Tzetlin Alexander, Vortsepneva Elena, Zhadan Anna
Biological Faculty, M. V. Lomonosov Moscow State University, Moscow, Russia.
J Morphol. 2023 Apr;284(4):e21568. doi: 10.1002/jmor.21568.
Representatives of the extant family Oenonidae (Annelida, Eunicida) have a prionognath jaw apparatus, with maxillae having forceps-like elements, a number of asymmetrical dentate plates and long slender carriers, which is characteristic of some fossil forms known from the Paleozoic epoch. Therefore, data on the fine structure and functional morphology of Oenonidae jaws are helpful for the interpretation of fossil materials. The fine structure of the jaw apparatus and the ventral pharyngeal organ is studied in one species of the Oenonidae (Annelida)-Drilonereis cf. filum. The material was collected in the soft bottom of Marseille Bay (Mediterranean) and examined with the help of TEM and histological techniques. A three-dimensional (3D) reconstruction was made from a complete series of semithin sections. The entire jaw apparatus is about 500 µm in length; it includes ventral mandibles and four pairs of maxillae, connected with long paired dorsal carriers and an unpaired ventral carrier. While retracted, it reaches the VIII-XI chaetigers. The most solid part of the maxillary apparatus, that is, maxillae I and II, are 2.5-5 µm thick. The plate consists of a monolithic array of merged scleroprotein granules in which perforations, that is, spaces remaining from microvilli, are visible; the basal part of the maxillary plate is a layer of loosely arranged collagen fibers penetrated with microvilli and has no signs of sclerotization. A study of the jaws of Drilonereis cf. filum showed the presence of common jaw patterns in Eunicida order. Like the jaws of Dorvilleidae, Eunicidae, Onuphidae, and Lumbrineridae, the jaws of Drilonereis are formed at the basis of a typical annelid cuticle's transformation with epi- and basicuticular layers, and its impregnation by merging scleroprotein granules. Through the nature of sclerotization, the jaws of D. cf. filum are similar to those of Dorvilleidae, Histriobdellidae, and the juvenile jaws of Mooreonuphis stigmatis (Onuphidae). Analysis of the 3D-reconstructions of the D. cf. filum jaw apparatus shows that the MxI of this species, and probably of other Oenonidae with dorsal and ventral carriers, can make grasping motions by fixing the joint of the right and left MxI in the two-door hinge type. In general, the overall structure of the jaw apparatus of D. cf. filum and the mechanics of its work shows greater similarity with that of Dorvilleidae than with the jaw apparatus of extant Labidognatha and Simmetrognatha (Onuphidae, Eunicidae, Lumbrineridae). The need for compactization of the jaw apparatus when moving in dense sediment or in the burrows is probably one of the factors determining its structure.
现存的欧诺虫科(环节动物门,矶沙蚕目)的代表具有一种锯颚咀嚼器,其上颚具有钳状结构、一些不对称的齿状板和细长的载体,这是一些已知的古生代化石形态的特征。因此,关于欧诺虫科颚的精细结构和功能形态的数据有助于解释化石材料。本文研究了欧诺虫科(环节动物门)的一个物种——似丝海蚯蚓(Drilonereis cf. filum)颚器和腹侧咽器官的精细结构。材料采集自马赛湾(地中海)的软底,并借助透射电子显微镜(TEM)和组织学技术进行检查。通过一系列完整的半薄切片进行了三维(3D)重建。整个颚器长度约为500微米;它包括腹侧下颌和四对上颚,与长的成对背侧载体和一个不成对的腹侧载体相连。收缩时,它到达第VIII - XI刚节。上颚器最坚实的部分,即第一和第二上颚,厚度为2.5 - 5微米。该板由融合的硬蛋白颗粒的整体阵列组成,其中可见穿孔,即微绒毛留下的空间;上颚板的基部是一层松散排列的胶原纤维,有微绒毛穿透,且没有硬化迹象。对似丝海蚯蚓颚的研究表明,矶沙蚕目存在共同的颚模式。与多维尔虫科、矶沙蚕科、蠕纹沙蚕科和蛰龙介科的颚一样,似丝海蚯蚓的颚是在典型环节动物角质层的表皮层和基底层转变的基础上形成的,并通过融合硬蛋白颗粒进行浸渍。通过硬化的性质,似丝海蚯蚓的颚与多维尔虫科、组织蛭科以及摩尔蠕纹沙蚕(蠕纹沙蚕科)的幼体颚相似。对似丝海蚯蚓颚器的3D重建分析表明,该物种以及可能其他具有背侧和腹侧载体的欧诺虫科的第一上颚,可能通过将左右第一上颚的关节固定在双门铰链类型来进行抓握动作。总体而言,似丝海蚯蚓颚器的整体结构及其工作机制与多维尔虫科的更为相似,而与现存的颚齿目和对称颚目(蠕纹沙蚕科、矶沙蚕科、蛰龙介科)颚器的相似性较低。在密集沉积物或洞穴中移动时对上颚器紧凑化的需求可能是决定其结构的因素之一。