Comparative anatomy of the rostrosoma of Solifugae, Pseudoscorpiones and Acari

We compare the microscopic anatomy of the mouthparts of representative species of Solifugae, Pseudoscorpiones and Parasitiformes (Acari). Specifically, we focus on the epistome, the labrum, the lateral lips (= endites of the pedipalpal coxae) and the musculature of the pharyngeal suction pump. We provide evidence that the labrum is reduced in Solifugae, but present and functional in Pseudoscorpiones and Acari. The epistome constitutes the entire dorsal face of the rostrosoma in Solifugae, but is internalized into the prosoma in Pseudoscorpiones. In Acari, the epistome shows an ancestral morphology, probably close to the ground pattern of chelicerates. The lateral lips of Solifugae contribute to the ventral face of the rostrosoma and the two lips of the mouth opening. In Solifugae, the ventral rostrosoma also includes a sclerite that might derive from a tritosternum. In Pseudoscorpiones, the lateral lips remain independent of the rostrosoma, they interlock ventral to the rostrosoma forming a perioral space. Here, the rostrosoma has an unpaired ventral lip of unresolved morphological origin, which is, however, clearly distinct from the lateral lips of Solifugae. The pharyngeal suction pump differs in all three clades in attachment, number of muscles and origin of muscles. We interpret the data as evidence for independent, parallel evolution of elements of the ground pattern of the (eu)chelicerate mouth parts. Based on the morphological elements of a common euchelicerate ground plan, the rostrosoma evolved independently in the three clades. We reject earlier hypotheses that consider the rostrosoma a character to support a phylogenetic relationship of the three clades

Malformations of the sacculus and the semicircular canals in spider morph pythons

Spider morph ball pythons are a frequently-bred designer morph with striking alterations of the skin color pattern. We created high-resolution μCT-image series through the otic region of the skulls, used 3D-reconstruction software for rendering anatomical models, and compared the anatomy of the semicircular ducts, sacculus and ampullae of wildtype Python regius (ball python) with spider morph snakes. All spider morph snakes showed the wobble condition (i.e., twisting movements of the head, impaired locomotion, difficulty striking or constricting prey items). We describe the inner ear structures in wildtype and spider morph snakes and report a deviant morphology of semicircular canals, ampullae and sacculus in the latter. We also report about associated differences in the desmal skull bones of spider morph snakes, which were characterized by wider semicircular canals, ampullae widened and difficult to discern in μCT, a deformed crus communis, and a small sacculus with a highly deviant X-ray morphology as compared to wildtype individuals. We observed considerable intra- and interindividual variability of these features. This deviant morphology in spider morph snakes could easily be associated with an impairment of sense of equilibrium and the observed neurological wobble condition. Limitations in sample size prevent statistical analyses, but the anatomical evidence is strong enough to support an association between the wobble condition and a malformation of the inner ear structures. A link between artificially selected alterations in pattern and specific color design with neural-crest associated developmental malformations of the statoacoustic organ as known from other vertebrates is discussed

Morphological responses to feeding in ticks (Ixodes ricinus)

Background: Ticks can survive long periods without feeding but, when feeding, ingest large quantities of blood, resulting in a more than 100-fold increase of body volume. We study morphological adaptations to changes in opisthosoma volume during feeding in the castor bean tick, Ixodes ricinus. We aim to understand the functional morphological features that accommodate enormous changes in volume changes. Methods: Using light and electron microscopy, we compare the cuticle and epidermis of the alloscutum, the epithelium of the midgut diverticula, and the tracheae of adult female ticks when fasting, semi-engorged, and fully engorged. Results: Our results add to an existing body of knowledge that the area of the epidermis increases by cellular differentiation, cellular hypertrophy, and changes in the shape of epithelial cells from pseudostratified to single layered prismatic in semi-engorged ticks, and to thin squamous epithelium in fully engorged ticks. We did not find evidence for cell proliferation. The midgut diverticula accommodate the volume increase by cellular hypertrophy and changes in cell shape. In fully engorged ticks, the epithelial cells of the midgut diverticula are stretched to an extremely thin, squamous epithelium. Changes in size and shape (and cell divisions) contribute to the accommodation of volume changes. Tracheae do not increase in size, but extend in length, thus following the volume changes of the opisthosoma in feeding ticks to secure oxygen supply to the internal organs. Conclusions: Changes of epithelial tissue configuration in the epidermis and the midgut diverticula are described as important components of the morphological response to feeding in ticks. We provide evidence for a previously unknown mechanism hosted in the endocuticle of the tracheae that allows the tracheae of castor bean ticks to expand when the body volume increases and the distance between the respiratory spiracle and the oxygen demanding tissue enlarges. This is the first report of expandable tracheae in arthropods

m6A potentiates Sxl alternative pre-mRNA splicing for robust Drosophila sex determination

