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Scale pattern of five species of Zeiformes (Teleostei) 五种叠形目(遥骨目)的鳞片格局
IF 1.3 4区 生物学 Q4 ANATOMY & MORPHOLOGY Pub Date : 2024-11-23 DOI: 10.1111/azo.12526
Lucas C. Garcia, Cristiano R. Moreira

The order Zeiformes comprises a small order of predominantly deep-sea fishes. However, little information is available on their morphology, such as squamation. Therefore, in the present study, we analysed and described the squamation pattern and scale morphology of five species of Zeiformes occurring on the Brazilian coast: Allocyttus verrucosus, Grammicolepis brachiusculus, Xenolepidichthys dalgleishi, Zenion hololepis and Zenopsis conchifer. Our analysis revealed greater morphological diversity than previously reported for this order. All species, except the scaleless Zenopsis conchifer, presented cycloid scales of spinoid subtype. Grammicolepis and Zenion presented exclusively scales of the spinoid subtype, whereas both Allocyttus and Xenolepidichthys also displayed true cycloid subtype. A. verrucosus is characterised by two distinct groups of scales of different sizes that form scale clusters. G. brachiusculus and X. dalgleishi presented a unique vertically elongated scale covering most of the body. Small bony plates were observed at the dorsal- and anal-fin bases of G. brachiusculus, X. dalgleishi and Z. hololepis. The variation observed is putatively useful for the systematics of the order.

形目包括一个小目,主要是深海鱼类。然而,关于它们的形态,如鳞片的信息很少。因此,本研究分析和描述了巴西沿海五种泽形目的鳞片形态和鳞片形态:Allocyttus verrucosus、Grammicolepis brachiusculus、Xenolepidichthys dalleishi、Zenion hololepis和Zenopsis conchifer。我们的分析显示,该目的形态多样性比以前报道的要大。除无鳞的苍苍兰外,其他种属均为棘类亚种的摆线鳞。Grammicolepis和Zenion仅表现为棘状鳞亚型,而Allocyttus和Xenolepidichthys也表现为真正的摆线鳞亚型。疣状棘球蚴的特点是两组不同大小的鳞片形成鳞片簇。腕骨棘鱼和大鳞棘鱼具有独特的覆盖大部分身体的垂直细长鳞片。在腕骨鱼、大鳍鱼和hololepis鱼的背鳍和肛门鳍基部观察到小的骨板。所观察到的变化被认为对该目的分类学是有用的。
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引用次数: 0
Morphology and chronology of critical events for early-life history survival of an old word killifish, Esmaeilius shirini (Teleostei: Aphaniidae): A conservation management approach 古鳉早期生活史生存的形态学和关键事件年代学:保护管理方法
IF 1.3 4区 生物学 Q4 ANATOMY & MORPHOLOGY Pub Date : 2024-11-19 DOI: 10.1111/azo.12524
Hamid Reza Esmaeili, Tayebeh Asrar, Golnaz Sayyadzadeh

In this study, the live cell imaging microscopy of an old word cyprinodontiform fish, the Shirini toothcarp Esmaeilius shirini during its embryogenesis, is described and illustrated for the first time through the detailed morphological description and chronology of the events of all developmental stages (from fertilization to hatching). External fertilization of transparent, spherical, sticky and macrolecithal eggs bearing adhesive filaments with an average diameter of 0.83–1 mm is the first step in the ontogenesis of E. shirini. Embryogenesis continued with a process of meroblastic cleavage (approximately 2 and half h post-fertilization, hpf), formation of blastula (blastulation), gastrula (gastrulation), neurulation, somatogenesis, organ formation (organogenesis) and pre-hatching. The development continued with hatching (a stage between the embryonic and larval periods) at approximately 131 haf (at 24 ± 1°C). The new larvae were about 3.9–5.8 mm in length with distinct fin rays and pigmentation on the head, body and fins. Basic data from this study can be used in forthcoming investigations on the reproduction processes, breeding, behaviour and conservation of E. shirini. Fish development comprises an integrated, ordered and well-mapped sequence of early life-history events controlled by intrinsic and extrinsic factors. Knowledge of fish developmental stages during embryogenesis/ontogeny is essential for a better understanding of their biology, breeding criteria, phylogenetic relationships, dietary requirements, conservation plans, environmental requirements and ecological preferences.

