Diversity of dermal denticle structure in sharks: Skin surface roughness and three‐dimensional morphology

Shark skin is covered with numerous placoid scales or dermal denticles. While previous research has used scanning electron microscopy and histology to demonstrate that denticles vary both around the body of a shark and among species, no previous study has quantified three‐dimensional (3D) denticle structure and surface roughness to provide a quantitative analysis of skin surface texture. We quantified differences in denticle shape and size on the skin of three individual smooth dogfish sharks (Mustelus canis) using micro‐CT scanning, gel‐based surface profilometry, and histology. On each smooth dogfish, we imaged between 8 and 20 distinct areas on the body and fins, and obtained further comparative skin surface data from leopard, Atlantic sharpnose, shortfin mako, spiny dogfish, gulper, angel, and white sharks. We generated 3D images of individual denticles and measured denticle volume, surface area, and crown angle from the micro‐CT scans. Surface profilometry was used to quantify metrology variables such as roughness, skew, kurtosis, and the height and spacing of surface features. These measurements confirmed that denticles on different body areas of smooth dogfish varied widely in size, shape, and spacing. Denticles near the snout are smooth, paver‐like, and large relative to denticles on the body. Body denticles on smooth dogfish generally have between one and three distinct ridges, a diamond‐like surface shape, and a dorsoventral gradient in spacing and roughness. Ridges were spaced on average 56 µm apart, and had a mean height of 6.5 µm, comparable to denticles from shortfin mako sharks, and with narrower spacing and lower heights than other species measured. We observed considerable variation in denticle structure among regions on the pectoral, dorsal, and caudal fins, including a leading‐to‐trailing edge gradient in roughness for each region. Surface roughness in smooth dogfish varied around the body from 3 to 42 microns.     

Biological characterization of the skin of shortfin mako shark Isurus oxyrinchus and preliminary study of the hydrodynamic behaviour through computational fluid dynamics

This study characterized the morphology, density and orientation of the dermal denticles along the body of a shortfin mako shark Isurus oxyrinchus and identified the hydrodynamic parameters of its body through a computational fluid‐dynamics model. The study showed a great variability in the morphology, size, shape, orientation and density of dermal denticles along the body of I. oxyrinchus. There was a significant higher density in dorsal and ventral areas of the body and their highest angular deviations were found in the lower part of the mouth and in the areas between the pre‐caudal pit and the second dorsal and pelvic fins. A detailed three‐dimensional geometry from a scanned body of a shark was carried out to evaluate the hydrodynamic properties such as drag coefficient, lift coefficient and superficial (skin) friction coefficient of the skin together with flow velocity field, according to different roughness coefficients simulating the effect of the dermal denticles. This preliminary approach contributed to detailed information of the denticle interactions. As the height of the denticles was increased, flow velocity and the effect of lift decreased whereas drag increased. The highest peaks of skin friction coefficient were observed around the pectoral fins.     

The denticle surface of thresher shark tails: Three-dimensional structure and comparison to other pelagic species

Shark skin denticles (scales) are diverse in morphology both among species and across the body of single individuals, although the function of this diversity is poorly understood. The extremely elongate and highly flexible tail of thresher sharks provides an opportunity to characterize gradients in denticle surface characteristics along the length of the tail and assess correlations between denticle morphology and tail kinematics. We measured denticle morphology on the caudal fin of three mature and two embryo common thresher sharks (Alopias vulpinus), and we compared thresher tail denticles to those of eleven other shark species. Using surface profilometry, we quantified 3D-denticle patterning and texture along the tail of threshers (27 regions in adults, and 16 regions in embryos). We report that tails of thresher embryos have a membrane that covers the denticles and reduces surface roughness. In mature thresher tails, surfaces have an average roughness of 5.6 μm which is smoother than some other pelagic shark species, but similar in roughness to blacktip, porbeagle, and bonnethead shark tails. There is no gradient down the tail in roughness for the middle or trailing edge regions and hence no correlation with kinematic amplitude or inferred magnitude of flow separation along the tail during locomotion. Along the length of the tail there is a leading-to-trailing-edge gradient with larger leading edge denticles that lack ridges (average roughness = 9.6 μm), and smaller trailing edge denticles with 5 ridges (average roughness = 5.7 μm). Thresher shark tails have many missing denticles visible as gaps in the surface, and we present evidence that these denticles are being replaced by new denticles that emerge from the skin below.     

