RESUMEN
SUMMARY: This study evaluated the morphology of alpacas skin. Biopsies were collected and samples were fixed in 10 % neutral buffered formalin for histological procedures. The sections were stained with hematoxylin and eosin, picrosirius red and Masson's trichrome. Types I, III and IV collagen were analyzed by immunohistochemistry. The derma presented sebaceous and sweat glands, as well as follicular groups with medullated fibers. Type I and type IV collagen were observed at epidermis and dermis as well as in glandular structures and hair follicles. The collagen III, was observed only in dermis.
Este estudio evaluó la morfología de la piel de alpacas. Se recogieron biopsias y las muestras se fijaron en formalina tamponada neutra al 10 % para procedimientos histológicos. Las secciones se tiñeron con hematoxilina y eosina, rojo picrosirius y tricrómico de Masson. El colágeno tipo I, III y IV se analizó mediante inmunohistoquímica. La dermis presentó glándulas sebáceas y sudoríparas, así como grupos foliculares con fibras medulares. Se observó colágeno tipo I y tipo IV en la epidermis y la dermis, así como en estructuras glandulares y folículos pilosos. El colágeno III, se observó únicamente en la dermis.
Asunto(s)
Animales , Camélidos del Nuevo Mundo/anatomía & histología , Integumento Común/anatomía & histología , Inmunohistoquímica , Microscopía Electrónica de RastreoRESUMEN
The skin is a barrier between the internal and external environment of an organism. Depending on the species, it participates in multiple functions. The skin is the organ that holds the body together, covers and protects it, and provides communication with its environment. It is also the body's primary line of defense, especially for anamniotes. All vertebrates have multilayered skin composed of three main layers: the epidermis, the dermis, and the hypodermis. The vital mission of the integument in aquatic vertebrates is mucus secretion. Cornification began in apmhibians, improved in reptilians, and endured in avian and mammalian epidermis. The feather, the most ostentatious and functional structure of avian skin, evolved in the Mesozoic period. After the extinction of the dinosaurs, birds continued to diversify, followed by the enlargement, expansion, and diversification of mammals, which brings us to the most complicated skin organization of mammals with differing glands, cells, physiological pathways, and the evolution of hair. Throughout these radical changes, some features were preserved among classes such as basic dermal structure, pigment cell types, basic coloration genetics, and similar sensory features, which enable us to track the evolutionary path. The structural and physiological properties of the skin in all classes of vertebrates are presented. The purpose of this review is to go all the way back to the agnathans and follow the path step by step up to mammals to provide a comparative large and updated survey about vertebrate skin in terms of morphology, physiology, genetics, ecology, and immunology.
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Evolución Biológica , Dinosaurios , Animales , Dinosaurios/fisiología , Plumas/anatomía & histología , Integumento Común/anatomía & histología , Integumento Común/fisiología , Aves/anatomía & histología , Mamíferos/anatomía & histologíaRESUMEN
We here report on a well-preserved juvenile lizard specimen in Albian amber (ca. 110 mya) from the Hkamti site (Myanmar). This new taxon is represented by an articulated skull and the anterior portion of the trunk, including the pectoral girdle and forelimbs. The scleral ossicles and eyelid are also visible, and the specimen exhibits pristine detail of the integument (of both head and body). In a combined molecular and morphological analysis, it was consistently recovered as a scincoid lizard (Scinciformata), as sister to Tepexisaurus + Xantusiidae. However, the phylogenetic position of the new taxon should be interpreted with caution as the holotype is an immature individual. We explored the possibility of miscoding ontogenetically variable characters by running alternative analyses in which these characters were scored as missing data for our taxon. With the exception of one tree, in which it was sister to Amphisbaenia, the specimen was recovered as a Pan-xantusiid. Moreover, we cannot rule out the possibility that it represents a separate lineage of uncertain phylogenetic position, as it is the case for many Jurassic and Cretaceous taxa. Nonetheless, this fossil offers a rare opportunity to glimpse the external appearance of one group of lizards during the Early Cretaceous.