在这项研究中,通过详细的形态学描述和所有发育阶段(从受精到孵化)事件的年表,首次描述和说明了一种古老的cyprinodontiform鱼,Shirini齿鱼esmaelius Shirini胚胎发育时期的活细胞成像显微镜。透明、球形、粘性和大卵泡卵的体外受精,卵上有平均直径0.83-1 mm的粘连细丝,是shirini个体发生的第一步。胚胎发生的过程包括:原细胞分裂(受精后约2.5小时)、囊胚形成(blastulation)、原胚形成(gastrulation)、神经发育、躯体发育、器官形成(organogenesis)和预孵化。孵化(胚胎期和幼虫期之间的阶段)在大约131°C(24±1°C)下继续发育。新幼虫体长约3.9 ~ 5.8 mm,鳍纹明显,头、身、鳍均有色素沉着。本研究所得的基本数据可用于今后对希里尼的繁殖过程、繁殖、行为和保护的调查。鱼类的发育包括由内在和外在因素控制的一个完整的、有序的、绘制良好的早期生活史事件序列。了解鱼类在胚胎发生/个体发育过程中的发育阶段对于更好地了解它们的生物学、繁殖标准、系统发育关系、饮食要求、保护计划、环境要求和生态偏好至关重要。
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引用次数: 0
Role of mucous cells in the defence system of the cnidarian Actinia equina (Linnaeus, 1758): The keystone of evolution 黏液细胞在刺胞动物马线虫防御系统中的作用(林奈,1758):进化的基石
IF 1.3 4区 生物学 Q4 ANATOMY & MORPHOLOGY Pub Date : 2024-11-08 DOI: 10.1111/azo.12523
Alessio Alesci, Sebastian Marino, Anthea Miller, Angelo Giliberti, Gabriele Rigano, Letterio Giuffrè, Jorge M. O. Fernandes, Serena Savoca, Gioele Capillo

Cnidarians are the first phylum of metazoans to exhibit a functional mucus layer. In evolution, mucous cells perform the classical secretory function of mucus and act as sentinels for external insults, particularly in mammals. However, this topic has not been much analysed in the lower metazoans. This study aims to investigate the role of mucous cells in Actinia equina (Anthozoa, Hexacorallia) (Linnaeus, 1758) by using confocal microscopy and bioinformatics. The results show colocalization of Toll-like receptor 2 (TLR2) and Mucin2 (MUC2) on these cells, suggesting a crosstalk between defence response and mucus secretion in cnidarians. Furthermore, the presence of TLR2 on the surface of mucous cells suggests their involvement in host defence responses, as occurs in vertebrates. Moreover, through in silico analysis, we confirmed the presence of orthologous genes for the tested antibodies, providing additional evidence to support our results from an evolutionary perspective. These data suggest that mucous cells play a defensive role throughout evolution, helping us understand how these cells have developed and adapted across phylogenesis. This information also provides new insights into the defensive strategies of invertebrates and clarifies their evolutionary history.

刺胞动物是后生动物中第一个具有功能性黏液层的门。在进化过程中,黏液细胞具有典型的分泌黏液的功能,并作为外部损伤的哨兵,特别是在哺乳动物中。然而,这一主题在低等后生动物中还没有得到太多的分析。本研究旨在利用共聚焦显微镜和生物信息学方法研究马放线菌(Actinia equina (anthzoa, Hexacorallia) (Linnaeus, 1758)中黏液细胞的作用。结果显示toll样受体2 (TLR2)和Mucin2 (MUC2)在这些细胞上共定位,提示刺胞动物的防御反应和粘液分泌之间存在串扰。此外,粘膜细胞表面TLR2的存在表明它们参与宿主防御反应,就像发生在脊椎动物身上一样。此外,通过计算机分析,我们证实了测试抗体的同源基因的存在,从进化的角度为我们的结果提供了额外的证据。这些数据表明,黏液细胞在整个进化过程中起着防御作用,帮助我们了解这些细胞是如何在整个系统发生中发展和适应的。这一信息也为无脊椎动物的防御策略提供了新的见解,并阐明了它们的进化史。
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引用次数: 0
Scanning electron microscopy comparison of an abnormal and normal otoliths of the Anguilla anguilla collected from the Bafa Lake in Western Anatolia 安那托利亚西部巴法湖鳗鲡异常与正常耳石的扫描电镜比较
IF 1.3 4区 生物学 Q4 ANATOMY & MORPHOLOGY Pub Date : 2024-10-12 DOI: 10.1111/azo.12522
Laith A. Jawad, Salim Serkan Güçlü, Habil Uğur Koca, Mehmet Cilbiz

