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.
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.
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.
Lynne A. Fieber
This chapter introduces working definitions of neuropeptides and neurotransmitters from the perspective of invertebrate physiological processes. Neuropeptides and neurotransmitters are intercellular chemical signaling agents used by all animals. Chemical signaling augments or substitutes for electrical communication in the nervous system. When these agents act as neurotransmitters, they convert electrical signals to chemical signals across the synapse. As hormones, they circulate from a site of release to act at a more distant site in the body of the organism. Neuropeptides and neurotransmitters are classified into these groups mostly on the basis of their molecular size. This article describes several neuropeptide superfamilies and their wide scope of actions in model invertebrates. The article also describes the main neurotransmitters used by invertebrates.
Do insects, like other animals, expect future events, predict the value of potential actions, and decide between behavioral options without having access to the indicating stimuli? These cognitive capacities are captured by the term intentionality. This chapter addresses the question at two levels, behavior and neural correlates. Behavioral studies are performed with freely flying bees in the natural environment and with harnessed bees in the laboratory by applying the proboscis extension response paradigm. Data are presented and discussed on context-dependent learning, selective attention, rule learning, navigation, communication, and sleep-dependent memory consolidation. Although behavioral analyses document the rich repertoire and the cognitive dimensions of honeybee behavior, intentionality is nearly impossible to prove by behavioral analyses only and neural correlates are essential.
The Vertical Lobe of Cephalopods—A Brain Structure Ideal for Exploring the Mechanisms of Complex Forms of Learning and Memory
Ana Turchetti-Maia, Tal Shomrat, and Binyamin Hochner
We show that the cephalopod vertical lobe (VL) is a promising system for assessing the function and organization of the neuronal circuitry mediating complex learning and memory behavior. Studies in octopus and cuttlefish VL networks suggest an independent evolutionary convergence into a matrix organization of a divergence-convergence (“fan-out fan-in”) network with activity-dependent long-term plasticity mechanisms. These studies also show, however, that the properties of the neurons, neurotransmitters, neuromodulators, and mechanisms of induction and maintenance of long-term potentiation are different from those evolved in vertebrates and other invertebrates, and even highly variable among these two cephalopod species. This suggests that complex networks may have evolved independently multiple times and that, even though memory and learning networks share similar organization and cellular processes, there are many molecular ways of constructing them.