The characterization, replication and testing of dermal denticles of Scyliorhinus canicula for physical mechanisms of biofouling prevention

There is a current need to develop novel non-toxic antifouling materials. The mechanisms utilized by marine organisms to prevent fouling of external surfaces are of interest in this regard. Biomimicry of these mechanisms and the ability to transfer the antifouling characteristics of these surfaces to artificial surfaces are a highly attractive prospect to those developing antifouling technologies. In order to achieve this, the mechanisms responsible for any antifouling ability must be elucidated from the study of the natural organism and the critical surface parameters responsible for fouling reduction. Dermal denticles of members of the shark family have been speculated to possess some natural, as yet unidentified antifouling mechanism related to the physical presence of denticles. In this study, the dermal denticles of one particular member of the slow-swimming sharks, Scyliorhinus canicula were characterized and it was found that a significant natural variation in denticle dimensions exists in this species. The degree of denticle surface contamination was quantified on denticles at various locations and it was determined that the degree of contamination of the dorsal surface of denticles varies with the position on the shark body. In addition, we successfully produced synthetic sharkskin samples using the real skin as a template. Testing of the produced synthetic skin in field conditions resulted in significant differences in material attachment on surfaces exhibiting denticles of different dimensions.     

Teeth outside the mouth: The evolution and development of shark denticles

Vertebrate skin appendages are incredibly diverse. This diversity, which includes structures such as scales, feathers, and hair, likely evolved from a shared anatomical placode, suggesting broad conservation of the early development of these organs. Some of the earliest known skin appendages are dentine and enamel-rich tooth-like structures, collectively known as odontodes. These appendages evolved over 450 million years ago. Elasmobranchs (sharks, skates, and rays) have retained these ancient skin appendages in the form of both dermal denticles (scales) and oral teeth. Despite our knowledge of denticle function in adult sharks, our understanding of their development and morphogenesis is less advanced. Even though denticles in sharks appear structurally similar to oral teeth, there has been limited data directly comparing the molecular development of these distinct elements. Here, we chart the development of denticles in the embryonic small-spotted catshark (Scyliorhinus canicula) and characterize the expression of conserved genes known to mediate dental development. We find that shark denticle development shares a vast gene expression signature with developing teeth. However, denticles have restricted regenerative potential, as they lack a sox2+ stem cell niche associated with the maintenance of a dental lamina, an essential requirement for continuous tooth replacement. We compare developing denticles to other skin appendages, including both sensory skin appendages and avian feathers. This reveals that denticles are not only tooth-like in structure, but that they also share an ancient developmental gene set that is likely common to all epidermal appendages.     

Cell polarity during wound healing in an insect epidermis

The insect integument displays uniform posterior orientation of cuticular denticles or bristles formed by the epidermal cells. We want to understand how cell polarities become uniformly oriented in the plane of the epidermal sheet. Here we test whether directed cell migration disturbs the orientation of denticles. Burning a circular area of epidermal cells beneath the cuticle causes cells to migrate into the resulting wound and the cuticle pattern observed after the subsequent moult depends on the time interval between burning and ecdysis. After a short wound-healing period cuticular protrusions tend to point away from the wound. With increasing would healing periods they tend to point more and more towards the wound centre. These results suggest that the migrating cells tend to orient cuticular protrusions in the direction of cell movement while continued cell movement will bend nascent cuticular protrusions outwards. Cell shape may also determine denticle orientation. I propose that the asymmetric localization of cell components known to determine the orientation of cell migration may also determine denticle orientation in insect epidermal cells.     

Placoderm fishes,pharyngeal denticles,and the vertebrate dentition

The correlation of the origin of teeth with jaws in vertebrate history has recently been challenged with an alternative to the canonical view of teeth deriving from separate skin denticles. This alternative proposes that organized denticle whorls on the pharyngeal (gill) arches in the fossil jawless fish Loganellia are precursors to tooth families developing from a dental lamina along the jaw, such as those occurring in sharks, acanthodians, and bony fishes. This not only indicates that homologs of tooth families were present, but also illustrates that they possessed the relevant developmental controls, prior to the evolution of jaws. However, in the Placodermi, a phylogenetically basal group of jawed fishes, the state of pharyngeal denticles is poorly known, tooth whorls are absent, and the presence of teeth homologous to those in extant jawed fishes (Chondrichthyes + Osteichthyes) is controversial. Thus, placoderms would seem to provide little evidence for the early evolution of dentitions, or of denticle whorls, or tooth families, at the base of the clade of jawed fishes. However, organized denticles do occur at the rear of the placoderm gill chamber, but are associated with the postbranchial lamina of the anterior trunkshield, assumed to be part of the dermal cover. Significantly, these denticles have a different organization and morphology relative to the external dermal trunkshield tubercles. We propose that they represent a denticulate part of the visceral skeleton, under the influence of pharyngeal patterning controls comparable to those for pharyngeal denticles in other jawed vertebrates and Loganellia.     