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Ámbar , Fósiles/anatomía & histología , Integumento Común/anatomía & histología , Lagartos/anatomía & histología , Animales , Evolución Biológica , Fósiles/diagnóstico por imagen , Integumento Común/diagnóstico por imagen , Lagartos/genética , Mianmar , Filogenia , Microtomografía por Rayos XRESUMEN
Evolution has created different castes of females in eusocial haplodiploids. The difference between them lies in their functions and vulnerability but above all in their reproductive potentials. Honeybee queens are highly fertile. On the other hand, the workers are facultatively sterile. However, rebel workers, i.e. workers that develop in a queenless colony, reproduce more often than normal workers. As a result, the fat body of these bees, which apart from acting as the energy reserve, is also the site of numerous metabolic processes, had to specialize in different functions perfected over millions of years of eusocial evolution. Assuming that the variety of functions manifests itself in the pleomorphic structure of the fat body cells, we predicted that also different parts of the fat body, e.g. from different segments of the abdomen, contain different sets of cells. Such differences could be expected between queens, rebels and normal workers, i.e. females with dramatically different reproductive potentials. We confirmed all these expectations. Although all bees had the same types of cells, their proportion and segmental character corresponded with the caste reproductive potential and physiological characteristics shaped in the evolutionary process. The females with an increased reproductive potential were characterized by the presence of oenocytes in the third tergite and high concentrations of compounds responsible for energy reserves, like glucose, glycogen and triglycerides. Queens had very large trophocytes, especially in the third tergite. Only in workers did we observe intercellular spaces in all the segments of the fat body, as well as high protein concentrations-especially in the sternite. As expected, the rebels combined many features of the queens and normal workers, what with other findings can help understand the ways that led to the origin of different castes in females of eusocial Hymenoptera.
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Abejas/fisiología , Cuerpo Adiposo/anatomía & histología , Integumento Común/anatomía & histología , Animales , Peso Corporal , Núcleo Celular/metabolismo , Cuerpo Adiposo/citología , Femenino , Glucosa/metabolismo , Glucógeno/metabolismo , Proteínas de Insectos/metabolismo , Reproducción , Triglicéridos/metabolismoRESUMEN
The second integument of the angiosperm ovule is unique among seed plants, with developmental genetics that are distinct from those of the inner integument1. Understanding how the second integument should be compared to structures in other seed plants is therefore crucial to resolving the long-standing question of the origin of angiosperms2-6. Attention has focused on several extinct plants with recurved cupules that are reminiscent of the anatropous organization of the basic bitegmic ovules of angiosperms1-6, but interpretations have been hampered by inadequate information on the relevant fossils. Here we describe abundant exceptionally well-preserved recurved cupules from a newly discovered silicified peat dating to the Early Cretaceous epoch (around 125.6 million years ago) in Inner Mongolia, China. The new material, combined with re-examination of potentially related fossils, indicates that the recurved cupules of several groups of Mesozoic plants are all fundamentally comparable, and that their structure is consistent with the recurved form and development of the second integument in the bitegmic anatropous ovules of angiosperms. Recognition of these angiosperm relatives (angiophytes) provides a partial answer to the question of angiosperm origins, will help to focus future work on seed plant phylogenetics and has important implications for ideas on the origin of the angiosperm carpel.
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Evolución Biológica , Extinción Biológica , Fósiles , Integumento Común/anatomía & histología , Magnoliopsida/anatomía & histología , Óvulo Vegetal/anatomía & histología , China , Historia Antigua , Magnoliopsida/ultraestructura , Mongolia , Óvulo Vegetal/ultraestructura , FilogeniaRESUMEN
Despite the relevance of chemical communication in vertebrates, comparative examinations of macroevolutionary trends in chemical signaling systems are scarce. Many turtle and tortoise species are reliant on chemical signals to communicate in aquatic and terrestrial macrohabitats, and many of these species possess specialized integumentary organs, termed mental glands (MGs), involved in the production of chemosignals. We inferred the evolutionary history of MGs and tested the impact of macrohabitat on their evolution. Inference of ancestral states along a time-calibrated phylogeny revealed a single origin in the ancestor of the subclade Testudinoidea. Thus, MGs represent homologous structures in all descending lineages. We also inferred multiple independent losses of MGs in both terrestrial and aquatic clades. Although MGs first appeared in an aquatic turtle (the testudinoid ancestor), macrohabitat seems to have had little effect on MG presence or absence in descendants. Instead, we find clade-specific evolutionary trends, with some clades showing increased gland size and morphological complexity, whereas others exhibiting reduction or MG loss. In sister clades inhabiting similar ecological niches, contrasting patterns (loss vs. maintenance) may occur. We conclude that the multiple losses of MGs in turtle clades have not been influenced by macrohabitat and that other factors have affected MG evolution.