Aberrations in the otolith can harmfully influence the imperative actions of the fish species. The chief drive of the current investigation is to describe the deformities in the otoliths of the European eel Anguilla anguilla. Out of 76 Anguilla anguilla individuals collected from Bafa Lake in Western Anatolia, Türkiye, four female specimens exhibited abnormal otoliths. Scanning electron microscope was used to determine the deformed otoliths examined. The outcomes of the abnormality analyses in A. anguilla otoliths, deformities were noticed such as numerous prominence structures and bumps of different sizes on the surface of the otolith produced by buildup of the different crystal structures in some parts of the otolith. These deformities were found in the left and/or right otoliths of the female individuals of different total lengths. The percentage of individuals with abnormal otoliths of A. anguilla is calculated as 5.3%. The current study presents for the first-time abnormal otolith information on left and right otoliths in female A. anguilla. Deformities in the otoliths of European eels are believed to result from a combination of genetic predisposition and stress caused by various environmental factors, such as water pollution, changes in salinity, temperature fluctuations and nutritional imbalances.

耳石的异常会对鱼类的重要活动产生有害的影响。目前调查的主要动力是描述欧洲鳗鲡耳石的畸形。在基耶省西部安纳托利亚巴法湖采集的76只安圭拉鳗鲡中,有4只雌性标本出现异常耳石。用扫描电子显微镜观察变形耳石。对鳗鲡耳石的异常分析结果表明,由于耳石某些部位的不同晶体结构的积累,在耳石表面出现了许多凸起结构和大小不等的凸起等畸形。这些畸形在不同总长度的女性个体的左耳石和/或右耳石中发现。安圭拉耳石异常个体的比例计算为5.3%。本研究首次报道了雌性安圭拉左右耳石的异常信息。欧洲鳗鱼的耳石畸形被认为是由遗传倾向和各种环境因素(如水污染、盐度变化、温度波动和营养不平衡)造成的压力共同造成的。
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引用次数: 0
Application of otolith morphometrics in delineating population structure of Aphaniops stoliczkanus (Teleostei: Aphaniidae) in southern Iranian waters 耳石形态计量学在划定伊朗南部海域斯托利卡鱼种群结构中的应用(Teleostei: aphaniopstoliczkanus)
IF 1.1 4区 生物学 Q4 ANATOMY & MORPHOLOGY Pub Date : 2024-10-11 DOI: 10.1111/azo.12521
Mahdi Iranmanesh, Majid Askari Hesni

Widely distributed Species in different geological and ecological environments often exhibit great intraspecific diversity. Aphaniops stoliczkanus (Day, 1872) is an endemic species of Middle East occurring over a vast geographic range in different basins. This species has been observed in freshwater, brackish water and hot sulphur springs; therefore, its populations are known to exhibit considerable morphological variation. In the present study, we have investigated the poorly studied populations of A. stoliczkanus of the Hormuz and Makran basins of Iran in terms of otolith morphology and morphometrics. According to the results, these populations can be divided into two distinct groups including east and west of the Strait of Hormuz. Our results also provide evidence that there is a relationship between otolith morphology and the environment, serving as environmental factors that may influence the formation of specific regions in the otolith shape. Despite the remarkable morphological similarities among otoliths in geographically close populations of Charak and Chiruyeh, our data purposed that geographically these populations have different dorsal tip shapes in their otolith. In comparison to the data from other studies, the otoliths of Qeshm Island appear to resemble the overall shape of otolith found in Oman very closely. However, it is believed that Qeshm Island population was separated from the Hormuz basin populations during the Holocene Sea level rise. Also, the outcomes of this study provide insights into the potential role of environmental factors in shaping intraspecific diversity and the migration patterns of this species within the Persian Gulf.