Anomalies in the expression of abdominal denticle belts of partial larvae produced by fragmentation of theDrosophila melanogaster egg

Summary Abnormal denticle belt patterns can occasionally be observed in abdominal belts of partial larvae obtained from egg fragments. The abdominal belts have the following features in common: 1) The number of denticles of an abdominal denticle belt may increase, depending on the space occupied by a distinct segment or the whole body region. The arrangement of the denticles in such enlarged belts is less regular than in normal belts. 2) Enlarged denticle belts are also found in the terminal segment of a fragment, or in the segment next to it when the larval pattern is interrupted by fragmentation. The denticle belt in the adjacent segment(s) may then be supressed. 3) All denticles in a belt (or part of a belt) are orientated posteriorly if the distance to the posteriorly adjacent belt (or part of a belt) is larger than normal, or if this denticle belt is suppressed. Conditions anterior to a segment do not seem to exert any influence on denticle orientation.     

Morphogenese und Musterbildung des Hautzähnchen-Skelettes von Heterodontus

Different denticle shapes in Heterodontus (Selachii) are derived from a single primary form by five geometrical transformations. Insertion of new denticles into the existing pattern is controlled by reference points transmitting positional information which is space- and time-dependent. Injuries interrupt the transmission of certain informations, causing anomalous denticles, which do not occur in normal ontogeny.
Dic verschiedenen Zähnchenformen lassen sich von einer einzigen Grundform durch fünf verschiedene geometrische Transformationen ablciten. Das Einfügen neuer Zähnchen in das existicrende Muster wird durch Referenzpunkte kontrolliert, die Lage- und Zeit-abhängige lnfoimationen übertragen. Verwundungen unterbrechcn die Übermittlung be-stimmter Informationen. Dies führt zu anomalen Zähnchen, die in der normalen Onto-gencsc nicht vorkommen.     

General descriptions of the dermis structure of a juvenile whale shark Rhincodon typus from the Gulf of California

The aim of this study is to provide preliminary observations on the microanatomy of Rhincodon typus skin using histology and electron microscopy analyses. Skin biopsies were obtained from a deceased juvenile male shark (548 cm total length) stranded in La Paz, Mexico, during February 2018. The results of this study evidenced the basic structure of the dermal denticles in the epidermis of the trunk of the shark, as well as the composition of the connective tissue in the hypodermis. Histological images of the hypodermis showed a high concentration of collagen fibres, formed by a large number of fine and wavy fibres of compact shape and little intercellular substance.     

A new species of <Emphasis Type="Italic">Trichodina</Emphasis> Ehrenberg, 1830 (Ciliophora: Trichodinidae) infecting farmed <Emphasis Type="Italic">Clarias gariepinus</Emphasis> (Burchell) (Siluriformes: Clariidae) in Cuba

A new species of Trichodina Ehrenberg, 1830 collected from the skin and fins of farmed North African catfish Clarias gariepinus (Burchell) fingerlings, is described. The new species can be distinguished from other trichodinids by the characteristics of the adhesive disc, especially by the great number of denticles. Trichodina merciae n. sp. is morphologically similar to T. renicola (Mueller, 1931) and T. marplatensis Martorelli, Marcotegui & Alda, 2008, in the number of denticles, but differs in the morphometric data, denticle morphology, environment and location. Trichodina merciae n. sp. has broad sickle-shaped blades and thin, straight rays, while T. marplatensis has broad club-shaped blades and wide S-shaped rays. Besides, denticle length, blade length, ray length, width of central part and denticle span of the new species are greater than T. marplatensis. However, the diameter of denticle ring and the diameter of the central area in T. marplatensis is larger than the ones in T. merciae n. sp. This is the first record of freshwater ectoparasite trichodinid with an average number of denticles greater than 50.     