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Evolución Biológica , Integumento Común/fisiología , Feromonas/química , Tortugas/fisiología , Comunicación Animal , Animales , Fenómenos Biomecánicos , Ecosistema , Integumento Común/anatomía & histología , Feromonas/biosíntesis , Filogenia , Transducción de Señal/genética , Tortugas/anatomía & histologíaRESUMEN
Holothuria atra is a black sea cucumber commonly found on the sandy bottom of Okinawan coral reefs. The body surface of H. atra is usually covered with sand; however, sand never covers the body of another black congener, Holothuria leucospilota, which is sympatrically distributed with H. atra. The epidermal structures were examined in these two species by means of transmission electron microscopy to determine how sand adheres to the surface of H. atra. While the epidermis was basically composed of support cells bearing microvilli and vacuolated cells probably corresponding to mucus cells, two types of granular cells, type 1 and 2, were also found at the tip of the tube feet. These granular cells were closely similar in structure to secretory cells that have been supposed to secrete adhesive substances in other holothurians. Type 1 granular cells were also found in the dorsal epidermis of H. atra but not in H. leucospilota. Therefore, adhesive secretion by type 1 granular cells probably enables the attachment of sand to the H. atra body.
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Holothuria/anatomía & histología , Arena , Animales , Ecosistema , Holothuria/fisiología , Integumento Común/anatomía & histología , Integumento Común/fisiologíaRESUMEN
Birds are the most diversified terrestrial vertebrates due to highly diverse integumentary organs that enable robust adaptability to various eco-spaces. Here we show that this complexity is built upon multi-level regional specifications. Across-the-body (macro-) specification includes the evolution of beaks and feathers as new integumentary organs that are formed with regional specificity. Within-an-organ (micro-) specification involves further modifications of organ shapes. We review recent progress in elucidating the molecular mechanisms underlying feather diversification as an example. (1) ß-Keratin gene clusters are regulated by typical enhancers or high order chromatin looping to achieve macro- and micro-level regional specification, respectively. (2) Multi-level symmetry-breaking of feather branches confers new functional forms. (3) Complex color patterns are produced by combinations of macro-patterning and micro-patterning processes. The integration of these findings provides new insights toward the principle of making a robustly adaptive bio-interface.
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Adaptación Fisiológica/genética , Evolución Biológica , Aves/fisiología , Integumento Común/fisiología , Animales , Pico/anatomía & histología , Pico/fisiología , Aves/genética , Plumas/anatomía & histología , Plumas/fisiología , Integumento Común/anatomía & histología , Piel/anatomía & histología , Vertebrados/genética , Vertebrados/fisiologíaRESUMEN
Sand deserts are common biotopes on the earth's surface. Numerous morphological and physiological adaptations have appeared to cope with the peculiar conditions imposed by sandy substrates, such as abrasion, mechanical resistance and the potential low oxygen levels. The psammophilous scincids (Lepidosauria) Scincus scincus and Eumeces schneideri are among those. S. scincus is a species frequently used to study displacement inside a sandy substrate. E. schneideri is a species phylogenetically closely related to S. scincus with a similar lifestyle. The aims of this study focus on the morphology of the integument and the muscular system. Briefly, we describe interspecific differences at the superficial architecture of the scales pattern and the thickness of the integument. We highlight a high cellular turnover rate at the level of the basal germinal layer of the epidermis, which, we suggest, corresponds to an adaptation to cutaneous wear caused by abrasion. We demonstrate the presence of numerous cutaneous holocrine glands whose secretion probably plays a role in the flow of sand along the integument. Several strata of osteoderms strengthen the skin. We characterize the corporal (M. longissimus dorsi and M. rectus abdominus) and caudal muscular fibers using immunohistochemistry, and quantify them using morphometry. The musculature exhibits a high proportion of glycolytic fast fibers that allow rapid burying and are well adapted to this mechanically resistant and oxygen-poor substrate. Oxidative slow fibers are low in abundance, less than 10% in S. scincus, but a little higher in E. schneideri.