分布广泛的物种在不同的地质和生态环境中往往表现出很大的种内多样性。Aphaniops stoliczkanus (Day, 1872)是一种中东特有物种,分布在不同盆地的广阔地理范围内。在淡水、半咸水和硫磺温泉中均有发现;因此,已知其种群表现出相当大的形态变异。在本研究中,我们从耳石形态和形态计量学的角度调查了伊朗霍尔木兹和马可兰盆地研究较少的A. stoliczkanus种群。根据结果,这些种群可以分为两个不同的群体,包括霍尔木兹海峡的东部和西部。我们的研究结果也提供了耳石形态与环境之间存在关系的证据,作为环境因素可能影响耳石形状中特定区域的形成。尽管在地理上相近的Charak和Chiruyeh人群中耳石在形态上有显著的相似性,但我们的数据表明,在地理上这些人群的耳石中有不同的背尖形状。与其他研究的数据相比,格什姆岛的耳石似乎与阿曼发现的耳石的整体形状非常相似。然而,人们认为在全新世海平面上升期间,格什姆岛种群与霍尔木兹盆地种群分离。此外,本研究的结果为环境因素在形成种内多样性和该物种在波斯湾内的迁移模式中的潜在作用提供了见解。
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引用次数: 0
Description of aberrant otolith morphology of Saurida lessepsianus (Aulopiformes, Synodontidae) collected from the coast of Muğla province, Aegean Sea, Turkey 土耳其爱琴海Muğla省海岸采集的小鳞蜥耳石异常形态描述
IF 1.1 4区 生物学 Q4 ANATOMY & MORPHOLOGY Pub Date : 2024-09-19 DOI: 10.1111/azo.12520
İsmail Reis, Laith Jawad, Celal Ateş

This is the first study investigating the aberrant otolith morphology in the Lessepsian lizardfish Saurida lessepsianus in the Aegean Sea, Turkey. A Leica Emspira 3 digital microscope was used to determine the aberrant otoliths examined. Two female specimens measuring 230 and 274 mm in total length (TL) were examined and showed an aberrant otolith on both sides of their head. These deformities might have been instigated by the impacts of changing water factors related to diverse pollutants in the Aegean Sea. Characteristics of otolith deformities are of immeasurable significance for aquatic habitats and other otolith studies.

这是对土耳其爱琴海小蜥蜴鱼(Saurida lesssepsianus)异常耳石形态的首次研究。采用徕卡Emspira 3型数码显微镜对异常耳石进行检测。两只雌性标本的总长度分别为230和274毫米,在头部两侧发现了异常的耳石。这些畸形可能是由爱琴海中各种污染物相关的水因素变化的影响引起的。耳石畸形特征对水生生境和其他耳石研究具有不可估量的意义。
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引用次数: 0
Immune system of fish with special reference to estrogenic immune regulation: A review 鱼类的免疫系统,特别是雌激素免疫调节:综述
IF 1.1 4区 生物学 Q4 ANATOMY & MORPHOLOGY Pub Date : 2024-09-06 DOI: 10.1111/azo.12519
Ragini Sinha, Dipak Kumar Mandal

Fish, first jawed vertebrate group appearing in evolution, have immune system similar to those of mammals. These early animals' innate and adaptive immune systems are completely developed, having head kidney and thymus as primary, spleen and mucosa associated lymphoid tissue as secondary immune organs. The adaptive immunity performs key role in defence against recurring infections and elimination of fish pathogens with the help of its memory cells, T-cell-receptors and immunoglobulins. Epidermal secretions including antibacterial peptides, lysozymes, lectins, complement and C-reactive proteins which function in disruption of antigens, phagocytosis, inflammatory response and mending of tissue injury play a significant role as elements of fish immune system. Fish, however, have greater innate immunity than humans do. Estrogens play crucial role in immunomodulation through nuclear oestrogen receptors, which are found in majority of immune cells and lymphoid organs in fish. There is mounting proof that xenoestrogens, that can attach as agonists to oestrogen receptors, pose significant ecotoxicological risk by disrupting the defence mechanism in fish. Though less potent than natural oestrogen, they can bioaccumulate to finally reach a substantial dose. This review's objective is to give an overview of the fish defence system, its estrogenic regulation and estrogenic-endocrine disruption.

鱼类是进化过程中最早出现的有颌脊椎动物,其免疫系统与哺乳动物相似。这些早期动物的先天性免疫系统和适应性免疫系统已完全发育成熟,头部肾脏和胸腺是主要免疫器官,脾脏和粘膜相关淋巴组织是次要免疫器官。在记忆细胞、T 细胞受体和免疫球蛋白的帮助下,适应性免疫在抵御反复感染和清除鱼类病原体方面发挥着关键作用。表皮分泌物包括抗菌肽、溶菌酶、凝集素、补体和 C 反应蛋白,它们在破坏抗原、吞噬、炎症反应和修复组织损伤方面发挥着重要作用。然而,鱼类的先天免疫力比人类更强。雌激素通过核雌激素受体在免疫调节中发挥着重要作用,而雌激素受体存在于鱼类的大多数免疫细胞和淋巴器官中。越来越多的证据表明,异雌激素可以作为激动剂附着在雌激素受体上,破坏鱼类的防御机制,从而造成重大的生态毒理学风险。虽然异雌激素的效力低于天然雌激素,但它们可以通过生物累积最终达到相当大的剂量。本综述旨在概述鱼类防御系统、雌激素调节和雌激素-内分泌干扰。
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引用次数: 0
Otolith-based species identification in the killifish Aphaniops (Teleostei; Cyprinodontiformes; Aphaniidae) using both morphometry and wavelet analysis 利用形态计量学和小波分析法,基于耳石鉴定鳉鱼Aphaniops(腕足动物;鲤形目;Aphaniidae)的物种
IF 1.1 4区 生物学 Q4 ANATOMY & MORPHOLOGY Pub Date : 2024-09-04 DOI: 10.1111/azo.12518
Eleni A. Charmpila, Azad Teimori, Bettina Reichenbacher