Cell shape and epithelial patterning in the Drosophila embryonic epidermis

The development of denticle rows on the ventral Drosophila embryo is a valuable system for studying the genetic control of epithelial patterning. During late embryogenesis, the apical surfaces of denticle-producing cells acquire a distinctive rectangular morphology with long anteroposterior boundaries, along which the denticles form, and short ventrolateral boundaries that stain strongly for adherens junction proteins. We observe that ventrolateral denticle cell boundaries are also convoluted, suggesting that the strong adherens staining results, at least in part, from the additional membrane in these regions. Embryos mutant for the Planar Cell Polarity (PCP) Effector gene multiple wing hairs (mwh), or expressing dominant negative form of the small GTPase Rac1, have cells present between the normal denticle cell rows. These 'Interloper Cells' do not have convoluted ventrolateral boundaries with strong adherens protein staining, but have normal denticle placement, suggesting that adherens protein localization is not critical for denticle cell PCP. Based on these and other observations, we propose that denticle cell morphology arises from an epithelial stretch without junction remodeling. A crude mechanical model suggests that this mechanism can generate both the straight anteroposterior boundaries and the compacted ventrolateral boundaries typical of denticle cells. We discuss the significance of cell adhesion for denticle cell morphogenesis, especially given the established role for Rac1 in cell adhesion.     

<Emphasis Type="Italic">Icriodus marieae</Emphasis>, a new icriodontid conodont species from the Middle Devonian

A new conodont species, Icriodus marieae, is described from pelagic limestone beds of the Carnic Alps (Austria). Specimens are obtained from the upper part of the Valentin Formation (Central Carnic Alps) and range from the latest Eifelian to middle Givetian. Significantly differing from other icriodontid conodonts is that the icriodontan element of the new species develops only three denticles on either lateral denticle row, which are constricted to the central part of the element. The anterior part of the element is free of lateral row denticles and consists of two to four denticles, which have a fan-shaped outline in lateral view. The anterior part as well as the posterior part (consisting of cusp and two to three pre-cusp denticles) is higher than the denticles of the central part of the element. Shape analysis confirms that the parameters chosen for landmarks (element size relation and denticle setting) show little variation between different specimens.     

Gill raker structure and development in indo-pacific anchovies (Teleostei: Engrauloidea), with a discussion of the structural evolution of engrauloid gill rakers

The postlarval development of gill raker denticles is described for the engrauloid (anchovy) genera Coilia, Lycothrissa, Setipinna, Thryssa, Stolephorus, and Encrasicholina based on scanning electron microscopy. The raker structure of adult Papuengraulis is also described. In the coiliid genera Coilia, Lycothrissa, Setipinna, Thryssa, and Papuengraulis, denticle development is not confined to particular region(s) of the raker. With few exceptions, the proliferation of denticles with growth is greatest along the upper raker edge; denticles are smaller and less dense on the raker faces and along the lower raker edge. Some Thryssa and Setipinna have a derived condition of denticle clustering along the upper raker edge. In Stolephorus and Encrasicholina, denticle development is confined to the upper raker half and includes the development of a single row of denticles along each raker face. A phylogenetic analysis of engrauloid raker structure, incorporating data from Bornbusch ( 88: Copeia 1988:174–182) and based on outgroup comparisons, indicates that for the Engrauloidea: (1) the pattern of denticle development shared by coiliids is plesiomorphic; and (2) the pattern of denticle development shared by Stolephorus, Encrasicholina, and most other engraulids is synapomorphic for the Engraulidae. There is no evidence that the studied coiliids Stolephorus and Encrasicholina are suspension feeders. The engraulid pattern of raker denticle development which is retained in suspension feeding engraulids of the genus Engraulis was thus derived before the derivation of suspension feeding in Engraulis. Comparative morphological and phylogenetic studies of clupeomorph raker structures and feeding behaviors can infer the historical origins of morphology-behavior associations, help define possible directions for analyses of raker denticle function, and thereby help elucidate the significance of structure-function couplings in the evolution of such clupeomorph trophic behaviors as suspension feeding. © 1992 Wiley-Liss, Inc.     

灌草与林带搭配条件下防护效应的数值模拟

在对林带和灌草进行参数化的基础上,将有、无灌草带存在情况下绕林带流场进行了数值模拟,分析比较了两种流场中沿流相对风速、不同断面风速廓线及湍动能的变化。结果表明,灌草在防护林体系中起着重要作用,在林带迎风面和带后一定区域内,均使风速减小。最后将数值模拟结果与实验结果进行了对比,并从实验角度分析了灌草的作用。     