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Integumento Común/anatomía & histología , Lagartos/anatomía & histología , Músculo Esquelético/anatomía & histología , Adaptación Fisiológica , Animales , División Celular , Dermis/anatomía & histología , Células Epidérmicas/citología , Epidermis/anatomía & histología , Femenino , Masculino , FilogeniaRESUMEN
Early in amniote evolution, epidermal scales evolved in stem reptiles as an efficient barrier against water loss and ultraviolet radiation, making them a key development in the transition to a fully terrestrial existence [1]. Accordingly, epidermal scales are not simple inert structures but highly-evolved organs suited to perform a broad suite of functions. Here, we provide new data on the epidermal complexity of a non-avian theropod, Juravenator starki, from the Torleite Formation (upper Kimmeridgian), Bavaria, Germany [2]. Although epidermal scales have been noted previously on the tail of Juravenator, we report a unique scale type with distinctive circular nodes that we identify as integumentary sense organs, analogous to those in modern crocodylians. The surprising presence of such structures suggests the tail had a sensory function, which is nevertheless congruent with the inferred ecology of Juravenator and the evolution of integumentary sense organs among archosaurs.
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Evolución Biológica , Dinosaurios/anatomía & histología , Dinosaurios/fisiología , Epidermis/anatomía & histología , Epidermis/fisiopatología , Fósiles , Órganos de los Sentidos/fisiología , Caimanes y Cocodrilos/fisiología , Animales , Integumento Común/anatomía & histología , Integumento Común/fisiología , Rayos UltravioletaRESUMEN
Euplatypus parallelus (F.) (Coleoptera: Curculionidea) is the most destructive cosmopolitan insect pest of the Platypodinae. Pheromone-based luring agents are used currently in controlling bark beetle. Antennae are the primary insect organs sensing volatiles of host trees and pheromones of pioneer males. We studied the external morphology of antennae and the type, distribution, and the number of the beetle sensilla. Our results show E. parallelus have a geniculate antenna composed of 6 segments, namely the scape, 4-segmented funicle and club. Ninety-seven percent of the antennal sensors were distributed in the club, and 3% were distributed in the scape and funicle. 6 types of sensilla on the antennae were found, including sensilla trichodea (subtypes: STI, STII and STIII), sensilla basiconica (subtypes: SBI, SBII, SBIII and SBIV), sensilla chaetica (subtypes: SChI, SChII and SChIII), as well as sensilla coeloconica, sensilla campaniform and sensilla furcatea. There was no significant difference in the type, distribution and number of sensilla in males and females. No significant difference in the shape and distribution of antennae was found between sexes, but the length of antennae and the number of SChI, SChII, STI, SBI, SBIII and SBIV were significantly larger in females than males. We revealed the external cuticular structure of the antennae in E. parallelus, which can be used to guide future electrophysiological investigations to understand the ability of this beetle to detect semiochemicals.