The killifish genus Aphaniops consists of nine species distributed in Eastern Africa and the Middle East. However, distinguishing these species from each other based on morphological traits is challenging. Here we investigate the utility of otoliths (sagittae) in distinguishing between A. dispar, A. ginaonis, A. hormuzensis, A. kruppi and A. stoliczkanus. Our approach is based on otoliths from 89 specimens and involves (1) otolith morphometrics, following prior recommendations, (2) shape analysis of otolith contours based on discrete wavelet transformation—a novel method in killifish otolith research—and comparative statistical analyses. Both methods reveal significant interspecific variation in the otolith regions of the rostrum, antirostrum and excisura. While method (1) effectively discriminates most species, method (2) struggles to differentiate A. hormuzensis, A. stoliczkanus and A. kruppi. Additionally, both methods encounter challenges in correctly classifying A. hormuzensis due to the high otolith variability of this species in our sample. Possible factors accounting for their variability are environmental fluctuations at the sampled hot sulphuric spring (Khurgo) and potential introgressive hybridization. We conclude that otolith morphometry is a valuable tool for Aphaniops species identification. Furthermore, we found that the distinctiveness of species-specific otolith traits increases with the divergence age of the species.

鳉鱼属有九个物种,分布在非洲东部和中东地区。然而,根据形态特征区分这些物种具有挑战性。在此,我们研究了耳石(sagittae)在区分 A. dispar、A. ginaonis、A. hormuzensis、A. kruppi 和 A. stoliczkanus 方面的作用。我们的方法以 89 个标本的耳石为基础,包括:(1)耳石形态计量学,遵循先前的建议;(2)基于离散小波变换的耳石轮廓形状分析--这是杀生鱼耳石研究中的一种新方法--以及比较统计分析。这两种方法都揭示了喙、反喙和外喙等耳石区域存在明显的种间差异。虽然方法(1)能有效区分大多数物种,但方法(2)却难以区分荷尔蒙鱼(A. hormuzensis)、斯托利茨卡努斯鱼(A. stoliczkanus)和克鲁皮鱼(A. kruppi)。此外,这两种方法在正确分类荷尔鲁津鱼时都遇到了挑战,因为在我们的样本中,该物种的耳石变异性很高。造成其变异性的可能因素是取样硫酸热泉(Khurgo)的环境波动和潜在的引种杂交。我们的结论是,耳石形态测量是鉴定 Aphaniops 物种的重要工具。此外,我们还发现,物种特异性耳石特征的独特性会随着物种的分化年龄而增加。
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引用次数: 0
Not all heroes wear shells: New data on the spicules' morphology in acochlidimorph mollusc 并非所有英雄都有贝壳:关于无脊椎软体动物脊柱形态的新数据
IF 1.1 4区 生物学 Q4 ANATOMY & MORPHOLOGY Pub Date : 2024-08-13 DOI: 10.1111/azo.12517
Anna L. Mikhlina, Ekaterina D. Nikitenko, Elena V. Vortsepneva

Calcareous skeletal structures are common among different invertebrate groups, including Porifera, Cnidaria, Platyhelminthes, Mollusca, and Echinodermata. They appear externally as shells, plates, or scales, or internally as subepidermal spicules. Although typically formed outside of the cells, in one class of gastropod molluscs, the Heterobranchia, subepidermal and intracellular spicules are found—a rare case, at least for the molluscs. This type of spicules is present in several phylogenetically distant groups of heterobranch molluscs (Rhodopemorpha, Acochlidimorpha, and Nudibranchia), each different in terms of morphology and autecology. This raises questions about the origin and homology of subepidermal spicules in heterobranch molluscs. To answer these questions, understanding the process of spicule formation is crucial. This is impeded by insufficient data on the fine structure of the subepidermal spicules and adjacent epithelia. To address this problem, we carried out a comprehensive morphological study of subepidermal spicules and the body wall in one species of Acochlidimorpha. We revealed variations in the cellular composition of the epidermis, musculature organization, and spicules' arrangement, which most likely reflect the different functional roles of the spicule complex within Heterobranchia representatives. However, the obtained data are insufficient to answer the question of the origin of spicules and their homology within Heterobranchia.