行带式柠条固沙林防风效果

传统观点研究认为：植被覆盖度低于40％,沙地处于半固定半流动状态．但在实践观察中发现：在低密度（或覆盖度）时,灌丛的水平分布格局对固定流沙和阻止风沙流的作用差异显著。以低覆盖度（20％-25％）的柠条固沙林为研究对象,采用多点式自记风速仪（GB-228）,在2。4月份的盛风季节,测定了行带式配置和随机分布的柠条固沙林以及完整的行带式配置和其中一带出现缺口的行带式固沙林内不同部位、不同高度的风速．统计分析风速测定结果发现：（1）当覆盖度在20％-25％时,行带式配置的柠条固沙林内的防风效果比同覆盖度随机分布的在20cm高处高48．4％;50era高处高30．7％;200cm高处高27．4％,且风速越大,行带式配置降低风速的效果越显著,当200cm高风速达到6—7m／s时,行带式的平均防风效果比同覆盖度随机分布的高48．2％;反映出在低覆盖度时,灌丛的水平分布格局成为制约固沙林沙防风效果的重要因素,行带式配置具有显著的防风效果;（2）在灌丛与灌丛之间形成的类似“狭管”流场的局部,有提升风速的作用,导致其林内观测的风速有约41．3％的观测结果超过旷野对照风速,行带式配置的柠条固沙林内没有一个观测结果超过对照风速;这个结果反映出随机分布的柠条固沙林内流场结构复杂、变化多样,也成为低覆盖度时,沙地处于半固定、半流动状态,疏林内同时存在风蚀和积沙和缺口处风速升高的重要因素;（3）行带式配置林内地表粗糙度比随机配置的高5．4-114．4倍,说明行带式配置具有显著的防止风蚀、固定流沙的作用;（4）行带式林内出现断带缺口处,其缺口处的风速降低有明显的累加现象;风速抬升现象在一定程度受到制约。这些结果为发展低覆盖度行带式配置的固沙林提供重要的科学依据。     

No sitting on the fence: protecting wetlands from feral pig damage by exclusion fences requires effective fence maintenance

Feral pigs damage the significant ecological and cultural values of tropical Australian wetlands. Control measures such as culling, baiting, and trapping can reduce overall pig populations, but do not eliminate the substantial physical damage to wetlands that can occur from just a few individuals. Exclusion fences have been adopted as a potential technique to prevent damage to selected wetlands. To test the effectiveness of exclusion fences we measured the physical damage caused by pigs to multiple wetlands in the Archer River catchment of tropical northern Australia. Wetlands were fenced using a typical cattle exclusion fence, a specific pig exclusion fence or had no fence. Initial analyses of these fence treatments showed no significant difference in the intensity of physical pig damage to exposed wetland sediments and fringing vegetation. However, several of the pig exclusion fences were found to have been compromised. Reanalysis indicated wetlands with functioning pig exclusion fences had no physical pig damage and this was significantly less damage than in all other treatments. In contrast, wetlands with compromised pig exclusion fences had damage that was statistically equivalent to sites without fences or with cattle exclusion fences, but in individual cases had the worst damage recorded in any of the treatments. Compromised pig exclusion fencing of wetlands can thus be worse than having no fencing at all. This suggests that the successful prevention of pig damage to wetlands by exclusion fences requires ongoing and effective fence monitoring and maintenance regimes.

     

Scanning electron microscopy of the lip denticiles of Ascaris suum adults of known ages.

Purebred Hampshire pigs, farrowed and maintained under conditions precluding extraneous helminth infection, were exposed to a single dose of 10,000 Ascaris suum infective eggs. The pigs were killed at intervals of 28, 41, 55, 86, 115, 145, 175, and 206 days after infection. At necropsy, no gross lesions were found in the lungs or livers of infected pigs. The worms were recovered from the small intestine, identified, counted, and fixed. The heads were excised, critical point dried, mounted en face, and examined by scanning electron microscopy. Worms 28 to 115 days old had unworn denticles that were triangular when viewed laterally but blunt when viewed tangentially. Wearing of the denticles was observed first with 145-day-old worms; wearing increased with age both in numbers of denticles affected and in degree of wear so that by 206 days after inoculation, almost all denticles in the center of the lip were worn. Worn denticles appear truncated when viewed from any angle. The denticles outside the central area were not affected by wear. The size of the denticles varies not only between specimens of the same age, but also on each specimen. However, average denticle size is directly related to the size and, accordingly, to the age of the worm. External to each denticle is a corresponding depression that we have called the denticular groove. One 28-day-old specimen had some extra denticles aligned irregularly along the lip; this irregularity gave the appearance of a double row. The denticles of the two subventral lips are similar to those of the dorsal and are equally affected by wear. There was no detectable difference in denticles of male and female worms. Since wear can now be specifically correlated with age, we conclude that the denticles are functional and become worn through use. Consequently, adult A. suum may be an even more injurious pathogen than heretofore supposed.