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Antenas de Artrópodos/anatomía & histología , Escarabajos/anatomía & histología , Sensilos/anatomía & histología , Animales , Antenas de Artrópodos/ultraestructura , Escarabajos/ultraestructura , Femenino , Integumento Común/anatomía & histología , Masculino , Sensilos/ultraestructuraRESUMEN
Scaly-tailed squirrels, the most poorly known group of gliding mammals, hold the record for variety of remarkable integument peculiarities. One of the most striking of these features is the scales on the tail, which apparently allow them to reduce energy costs when positioning themselves on a tree trunk. No less interesting is a peculiar spur that supports the flying membrane: the unciform element ('spur'). Despite the peculiarity of such elements, their nature has not yet been studied. Using anatomical, histological methods and scanning electron microscopy we studied the structure of the skin and its derivatives in five of the six species from both genera of extant gliding scaly-tailed squirrels (Anomaluridae, Rodentia): Idiurus macrotis, Idiurus zenkeri, Anomalurus beecrofti, Anomalurus pusillus and Anomalurus derbianus. In addition to the common mammalian skin structures, such as hair, vibrissae, sebaceous glands, meibomian glands of eyelids and eccrine sweat glands of the palmar and plantar pads, these animals have unique species-specific skin derivatives (the tail scaly organ and its specific glands, vibrissae of the withers, patagium and its hair brush) that play a significant role in their adaptation to gliding and to their environment in general. The structure of the elbow spur is also described and hypotheses on its evolutionary origin from the tendon of the triceps muscle are presented.
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Adaptación Fisiológica/fisiología , Integumento Común/anatomía & histología , Locomoción/fisiología , Roedores/anatomía & histología , Animales , Integumento Común/fisiología , Roedores/fisiología , Especificidad de la EspecieRESUMEN
Cylindrical silk gland (CY) spigots distinguish a large clade of modern spiders, the CY spigot clade, which includes all entelegyne spiders and their closest relatives. Following a widespread paradigm, CYs and their spigots are only known to occur in female spiders and they produce silk used in the construction of egg sacs. Here we report the occurrence of a CY spigot or CY nubbin on each posterior median spinneret (PMS) in males (5th stadium and later) of the spider Australomimetus maculosus. Late juvenile males had a CY spigot on each PMS, whereas adult males either had a CY spigot or, more often, a non-functional CY nubbin. This indicates that potential CY use by males is at least largely limited to late juvenile instars and is not involved with egg sac construction. Despite the presence of CY spigots in both sexes, sexual dimorphism with respect to CYs was still evident since males lacked the CY spigot on each posterior lateral spinneret present in late juvenile and adult females, and CY spigots of males never had the wide shaft and opening of adult females. This study adds to our knowledge of spinning apparatus variability in modern spiders and demonstrates an exception to the paradigm that, in the CY spigot clade, such spigots are restricted to female spiders.
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Integumento Común/fisiología , Caracteres Sexuales , Seda/biosíntesis , Arañas/fisiología , Animales , Femenino , Integumento Común/anatomía & histología , Masculino , Microscopía Electrónica de Rastreo , Arañas/anatomía & histología , Arañas/ultraestructuraRESUMEN
During development, cell fate decisions are often highly stochastic, but with the frequency of the different possible fates tightly controlled. To understand how signaling networks control the cell fate frequency of such random decisions, we studied the stochastic decision of the Caenorhabditis elegans P3.p cell to either fuse to the hypodermis or assume vulva precursor cell fate. Using time-lapse microscopy to measure the single-cell dynamics of two key inhibitors of cell fusion, the Hox gene LIN-39 and Wnt signaling through the ß-catenin BAR-1, we uncovered significant variability in the dynamics of LIN-39 and BAR-1 levels. Most strikingly, we observed that BAR-1 accumulated in a single, 1-4 âh pulse at the time of the P3.p cell fate decision, with strong variability both in pulse slope and time of pulse onset. We found that the time of BAR-1 pulse onset was delayed relative to the time of cell fusion in mutants with low cell fusion frequency, linking BAR-1 pulse timing to cell fate outcome. Overall, a model emerged where animal-to-animal variability in LIN-39 levels and BAR-1 pulse dynamics biases cell fate by modulating their absolute level at the time cell fusion is induced. Our results highlight that timing of cell signaling dynamics, rather than its average level or amplitude, could play an instructive role in determining cell fate.