石灰质骨骼结构常见于不同的无脊椎动物类群,包括多孔动物、蛇形纲、扁形动物、软体动物和棘皮动物。它们在外部表现为壳、板或鳞片,在内部则表现为表皮下的棘皮。虽然通常是在细胞外形成,但在腹足类软体动物中,异鳃纲发现了表皮下和细胞内的骨刺--至少在软体动物中这种情况很少见。这种类型的棘皮存在于几个系统发育距离较远的异鳃纲软体动物类群(Rhodopemorpha、Acochlidimorpha 和 Nudibranchia)中,每个类群在形态学和自生态学方面都各不相同。这就提出了关于异支软体动物表皮下棘皮的起源和同源性的问题。要回答这些问题,了解棘皮的形成过程至关重要。表皮下棘皮和邻近上皮的精细结构数据不足,阻碍了这一工作的开展。为了解决这个问题,我们对一个物种的表皮下棘皮和体壁进行了全面的形态学研究。我们发现了表皮细胞组成、肌肉组织和棘突排列的变化,这些变化很可能反映了棘突复合体在异鳃纲动物中的不同功能作用。然而,所获得的数据还不足以回答棘皮动物的起源及其在异鳃亚目中的同源性问题。
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引用次数: 0
Development, differentiation and toughness in lizard claws derive from two types of cysteine–glycine-rich proteins 蜥蜴爪子的发育、分化和韧性源自两种富含半胱氨酸-甘氨酸的蛋白质
IF 1.1 4区 生物学 Q4 ANATOMY & MORPHOLOGY Pub Date : 2024-08-07 DOI: 10.1111/azo.12516
Lorenzo Alibardi

The present study reports the formation of claws in lizards and their cornification. From a flat epidermis with lose mesenchyme, the skin forms undulations that become symmetric domes and later asymmetric for the elongation of their proximal part that gives rise to the outer scale surface. At the tip of digits claws develop in association with a mesenchyme that surrounds the cartilage of the last phalange. Beta-cells of claws appear 2–3 embryonic stages before those of scales formed in other body regions, including digital scales. The epidermis of the developing claw initially forms 3–4 embryonic layers under which elongated beta-cells accumulate long bundles of corneous material, immunopositive for CBPs. Claw beta-keratinocytes incorporate tritiated proline and accumulate numerous, oriented corneous bundles that rapidly occupy the entire cell. The bundles are electron-dense under transmission electron microscopy, suggesting that, in association to CBPs, other proteins are synthesized and increase the mechanical resistance of claws. Immunolabelling for cysteine-rich IFKs, HA and HB, shows that these keratins co-localize with CBPs only in claws but are low to absent in digital scales and adhesive setae. This indicates that the hardness of claws mainly depends from the association between these different proteins rich in glycine and cysteine.

本研究报告了蜥蜴爪的形成及其角化过程。皮肤从失去间质的扁平表皮开始形成起伏,这些起伏成为对称的圆顶,后来由于近端部分的伸长而形成不对称的圆顶,进而形成外鳞片表面。在趾甲的顶端,趾爪与围绕最后一节趾骨软骨的间质一起发育。爪的 Beta 细胞比其他身体区域(包括数字鳞片)形成的鳞片的 Beta 细胞早 2-3 个胚胎阶段出现。发育中的爪表皮最初会形成 3-4 个胚层,在这些胚层下,拉长的 beta 细胞会积聚长长的角质材料束,CBPs 免疫呈阳性。爪β角质细胞结合三尖脯氨酸,并积累大量定向角质束,这些角质束迅速占据整个细胞。在透射电子显微镜下,角质束呈电子致密状,这表明除了 CBPs 外,还合成了其他蛋白质,从而增强了爪的机械阻力。对富含半胱氨酸的 IFKs、HA 和 HB 的免疫标记表明,这些角蛋白只在爪中与 CBPs 共定位,而在数字鳞片和粘着刚毛中则很少或没有。这表明,爪的硬度主要取决于这些富含甘氨酸和半胱氨酸的不同蛋白质之间的关联。
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