Cerio Donald G, Witmer Lawrence M
Department of Biological Sciences, Ohio University, Athens, OH, USA.
Department of Biomedical Sciences, Heritage College of Osteopathic Medicine, Ohio University, Athens, OH, USA.
PeerJ. 2019 Jul 23;7:e7355. doi: 10.7717/peerj.7355. eCollection 2019.
The cochlea and semicircular canals (SCCs) of the inner ear are vital neurosensory devices. There are associations between the anatomy of these sensorineural structures, their function, and the function of related biological systems, for example, hearing ability, gaze stabilization, locomotor agility, and posture. The endosseous labyrinth is frequently used as a proxy to infer the performance of the hearing and vestibular systems, locomotor abilities, and ecology of extinct species. Such fossil inferences are often based on single specimens or even a single ear, representing an entire species. To address whether a single ear is representative of a population, we used geometric morphometrics to quantitatively assess the variation in shape and symmetry in a sample of endosseous labyrinths of wild turkeys of southern Ohio. We predicted that ears would be symmetrical both within individuals and across the sample; that labyrinth shape and size would covary; that labyrinth shape would vary with the size of the brain, measured as width of the endocranium at the cerebellum; and that labyrinths would be morphologically integrated. To test these predictions, we microCT-scanned the heads of 26 cadaveric turkeys, digitally segmented their endosseous labyrinths in Avizo, and assigned 15 manual landmarks and 20 sliding semilandmarks to each digital model. Following Procrustes alignment, we conducted an analysis of bilateral symmetry, a Procrustes regression analysis for allometry and other covariates including side and replicate, and analyses of global integration and modularity. Based on Procrustes distances, no individual's left and right ears were clearly different from each other. When comparing the ears of different specimens, statistically clear differences in shape were found in only 66 of more than 1,300 contrasts. Moreover, effects of both directional and fluctuating asymmetry were very small-generally, two orders of magnitude smaller than the variance explained by individual variation. Statistical tests disagreed on whether these asymmetric effects crossed the threshold of significance, possibly due to non-isotropic variation among landmarks. Regardless, labyrinths appeared to primarily vary in shape symmetrically. Neither labyrinth size nor endocranial width was correlated with labyrinth shape, contrary to our expectations. Finally, labyrinths were found to be moderately integrated in a global sense, but four weakly separated modules-the three SCCs and cochlea-were recovered using a maximum-likelihood analysis. The results show that both fluctuating and directional asymmetry play a larger role in shape variation than expected-but nonetheless, endosseous labyrinths are symmetrical within individuals and at the level of the population, and their shape varies symmetrically. Thus, inferences about populations, and very possibly species, may be confidently made when only a single specimen, or even a single ear, is available for study.
内耳的耳蜗和半规管是重要的神经感觉器官。这些感觉神经结构的解剖结构、其功能以及相关生物系统的功能之间存在关联,例如听力、注视稳定、运动敏捷性和姿势。骨迷路经常被用作推断已灭绝物种的听觉和前庭系统性能、运动能力及生态的替代指标。此类化石推断通常基于单个标本甚至单个耳朵,却代表了整个物种。为了探究单个耳朵是否能代表一个种群,我们使用几何形态测量学定量评估了俄亥俄州南部野生火鸡骨迷路样本的形状和对称性变化。我们预测,耳朵在个体内部以及整个样本中都是对称的;骨迷路的形状和大小会共同变化;骨迷路的形状会随大脑大小而变化,以小脑处颅内膜的宽度来衡量;并且骨迷路在形态上是整合的。为了检验这些预测,我们对26只火鸡尸体的头部进行了显微CT扫描,在Avizo软件中对其骨迷路进行数字分割,并为每个数字模型指定了15个手动地标点和20个滑动半地标点。经过普氏叠加对齐后,我们进行了双侧对称性分析、关于异速生长和其他协变量(包括侧别和重复样本)的普氏回归分析,以及整体整合和模块性分析。基于普氏距离,没有个体的左耳和右耳明显不同。在比较不同标本的耳朵时,在1300多个对比中,只有66个在形状上存在统计学上的明显差异。此外,方向性不对称和波动不对称的影响都非常小——一般来说,比个体变异所解释的方差小两个数量级。关于这些不对称效应是否超过显著性阈值的统计检验结果不一致,可能是由于地标点之间存在非各向同性变异。无论如何,骨迷路在形状上似乎主要是对称变化的。与我们的预期相反,骨迷路大小和颅内膜宽度均与骨迷路形状无关。最后,从整体上看,骨迷路被发现有适度的整合,但通过最大似然分析发现了四个弱分离的模块——三个半规管和耳蜗。结果表明,波动不对称和方向性不对称在形状变异中所起的作用比预期的更大——但尽管如此,骨迷路在个体内部和种群水平上是对称的,并且其形状对称变化。因此,当只有单个标本甚至单个耳朵可供研究时,关于种群乃至很可能关于物种的推断仍可自信地做出。