We present systematic and detailed measurements of interparticle contact forces inside three-dimensional piles of frictionless liquid droplets. We measured long-range chainlike correlations of the directions and magnitudes of large forces, thereby establishing the presence of force chains in three dimensions. Our correlation definition provides a chain persistence length of 10 mean droplet diameters, decreasing as load is applied to the pile. We also measured the angles between contacts and showed that the chainlike arrangement arises from the balance of forces. Moreover, we found that piles whose height was comparable to the chain persistence length exhibited substantially greater strain hardening than did tall piles, which we attributed to the force chains. Together, the results establish a connection between the microscopic force network and the elastic response of meso- or macroscopic granular piles. The conclusions drawn here should be relevant in jammed systems generally, including concentrated emulsions and piles of sand or other heavy particles.
Spiders possess a wide array of sensory-driven behaviors and therefore provide rich models for studying evolutionary hypotheses about the relationship between brain morphology, sensory systems, and behavior. Despite this, only a handful of studies have examined brain variation across the order of Araneae. In this study, I present descriptions of the gross brain morphology for 19 families of spiders that vary in eye morphology. Spiders showed the most variation in the secondary eye visual pathway. Based on this variation, spiders could be categorized into four groups. Group 1 spiders had small, underdeveloped laminae, no medullae, and no mushroom bodies. Group 2 spiders had large laminae, no medullae and large mushroom bodies. Group 3 spiders had laminae and some evidence of reduced medullae and mushroom bodies. Group 4 spiders had the most complex systems, with large laminae, medullae formed from optical glomeruli, and robust mushroom bodies. Within groups, there was large variation in the shape and size of individual regions, indicating possible variation in neuronal organization. The possible evolutionary implications of the loss of a dedicated olfactory organ in spiders and its effects on the mushroom body are also discussed.
Whip spiders (Amblypygi) are known for their nocturnal navigational abilities, which rely on chemosensory and tactile cues and, to a lesser degree, on vision. Unlike true spiders, the first pair of legs in whip spiders is modified into extraordinarily long sensory organs (antenniform legs) covered with thousands of mechanosensory, olfactory, and gustatory sensilla. Olfactory neurons send their axons through the leg nerve into the corresponding neuromere of the central nervous system, where they terminate on a particularly large number (about 460) of primary olfactory glomeruli, suggesting an advanced sense of smell. From the primary glomeruli, olfactory projection neurons ascend to the brain and terminate in the mushroom body calyx on a set of secondary olfactory glomeruli, a feature that is not known from olfactory pathways of other animals. Another part of the calyx receives visual input from the secondary visual neuropil (the medulla). This calyx region is composed of much smaller glomeruli ("microglomeruli"). The bimodal input and the exceptional size of their mushroom bodies may support the navigational capabilities of whip spiders. In addition to input to the mushroom body, we describe other general anatomical features of the whip spiders' central nervous system.
Some species have sensory systems divided into subsystems with morphologically different sense organs that acquire different types of information within the same modality. Jumping spiders (family Salticidae) have eight eyes. Four eyes are directed anteriorly to view objects in front of the spider: a pair of principal eyes track targets with their movable retinae, while the immobile anterior lateral (AL) eyes have a larger field of view and lower resolution. To test whether the principal eyes, the AL eyes, or both together mediate the response to looming stimuli, we presented spiders with a video of a solid black circle that rapidly expanded (loomed) or contracted (receded). Control spiders and spiders with their principal eyes masked were significantly more likely to back away from the looming stimulus than were spiders with their AL eyes masked. Almost no individuals backed away from the receding stimulus. Our results show that the AL eyes alone mediate the loom response to objects anterior to the spider.
The diversity of spider behavior and sensory systems provides an excellent opportunity for comparative studies of the relationship between the brain and behavior. However, the morphology of spiders poses a challenge for histologists since the spider cephalothorax contains heterogeneous tissues and has both tough external and internal sclerotized regions. Unlike the heads of insects, the cephalothorax is highly pressurized, which can cause tissues to shift during processing and can reduce tissue cohesion in thick sections. This work describes a novel protocol for producing thick whole-head sections for morphological study by softening the exoskeleton and stabilizing friable tissue, without freezing or dehydration. It also presents an effective whole-head DiI staining method that uses minimal dehydration and highlights neural structures.
scite is a Brooklyn-based organization that helps researchers better discover and understand research articles through Smart Citations–citations that display the context of the citation and describe whether the article provides supporting or contrasting evidence. scite is used by students and researchers from around the world and is funded in part by the National Science Foundation and the National Institute on Drug Abuse of the National Institutes of Health.