Klingenberg C P, Badyaev A V, Sowry S M, Beckwith N J
Laboratory of Development and Evolution, University Museum of Zoology, Department of Zoology, Downing Street, Cambridge CB2 3EJ, United Kingdom.
Am Nat. 2001 Jan;157(1):11-23. doi: 10.1086/317002.
Organisms are built from distinct modules, which are internally coherent but flexible in their relationships among one another. We examined morphological variation within and between two candidate modules: the fore- and hindwings of bumblebees (Hymenoptera: Apidae: Bombus empatiens). We used the techniques of geometric morphometrics (Procrustes superimposition) to analyze the variation of landmark configurations in fore- and hindwings. Regression was used to correct for size-related shape variation (allometry). Principal component analysis revealed patterns of variation that were remarkably similar for individual variation and fluctuating asymmetry (FA). Because covariation of FA among parts must be due to direct transmission of the developmental perturbations causing FA, this agreement of patterns suggests that much of individual variation is also due to direct developmental interactions within each developing wing. Moreover, partial least squares analysis indicated that the patterns of shape covariation between fore- and hindwings were nearly the same as the patterns of within-wing variation. Shape covariation of FA was only found in bees that had been reared under elevated CO(2) concentration but not in bees from the control treatment, suggesting that the mechanisms of developmental interactions between fore- and hindwings are related to gas exchange. We conclude that the fore- and hindwings are developmental modules that maintain internal coherence through direct developmental interactions and are connected to each other only by relatively few links that use the system of interactions within modules.
生物体由不同的模块组成，这些模块在内部是连贯的，但它们之间的关系具有灵活性。我们研究了两个候选模块内部和之间的形态变异：熊蜂（膜翅目：蜜蜂科：耐氏熊蜂）的前翅和后翅。我们使用几何形态测量学技术（普氏叠加）来分析前翅和后翅地标配置的变异。回归用于校正与大小相关的形状变异（异速生长）。主成分分析揭示了个体变异和波动不对称（FA）的变异模式非常相似。由于各部分之间FA的协变一定是由于导致FA的发育扰动的直接传递，这种模式的一致性表明，许多个体变异也归因于每个发育中的翅膀内的直接发育相互作用。此外，偏最小二乘分析表明，前翅和后翅之间的形状协变模式与翅内变异模式几乎相同。FA的形状协变仅在高二氧化碳浓度下饲养的蜜蜂中发现，而在对照处理的蜜蜂中未发现，这表明前翅和后翅之间的发育相互作用机制与气体交换有关。我们得出结论，前翅和后翅是发育模块，它们通过直接发育相互作用保持内部连贯性，并且仅通过相对较少的利用模块内相互作用系统的联系相互连接。
