia aurata, Potosia cuprea (Coleoptera, Scarabaeidae, Cetoniinae), all possessing left-circularly polarizing exocuticle. From the results of our experiments we conclude that the studied four scarab species are not attracted to CP light when feeding or looking for mate or conspecifics. We demonstrated that the light reflected by host plants of the investigated scarabs is circularly unpolarized. Our results finally solve a puzzle raised over one hundred years ago, when Michaelson discovered that scarab beetles reflect circularly polarized light.

Müllerian mimicry: sharing the load reduces the legwork.

Color pattern mimicry has long been held up as a powerful example of natural selection. A recent study supports the theory by describing Müllerian mimicry rings in Appalachian millipedes that are analogous to those observed in tropical butterflies.

Alterações músculo-tegumentares e aminoácidos plasmáticos no hipertireoidismo (contribuição à interpretação fisiopatológica) / Changes muscle tissue and plasma amino acids in hyperthyroidism (contribution to the interpretation pathophysiological

Vários autores assinalam que o excesso de hormônio tireoidiano estimularia a proteólise muscular e, ao mesmo tempo, a captação esplênica de aminoácidos, determinando variações em suas concentrações plasmáticas...Alterações tegumentares na tireotoxicose, avaliadas pela presença de unhas de Plummer, correlacionaram-se signigicativamente com a redução de concentração plasmática de cistina (p,0,05)