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Proteínas de Caenorhabditis elegans/metabolismo , Caenorhabditis elegans/citología , beta Catenina/metabolismo , Animales , Sistemas CRISPR-Cas , Caenorhabditis elegans/genética , Proteínas de Caenorhabditis elegans/biosíntesis , Proteínas de Caenorhabditis elegans/genética , Diferenciación Celular , Fusión Celular , Linaje de la Célula , Proteínas del Citoesqueleto/metabolismo , Femenino , Regulación del Desarrollo de la Expresión Génica , Genotipo , Proteínas de Homeodominio/metabolismo , Hibridación Fluorescente in Situ , Integumento Común/anatomía & histología , Glicoproteínas de Membrana/biosíntesis , Glicoproteínas de Membrana/genética , Proteínas Recombinantes de Fusión/metabolismo , Análisis de la Célula Individual , Procesos Estocásticos , Imagen de Lapso de Tiempo , Vulva/citología , Vía de Señalización WntRESUMEN
A striking characteristic of insect cuticle is the wide range of its material property values, with respect to stiffness, strength and toughness. The elastic modulus of cuticle, for instance, ranges over seven orders of magnitude in different structures and different species. Previous studies suggested that this characteristic is influenced by the microstructure and sclerotization of cuticle. However, the relative role of the two factors in determining the material properties of cuticle is unknown. Here we used a combination of scanning electron microscopy (SEM), confocal laser scanning microscopy (CLSM) and nanoindentation, to investigate the effect of microstructure and sclerotization on the elastic modulus of tibiae of desert locusts. Our results showed that tibial cuticle is an anisotropic material with the highest elastic modulus along the tibial axis. This is likely because majority of the fibers in the cuticle are oriented along this axis. We also found that the hind tibia has a significantly higher elastic modulus, compared with the fore and mid tibiae. This is likely due to the higher sclerotization level of the hind tibia cuticle, and seems to be an adaptation to the locust locomotion by jumping, in which axial loads in the hind tibiae may reach several times the insect body weight. Our results suggest that while sclerotization determines the difference between the elastic moduli of the tibiae, anisotropic properties of each tibia is controlled by the specific fiber orientation. Our study provides one of only a few comprehensive investigations on insect cuticle, and helps to better understand the structure-material-function relationship in this complex biological composite. STATEMENT OF SIGNIFICANCE: Insect cuticle is a biological composite with strong anisotropy and wide ranges of material properties. Using an example of the tibial cuticle of desert locusts, we examined the role of two influential factors on the elastic modulus of cuticle: microstructure and sclerotization. Our results suggested the strong influence of sclerotization on the variation of the elastic modulus among fore, mid and hind tibiae, and that of the microstructure on the anisotropy of each tibia. Our results deepens the current understanding of the structure-material-function relationship in complex insect cuticle.
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Módulo de Elasticidad , Saltamontes/anatomía & histología , Saltamontes/fisiología , Integumento Común/anatomía & histología , Tibia/anatomía & histología , Tibia/fisiología , Animales , Saltamontes/ultraestructura , Imagenología Tridimensional , Tibia/ultraestructuraRESUMEN
In many cases, strong friction reduction is critical for success of both living organisms and engineering systems. Some arthropods exhibit good antifriction abilities in their specific living environments and have inspired many innovations for solving industry challenges. However, the current literature mainly focused on terrestrial insects, such as beetles, grasshoppers and katydids. The antifriction mechanisms in amphibious arthropods are still unknown, even if their surfaces are optimized for both air and water environments. Herein the tribological properties of the cuticle surface of the sandhopper Talitrus saltator were studied using a universal microtribometer. Further investigations were developed to identify the microstructural, compositional, wettability, and mechanical properties of the sandhopper shell cuticles. It was found that increasing normal force can significantly reduce the coefficient of friction of the shell cuticle, especially for the alive and rewet sandhopper shells. The shell consists of bottle-like nano-caves in its exocuticle, nano-tubes in its mesocuticle, and gauze-like multilayers in its endocuticle. Under physiological conditions, glycoprotein-like fluid fillings exist in both the bottle-like caves and the nano-tubes below and cover on the shell surface. More importantly, a new antifriction mechanism of lubricant-squeezing nano-porous system was established for the sandhopper shell. This work can deepen our understanding in antifriction surfaces of amphibiotic crustaceans, and provide a potential approach to resolve the friction challenge in micro-machines, especially for the applications under aqueous condition. STATEMENT OF SIGNIFICANCE: Friction regulation is one of the critical mechanisms for animal locomotion in natural environments. However, not much is known about the mechanism of amphibious arthropods to reduce friction between their body and diverse environments, particularly achieving adaption under both air and aqueous conditions. We quantitatively study the microstructural, compositional and mechanical properties of the sandhopper (Talitrus saltator) shell cuticle and tribological behaviors under different conditions. Our results reveal the nano-porous system with fluid fillings for the sandhopper's shell and demonstrate the potential antifriction mechanism of this amphibious animal. We anticipate this work will inspire some effective antifriction designs for micro-machines, especially for their applications in complex environment like human body.
