Bock Natika L, Paul Dorothy H
Department of Biology, University of Victoria, Victoria, BC, Canada.
Brain Behav Evol. 2009;73(3):188-205. doi: 10.1159/000222563. Epub 2009 Jun 3.
Squat lobsters (Galatheidae) and mole sand crabs (Hippidae) differ in posture and locomotion from each other and from crayfish, their surrogate ancestor for neurobehavioral features. Galatheids resemble crayfish more closely in general behavior and niche, but are intermediate between crayfish and hippids with respect to morphology and neuromusculature. The tailfan is inverted under the abdomen in both, due to the flexed abdominal posture, but its morphology has diverged considerably. Nothing is known about adaptations of the tailfan exteroceptors to the new sensory world of either group. We used SEM, vital staining, and extracellular electrophysiological techniques to survey the sensory structures on the telsons of the galatheid Munida quadrispina and the hippid Emerita analoga for comparison with published data on the homologous mechanosensory system in crayfish. Both telsons bear plumose, peg, and non-annulate (natatory or guard) setae. In addition, M. quadrispina has singly-innervated smooth setae and E. analoga a previously undescribed type of small seta the outer face of which is covered by transversely-oriented, thin setules that are much broader than they are long and angled outward toward the seta's distal end, overlapping loosely. The 'stack-of-scales' appearance of its distal portion viewed from the side engendered the name: scaly seta. Some shared features with other small setae that are chemo- and mechanoreceptive suggest scaly setae might be bimodal sensilla. The telson of M. quadrispina is very flexible. Plumose setae on its dorsal surface are arranged into hemi-circlets and most, if not all, appear not to be innervated. They may contribute to adjacent smooth setae's mechanosensitivity via mechanical coupling through adjacent cuticle, as occurs between feathered and smooth setae on crayfish antennae. Sensory nerve recordings show many afferents to have low thresholds to mechanical disturbance, suggesting they are hydrodynamic receptors. The telson of E. analoga is rigid, and all dorsal setae are relegated to the margins. Patches of scaly setae on the anterior lateral dorsal telson are strategically located to sense the substrate when the crabs are in sand. Scaly and peg setae are arrayed along shallow grooves, one along each side, that are flanked laterally by a fringe of plumose and pappose setae. Substantial deflection from resting position of the latter was required to reliably elicit afferent activity, suggesting most function as touch receptors. The different, non-random distributions of tailfan setae match these animals' divergent sensory worlds and might have engendered species-specific alterations in their central sensory systems.
铠甲虾(铠甲虾科)和鼹鼠沙蟹(蝉蟹科)在姿势和运动方式上彼此不同,也与小龙虾不同,小龙虾是它们神经行为特征的替代祖先。一般来说,铠甲虾在行为和生态位上与小龙虾更为相似,但在形态和神经肌肉方面介于小龙虾和蝉蟹之间。由于腹部弯曲的姿势,两者的尾扇都倒置在腹部下方,但其形态已经有了很大的差异。对于尾扇外感受器如何适应这两类动物新的感觉世界,我们一无所知。我们使用扫描电子显微镜、活体染色和细胞外电生理技术,对铠甲虾四刺芒眼虾和蝉蟹艾氏蝉蟹尾节上的感觉结构进行了研究,以便与已发表的关于小龙虾同源机械感觉系统的数据进行比较。两者的尾节都有羽状、栓状和无环(游泳或防护)刚毛。此外,四刺芒眼虾有单根神经支配的光滑刚毛,而艾氏蝉蟹有一种以前未描述过的小型刚毛,其外表面覆盖着横向排列的细毛,这些细毛比它们的长度宽得多,并且朝着刚毛的远端向外倾斜,松散地重叠。从侧面看,其远端部分呈现出“鳞片堆叠”的外观,因此得名:鳞片刚毛。与其他具有化学感受和机械感受功能的小型刚毛有一些共同特征,这表明鳞片刚毛可能是双模态感受器。四刺芒眼虾的尾节非常灵活。其背表面的羽状刚毛排列成半圆形,而且大多数(如果不是全部的话)似乎没有神经支配。它们可能通过相邻角质层的机械耦合,对相邻光滑刚毛的机械敏感性做出贡献,就像小龙虾触角上的羽状刚毛和平滑刚毛之间的情况一样。感觉神经记录显示,许多传入神经对机械干扰的阈值较低,这表明它们是流体动力感受器。艾氏蝉蟹的尾节是刚性的,所有背侧刚毛都分布在边缘。在尾节前外侧背表面的鳞片刚毛斑块位置经过精心安排,以便在螃蟹处于沙中时感知底物。鳞片刚毛和栓状刚毛沿着浅沟排列,每条浅沟位于一侧,其两侧有一排羽状和毛状刚毛。后者从静止位置发生相当大的偏转才能可靠地引发传入神经活动,这表明它们大多起到触觉感受器的作用。尾扇刚毛不同的、非随机分布与这些动物不同的感觉世界相匹配,可能导致了它们中枢感觉系统中物种特异性的变化。