Yun Doo Chung and Jeongmi Lee
Hearing in invertebrates has evolved independently as an adaptation to avoid predators or to mediate intraspecific communication. Although many invertebrate groups are able to respond to sound stimuli, insects are the only group in which hearing is widely used. Therefore, we will focus here on the auditory systems of some well-known insect models. Appearance of the ability to perceive sound in insects is presumably a quite recent event in evolution. As a result of independent evolution, diverse types of hearing organs are evolved in insects. Here we will introduce basic features of insect ears and the mechanisms through which sound stimuli are converted into neuronal electric signals. We will also summarize our current understanding of neural processing of auditory information, including tonotopy, sound localization, and pattern recognition.
Reception of chemicals via olfaction and gustation are prerequisites to find, distinguish, and recognize food and mates and to avoid dangers. Several receptor gene superfamilies are employed in arthropod chemosensation: inverse 7-transmembrane (7-TM) gustatory and olfactory receptors (GRs, ORs), 3-TM ionotropic glutamate-related receptors (IRs), receptor-guanylyl cyclases, transient receptor potential ion channels, and epithelial sodium channels. Some of these receptor gene families have ancient origins and expanded in several taxa, producing very large, variant gene families adapted to the respectively relevant odor ligands in species-specific environments. Biochemical and electrophysiological studies in situ as well as molecular genetics found evidence for G-protein-dependent signal transduction cascades for ORs, GRs, and IRs, suggesting that signal amplification is paramount for chemical senses. In contrast, heterologous expression studies argued for primarily ionotropic transduction as a prerequisite to interstimulus intervals in the range of microseconds.
Important cnidarian contributions to our understanding of nervous system evolution may be found in the arrangement of conducting systems and their interactions. We see multiple, diffuse systems that interact to produce specific behaviors, the compression of conducting systems into compact directional or bidirectional conduction systems, and accumulation of multiple compressed conducting systems into integrating structures like nerve rings. We even see ganglion-like rhopalia that contain bilateral and directional conducting pathways. We now know that this compression and specificity of connections is controlled by conserved sets of genetic commands similar to those found in bilateral animals, and likely in common ancestors. This gradation in centralization is only limited in a directed pathway by the unique radial symmetry of cnidarians. Based on the compression of cnidarian conducting systems into integrating centers (nerve rings and rhopalia), the primary hurdle to cephalization is body symmetry. Medusoid cnidarians possess multiple “brains” connected by conducting systems that, by necessity, are nonpolarized.
Roy E. Ritzmann and Sasha N. Zill
This article discusses legged locomotion in insects. It describes the basic patterns of coordinated movement both within each leg and among the various legs. The nervous system controls these actions through groups of joint pattern generators coupled through interneurons and interjoint reflexes in a range of insect species. These local control systems within the thoracic ganglia rely on leg proprioceptors that monitor joint movement and cuticular strain interacting with central pattern generation interneurons. The local control systems can change quantitatively and qualitatively as needed to generate turns or more forceful movements. In dealing with substantial obstacles or changes in navigational movements, more profound changes are required. These rely on sensory information processed in the brain that projects to the multimodal sensorimotor neuropils collectively referred to as the central complex. The central complex affects descending commands that alter local control circuits to accomplish appropriate redirected movements.
This review focuses on the neural control of feeding in Aplysia. Its purpose is to highlight distinctive features of the behavior and to describe their neural basis. In a number of mollusks, food is grasped by a radula that protracts, retracts, and hyperretracts. In Aplysia, however, hyperretraction can require afferent activation. Phase-dependent regulation of sensorimotor transmission occurs in this context. Aplysia also open and close the radula, generating egestive as well as ingestive responses. Thus, the feeding network multitasks. It has a modular organization, and behaviors are constructed by combinations of behavior-specific and behavior-independent neurons. When feeding is initially triggered in Aplysia, responses are poorly defined. Motor activity is not properly configured unless responses are repeatedly induced and modulatory neurotransmitters are released from inputs to the central patter generator (CPG). Persistent effects of modulation have interesting consequences for task switching.
The main function of brains is to generate adaptive behavior. Far from being the stereotypical, robot-like insect, the fruit fly Drosophila exhibits astounding flexibility and chooses different courses of actions even under identical external circumstances. Due to the power of genetics, we now are beginning to understand the neuronal mechanisms underlying this behavioral flexibility. Interestingly, the evidence from studies of disparate behaviors converges on common organizational principles common to many if not all behaviors, such as modified sensory processing, involvement of biogenic amines in network remodeling, ongoing activity, and modulation by feedback. Seemingly foreseeing these recent insights, the first research fields in Drosophila behavioral neurogenetics reflected this constant negotiation between internal and external demands on the animal as the common mechanism underlying adaptive behavioral choice in Drosophila.
Jiaxing Li and Catherine A. Collins
In the face of acute or chronic axonal damage, neurons and their axons undergo a number of molecular, cellular, and morphological changes. These changes facilitate two types of responses, axonal degeneration and regeneration, both of which are remarkably conserved in both vertebrates and invertebrates. Invertebrate model organisms, including Drosophila and C. elegans, have offered a powerful platform with accessible genetic tools for manipulation and amenable nervous system for visualization. Thus far, several critical components and pathways in axonal degeneration and regeneration have been identified in invertebrate studies, including Sarm and Wallenda/DLK. This article highlights important findings in Drosophila, C. elegans, and other invertebrate injury models that have shed light upon the mechanism in axonal injury response.
Despite their often small numbers, the neurons in invertebrate nervous systems can nevertheless constitute many classes, and the nervous systems of little studied or entirely new species still offer significant opportunities for discovery. Circuit analyses and connectomic data are of particular significance, as are the relationships of these to behavior, and the organization of simple larval brains. Functional analyses of synaptic circuits still require knowledge of the neurotransmitter and neurotransmitter receptor for each identified neuron. Synapse complexity ranges widely; undifferentiated pathways in basal species may have unpolarized synapses with presynaptic sites opposite each other, and specialized pathways may have polyadic synapses.
Alex S. Mauss and Alexander Borst
Visual perception seems effortless to us, yet it is the product of elaborate signal processing in intricate brain circuits. Apart from vertebrates, arthropods represent another major animal group with sophisticated visual systems in which the underlying mechanisms can be studied. Arthropods feature identified neurons and other experimental advantages, facilitating an understanding of circuit function at the level of individual neurons and their synaptic interactions. Here, focusing on insect and crustacean species, we summarize and connect our current knowledge in four related areas of research: (1) elementary motion detection in early visual processing; (2) the detection of higher level visual features such as optic flow fields, small target motion and object distance; (3) the integration of such signals with other sensory modalities; and (4) state-dependent visual motion processing.
Guy Levy, Nir Nesher, Letizia Zullo, and Binyamin Hochner
Motor Control is essentially the computations required for producing coordinated sequences of commands from the controlling system (i.e., nervous system) to the actuation system (i.e., muscles) to generate efficient motion. The level of motor control complexity depends on the number of free parameters (degrees of freedom) that have to be coordinated. This number is much smaller in skeletal animals because they have a rather limited number of joints. In soft bodied animals, like the octopus, this number is virtually infinite. Here we show that the efficient motor control system of the octopus uses solutions that are very different from those of articulated animals, and it involves embodied co-evolution of the unique morphology together with the organization of the nervous and muscular systems to enable control strategies that are best suited for a highly active soft-bodied animal like the octopus.