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Anfípodos/anatomía & histología , Anfípodos/fisiología , Fricción , Integumento Común/anatomía & histología , Integumento Común/fisiología , Anfípodos/ultraestructura , Exoesqueleto/anatomía & histología , Exoesqueleto/ultraestructura , Animales , Módulo de Elasticidad , Modelos Lineales , Reología , Propiedades de Superficie , HumectabilidadRESUMEN
The cuticle of stomatopod is an example of a natural mineralized biomaterial, consisting of chitin, amorphous calcium carbonate and protein components with a multiscale hierarchical structure, and forms a protective shell with high impact resistance. At the ultrastructural level, cuticle mechanical functionality is enabled by the nanoscale architecture, wherein chitin fibrils are in intimate association with enveloping mineral and proteins. However, the interactions between these ultrastructural building blocks, and their coupled response to applied load, remain unclear. Here, we elucidate these interactions via synchrotron microbeam wide-angle X-ray diffraction combined with in situ tensile loading, to quantify the chitin crystallite structure of native cuticle - and after demineralization and deproteinization - as well as time-resolved changes in chitin fibril strain on macroscopic loading. We demonstrate chitin crystallite stabilization by mineral, seen via a compressive pre-strain of approximately 0.10% (chitin/protein fibre pre-stress of â¼20â¯MPa), which is lost on demineralization. Clear reductions of stiffness at the fibrillar-level following matrix digestion are linked to the change in the protein/matrix mechanical properties. Furthermore, both demineralization and deproteinization alter the 3D-pattern of deformation of the fibrillar network, with a non-symmetrical angular fibril strain induced by the chemical modifications, associated with loss of the load-transferring interfibrillar matrix. Our results demonstrate and quantify the critical role of interactions at the nanoscale (between chitin-protein and chitin-mineral) in enabling the molecular conformation and outstanding mechanical properties of cuticle, which will inform future design of hierarchical bioinspired composites. STATEMENT OF SIGNIFICANCE: Chitinous biomaterials (e.g. arthropod cuticle) are widespread in nature and attracting attention for bioinspired design due to high impact resistance coupled with light weight. However, how the nanoscale interactions of the molecular building blocks - alpha-chitin, protein and calcium carbonate mineral - lead to these material properties is not clear. Here we used X-ray scattering to determine the cooperative interactions between chitin fibrils, protein matrix and biominerals, during tissue loading. We find that the chitin crystallite structure is stabilized by mineral nanoparticles, the protein phase prestresses chitin fibrils, and that chemical modification of the interfibrillar matrix significantly disrupts 2D mechanics of the microfibrillar chitin plywood network. These results will aid rational design of advanced chitin-based biomaterials with high impact resistance.
