Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada.
Faculty of Pharmacy & Pharmaceutical Sciences and Department of Biomedical Engineering, University of Alberta, Edmonton, AB, Canada.
J Anat. 2020 Apr;236(4):668-687. doi: 10.1111/joa.13132. Epub 2020 Jan 5.
The development of the iliosacral joint (ISJ) in tetrapods represented a crucial step in the evolution of terrestrial locomotion. This structure is responsible for transferring forces between the vertebral column and appendicular skeleton, thus supporting the bodyweight on land. However, most research dealing with the water-to-land transition and biomechanical studies in general has focused exclusively on the articulation between the pelvic girdle and femur. Our knowledge about the contact between the pelvic girdle and vertebral column (i.e. the ISJ) at a tissue level is restricted so far to human anatomy, with little to no information available on other tetrapods. This lack of data limits our understanding of the development and evolution of such a key structure, and thus on the pattern and processes of the evolution of terrestrial locomotion. Therefore, we investigated the macro- and microanatomy of the ISJ in limb-bearing squamates that, similar to most non-mammalian, non-avian tetrapods, possess only two sacral ribs articulating with the posterior process of the ilium. Using a combination of osteology, micro-computed tomography and histology, we collected data on the ISJ apparatus of numerous specimens, sampling different taxa and different ontogenetic stages. Osteologically, we recorded consistent variability in all three processes of the ilium (preacetabular, supracetabular and posterior) and sacral ribs that correlate with posture and locomotion. The presence of a cavity between the ilium and sacral ribs, abundant articular cartilage and fibrocartilage, and a surrounding membrane of dense fibrous connective tissue allowed us to define this contact as a synovial joint. By comparison, the two sacral ribs are connected to each other mostly by dense fibrous tissue, with some cartilage found more distally along the margins of the two ribs, defining this joint as a combination of a syndesmosis and synchondrosis. Considering the intermediary position of the ISJ between the axial and appendicular skeletons, the shape of the articular surfaces of the sacral ribs and ilium, and the characteristics of the muscles associated with this structure, we argue that the mobility of the ISJ is primarily driven by the movements of the hindlimb during locomotion. We hypothesize that limited torsion of the ilium at the ISJ happens when the hip is abducted, and the joint is likely able to absorb the compressional and extensional forces related to the protraction and retraction of the femur. The mix of fibres and cartilage between the two sacral ribs instead serves primarily as a shock absorber, with the potential for limited vertical translation during locomotion.
四足动物的荐髂关节 (ISJ) 的发展是陆地运动进化的关键一步。该结构负责在脊柱和附肢骨骼之间传递力,从而支撑身体在陆地上的重量。然而,大多数涉及水到陆的过渡和一般生物力学研究的研究都只专注于骨盆带和股骨之间的关节。我们对骨盆带和脊柱(即 ISJ)之间在组织水平上的接触的了解目前仅限于人体解剖学,而关于其他四足动物的信息则很少。这种数据的缺乏限制了我们对这种关键结构的发育和进化的理解,以及对陆地运动进化的模式和过程的理解。因此,我们研究了有肢鳞龙类动物的 ISJ 的宏观和微观解剖结构,这些动物与大多数非哺乳动物、非鸟类四足动物一样,只有两根荐骨与髂骨的后突关节。我们结合骨骼学、微计算机断层扫描和组织学,收集了大量样本的 ISJ 设备数据,样本来自不同的分类群和不同的个体发育阶段。骨骼学上,我们记录了髂骨的三个过程(髋臼前、髋臼上和后)和荐骨的一致性变化,这些变化与姿势和运动有关。髂骨和荐骨之间存在一个腔,丰富的关节软骨和纤维软骨,以及致密的纤维结缔组织膜,使我们能够将这种接触定义为滑膜关节。相比之下,两根荐骨主要通过致密的纤维组织相互连接,在两根肋骨的边缘处有一些软骨,将这个关节定义为一个联合和一个骺软骨的组合。考虑到 ISJ 在轴状骨骼和附肢骨骼之间的中间位置、荐骨和髂骨的关节表面的形状,以及与该结构相关的肌肉的特征,我们认为 ISJ 的活动性主要是由运动时后肢的运动驱动的。我们假设,当髋关节外展时,ISJ 处的髂骨旋转受到限制,关节可能能够吸收与股骨的前伸和后缩相关的压缩和拉伸力。两根荐骨之间的纤维和软骨混合物则主要作为减震器,在运动时可能会有有限的垂直平移。