N6-methyladenosine (m6A) is the most common internal modification of eukaryotic messenger RNA (mRNA) and is decoded by YTH domain proteins1, 2, 3, 4, 5, 6, 7. The mammalian mRNA m6A methylosome is a complex of nuclear proteins that includes METTL3 (methyltransferase-like 3), METTL14, WTAP (Wilms tumour 1-associated protein) and KIAA1429. Drosophila has corresponding homologues named Ime4 and KAR4 (Inducer of meiosis 4 and Karyogamy protein 4), and Female-lethal (2)d (Fl(2)d) and Virilizer (Vir)8, 9, 10, 11, 12. In Drosophila, fl(2)d and vir are required for sex-dependent regulation of alternative splicing of the sex determination factor Sex lethal (Sxl)13. However, the functions of m6A in introns in the regulation of alternative splicing remain uncertain3. Here we show that m6A is absent in the mRNA of Drosophila lacking Ime4. In contrast to mouse and plant knockout models5, 7, 14, Drosophila Ime4-null mutants remain viable, though flightless, and show a sex bias towards maleness. This is because m6A is required for female-specific alternative splicing of Sxl, which determines female physiognomy, but also translationally represses male-specific lethal 2 (msl-2) to prevent dosage compensation in females. We further show that the m6A reader protein YT521-B decodes m6A in the sex-specifically spliced intron of Sxl, as its absence phenocopies Ime4 mutants. Loss of m6A also affects alternative splicing of additional genes, predominantly in the 5′ untranslated region, and has global effects on the expression of metabolic genes. The requirement of m6A and its reader YT521-B for female-specific Sxl alternative splicing reveals that this hitherto enigmatic mRNA modification constitutes an ancient and specific mechanism to adjust levels of gene expression

Phylogeny and evolutionary history of the amniote egg

We review morphological features of the amniote egg and embryos in a comparative phylogenetic framework, including all major clades of extant vertebrates. We discuss 40 characters that are relevant for an analysis of the evolutionary history of the vertebrate egg. Special attention is given to the morphology of the cellular yolk sac, the eggshell, and extraembryonic membranes. Many features that are typically assigned to amniotes, such as a large yolk sac, delayed egg deposition, and terrestrial reproduction have evolved independently and convergently in numerous clades of vertebrates. We use phylogenetic character mapping and ancestral character state reconstruction as tools to recognize sequence, order, and patterns of morphological evolution and deduce a hypothesis of the evolutionary history of the amniote egg. Besides amnion and chorioallantois, amniotes ancestrally possess copulatory organs (secondarily reduced in most birds), internal fertilization, and delayed deposition of eggs that contain an embryo in the primitive streak or early somite stage. Except for the amnion, chorioallantois, and amniote type of eggshell, these features evolved convergently in almost all major clades of aquatic vertebrates possibly in response to selective factors such as egg predation, hostile environmental conditions for egg development, or to adjust hatching of young to favorable season. A functionally important feature of the amnion membrane is its myogenic contractility that moves the (early) embryo and prevents adhering of the growing embryo to extraembryonic materials. This function of the amnion membrane and the liquid-filled amnion cavity may have evolved under the requirements of delayed deposition of eggs that contain developing embryos. The chorioallantois is a temporary embryonic exchange organ that supports embryonic development. A possible evolutionary scenario is that the amniote egg presents an exaptation that paved the evolutionary pathway for reproduction on land. As shown by numerous examples from anamniotes, reproduction on land has occurred multiple times among vertebrates—the amniote egg presenting one “solution” that enabled the conquest of land for reproduction

Physiological and morphological responses to feeding in broad-nosed caiman (Caiman latirostris)

Broad nosed caiman are ectotherm sauropsids that naturally experience long fasting intervals. We have studied the postprandial responses by measuring oxygen consumption using respirometry, the size changes of the duodenum, the distal small intestine, and the liver, using repeated non-invasive ultrasonography, and by investigating structural changes on the level of tissues and cells by using light- and electron microscopy. The caimans showed the same rapid and reversible changes of organ size and identical histological features, down to the ultrastructure level, as previously described for other ectothermic sauropsids. We found a configuration change of the mucosa epithelium from pseudostratified during fasting to single layered during digestion, in association with hypertrophy of enterocytes by loading them with lipid droplets. Similar patterns were also found for the hepatocytes of the liver. By placing the results of our study in comparative relationship and by utilizing the phylogenetic bracket of crocodiles, birds and squamates, we suggest that the observed features are plesiomorphic characters of sauropsids. By extending the comparison to anurans, we suggest that morphological and physiological adjustments to feeding and fasting described here may have been a character of early tetrapods. In conclusion, we suggest that the ability to tolerate long fasting intervals and then swallow a single large meal as described for many sit-an-wait foraging sauropsids is a functional feature that was already present in ancestral tetrapods