Faculty of Veterinary, Department of Anatomy, Animal Production and Clinical Veterinary Sciences, University of Santiago de Compostela, Lugo, Spain.
J Anat. 2018 Dec;233(6):814-827. doi: 10.1111/joa.12884. Epub 2018 Sep 25.
The characterization of the rabbit mammary pheromone, which is sensed by the main olfactory system, has made this species a unique model for the study of pheromonal communication in mammals. This discovery has brought attention to the global understanding of chemosensory communication in this species. Chemocommunication is mediated by two distinct organs located in the nasal cavity, the main olfactory epithelium and the vomeronasal organ (VNO). However, there is a lack of knowledge about the vomeronasal system in rabbits. To understand the role of this system, an exhaustive anatomical and histological study of the rabbit VNO was performed. The rabbit VNO was studied macroscopically by light microscopy, and by histochemical and immunohistochemical techniques. We employed specific histological staining techniques (periodic acid-Schiff, Alcian blue, Gallego's trichrome), confocal autofluorescence, histochemical labelling with the lectin Ulex europaeus agglutinin (UEA-I), and immunohistochemical studies of the expression of the Gαi2 and Gαo proteins and olfactory marker protein. The opening of the vomeronasal duct into the nasal cavity and its indirect communication with the oral cavity through a functional nasopalatine duct was demonstrated by classical dissection and microdissection. In a series of transverse histological sections, special attention was paid to the general distribution of the various soft-tissue components of this organ (duct, glands, connective tissue, blood vessels and nerves) and to the nature of the capsule of the organ. Among the main morphological features that distinguish the rabbit VNO, the presence of a double envelope, which is bony externally and cartilaginous internally, and highly developed venous sinuses stand out. This observation indicates the crucial role played in this species by the pumping mechanism that introduces chemical signals into the vomeronasal duct. The functional properties of the organ are also confirmed by the presence of a well-developed neuroepithelium and profuse glandular tissue that is positive for neutral mucopolysaccharides. The role of glycoconjugates was assessed by the identification of the α1-2 fucose glycan system in the neuroepithelium of the VNO employing UEA-I lectin. The pattern of labelling, which was concentrated around the commissures of the sensory epithelium and more diffuse in the central segments, is different from that found in most mammals studied. According to the expression of G-proteins, two pathways have been described in the VNOs of mammals: neuroreceptor cells expressing the Gαi2 protein (associated with vomeronasal receptor type 1); and cells expressing Gαo (associated with vomeronasal receptor type 2). The latter pathway is absent in most mammals studied. The expression of both G-protein families in the rabbit VNO places Lagomorpha together with rodents and insectivores in a small group of mammals belonging to the two-path model. These findings support the notion that the rabbit possesses a highly developed VNO, with many specific morphological features, which highlights the significance of chemocommunication in this species.
兔类气味信息素的特征是被主要嗅觉系统感知,这使得该物种成为研究哺乳动物气味信息素交流的独特模型。这一发现引起了人们对该物种化学感觉通讯的全球理解。化学通讯是由鼻腔中两个不同的器官介导的,即主嗅觉上皮和犁鼻器(VNO)。然而,关于兔类犁鼻器系统的知识还很缺乏。为了了解该系统的作用,我们对兔类 VNO 进行了全面的解剖学和组织学研究。我们通过光学显微镜、组织化学和免疫组织化学技术对兔类 VNO 进行了宏观研究。我们采用了特定的组织学染色技术(过碘酸-雪夫、阿利新蓝、加勒戈三色)、共聚焦自发荧光、凝集素 Ulex europaeus agglutinin(UEA-I)的组织化学标记以及 Gαi2 和 Gαo 蛋白和嗅觉标记蛋白表达的免疫组织化学研究。通过经典解剖和微解剖证明了犁鼻管开口进入鼻腔,并通过功能性鼻腭管与口腔间接相通。在一系列横切组织学切片中,特别注意到该器官(导管、腺体、结缔组织、血管和神经)的各种软组织成分的一般分布,以及器官包膜的性质。在区分兔类 VNO 的主要形态特征中,外部为骨、内部为软骨的双层包膜和高度发达的静脉窦尤为突出。这一观察表明,在该物种中,泵送机制在将化学信号引入犁鼻管中起着至关重要的作用。通过存在发达的神经上皮和富含中性粘多糖的丰富腺体组织,也证实了该器官的功能特性。通过识别 UEA-I 凝集素在 VNO 神经上皮中的α1-2 岩藻糖糖系统,评估了糖缀合物的作用。标记模式集中在感觉上皮的连接体周围,在中央部分更为扩散,与大多数研究的哺乳动物不同。根据 G 蛋白的表达,哺乳动物的 VNO 中已经描述了两种途径:表达 Gαi2 蛋白的神经受体细胞(与犁鼻受体 1 型相关);和表达 Gαo 的细胞(与犁鼻受体 2 型相关)。后者途径在大多数研究的哺乳动物中都不存在。兔类 VNO 中两种 G 蛋白家族的表达将兔形目动物与啮齿目动物和食虫目动物一起归入属于双途径模型的一小群哺乳动物中。这些发现支持了这样一种观点,即兔子拥有高度发达的 VNO,具有许多特定的形态特征,这突出了化学通讯在该物种中的重要性。