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Calcificación Fisiológica , Quitina/química , Matriz Extracelular/metabolismo , Animales , Materiales Biocompatibles , Fenómenos Biomecánicos , Crustáceos/anatomía & histología , Integumento Común/anatomía & histología , Estrés Mecánico , Resistencia a la TracciónRESUMEN
Intraspecific variation during the anuran larval period has been analyzed mainly in relation to the timing of metamorphosis and body size at metamorphosis. However, other traits may vary as well. We examined two developmental series of Boana riojana from the same population in two consecutive years and describe intraspecific variation in larvae of this species. We discuss how variation, if present, may influence its life cycle. We found that both larval series differed in the larval period length, one twice as long as the other. This variation primarily depended on when breeding occurred, metamorphosis was achieved during late spring in both generations and at similar sizes, and only the rate of larval development during premetamorphosis varied extensively between years. This is consistent with thyroid gland activity because when it became active the developmental trajectory became more canalized. No variation of staging sequence occurred in relation to the different durations of the larval period. However, in the long-lasting series we found two different morphs. Also, integument, thyroid gland, skeleton, and testis differentiation events occurred at the same developing stages. In contrast, ovarian differentiation proceeded at the same absolute age in both series. Sexual dimorphism becomes evident within the year after metamorphosis. The intraspecific heterochrony that we describe for the larval development of B. riojana does not lead to phenotypic variation at the end of metamorphosis. We discuss the importance of analyzing growth and development independently. Each proceeds differently in time, but with an interdependence at some point, because size and shape do not vary at the end of metamorphosis.
Asunto(s)
Anuros/crecimiento & desarrollo , Estadios del Ciclo de Vida , Envejecimiento/fisiología , Animales , Anuros/anatomía & histología , Tamaño Corporal , Femenino , Integumento Común/anatomía & histología , Larva/anatomía & histología , Larva/crecimiento & desarrollo , Sistema de la Línea Lateral/anatomía & histología , Masculino , Fenotipo , Especificidad de la Especie , Factores de TiempoRESUMEN
Arthropod cuticle has extraordinary properties. It is very stiff and tough whilst being lightweight, yet it is made of rather ordinary constituents. This desirable combination of properties results from a hierarchical structure, but we currently have a poor understanding of how this impedes damage propagation. Here we use non-destructive, time-lapse in situ tensile testing within an X-ray nanotomography (nCT) system to visualise crack progression through dry beetle elytron (wing case) cuticle in 3D. We find that its hierarchical pseudo-orthogonal laminated microstructure exploits many extrinsic toughening mechanisms, including crack deflection, fibre and laminate pull-out and crack bridging. We highlight lessons to be learned in the design of engineering structures from the toughening methods employed. STATEMENT OF SIGNIFICANCE: We present the first comprehensive study of the damage and toughening mechanisms within arthropod cuticle in a 3D time-lapse manner, using X-ray nanotomography during crack growth. This technique allows lamina to be isolated despite being convex, which limits 2D analysis of microstructure. We report toughening mechanisms previously unobserved in unmineralised cuticle such as crack deflection, fibre and laminate pull-out and crack bridging; and provide insights into the effects of hierarchical microstructure on crack propagation. Ultimately the benefits of the hierarchical microstructure found here can not only be used to improve biomimetic design, but also helps us to understand the remarkable success of arthropods on Earth.
Asunto(s)
Escarabajos/anatomía & histología , Imagenología Tridimensional , Integumento Común/anatomía & histología , Estrés Mecánico , Imagen de Lapso de Tiempo , Animales , Módulo de Elasticidad , Tomografía Computarizada por Rayos XRESUMEN
Using in vitro procedures to prepare newly excysted metacercariae and gut-penetrated juvenile Fasciola gigantica, the ultrastructural features of the tegumental syncytium and perikarya of these ephemeral stages in the host-invasion process were compared. The T0-type tegumental cells in newly excysted metacercariae are packed with stored T0 granules which, following transport to the surface membrane of the syncytium, discharge by exocytosis to maintain the glycocalyx. The T0 cells become depleted of T0 granules during the penetration process, shrink in size, and initiate autophagy in the cytoplasm to facilitate metamorphosis from a storage function to active biosynthesis. The novel products appear to include lysosomes which contribute to the autophagosomes, and T1 granules, necessary for maintenance of the glycocalyx and immunoprotection, as the invasion process continues into the host liver. Residual bodies, the end-products of autophagy, are eliminated from the apical membrane of the tegumental syncytium into the host-parasite interface. There they may present a transient source of parasite-derived molecules, including lysosomal cathepsin-type proteases, with potential for interaction with the host's immune system, and so might be exploited as targets for vaccinal and immunomodulatory studies.