- Synaptogenesis
The formation of the chemical synaptic junction that mediates communication between neurons and other neurons or muscle cells. These junctions can be identified in electron micrographs as darkened specialized areas on the presynaptic side of the junction that contains clusters of synaptic vesicles.
- Gap junction biogenesis and organization
Gap junctions, along with chemical synapses, mediate communication between neurons. Innexins are major components of these junctions. Gap junctions can be identified in electron micrographs as parallel membranes of closely apposed cells.
- Olfaction
Volatile organic molecules are sensed through olfaction. C. elegans can distinguish and respond to many volatile odorants through attractive or repulsive chemotactic behaviors. In some instances volatile compounds can induce both behaviors depending on its concentration. Olfaction studies in C. elegans has revealed a complex sensory system where only three types of neurons (AWC, AWA, and AWB) have been found to be responsible for processing over seven classes of volatile odorants, including alcohols, ketones, organic acids, sulfhydrals, and heterocyclic compounds. Detailed study of the molecular machinery behind odor reception has shown that each neuron controls a particular attractive or repulsive behavioral response, for example, AWC controls attractive chemotaxis responses and AWB controls repulsive chemotaxis responses. One distinguishing feature of C. elegans sensory system is that the sensory neurons are polymodal in their stimulus detecting ablility; that is, individual neurons in C. elegans express multiple odorant receptors allowing multiple sensory functions, whereas vertebrate neurons express a single receptor limiting their function to detecting a single odorant.
- Neurotransmission
Neurons communicate across synaptic junctions with target cells, such as neurons, muscles, or specialized secretory cells through chemical messengers that are released from the neuron and bind to and activate receptors on the target cell. Pre-synaptic release of neurotransmitters can be evoked, such as through mechanical or chemical stimulation, as well as can occur spontaneously at a low rate. Depending on the neurotransmitter released and or the receptors of the post-synaptic cell, the activation of receptors can trigger excitatory or inhibitory actions in the target cell. These neuronal communications can also result in short term post-synapatic cellular changes to the membrane potential or can cause the activation of signaling cascades, resulting in longer term changes in the cell.
- Mechanosensation
Mechanosensation converts mechanical energy into electrical signals allowing an organism to use physical cues from the environment or from internal sensors to affect its behavior. Mechanical stimuli are received through mechanosensory receptor neurons (MRNs). In C. elegans, there are 30 putative MRNs in hermaphrodites while an additional 52 MRNs are found in males. More than 40 of these male-specific MRNs are found in the male tail, hook, post-cloacal sensilla and spicule and are required for male mating. MRNs transmit electrical signals to other neurons through electrical or chemical synapses. MRNs may or may not have ciliated dendrite endings, which is some cases are exposed to the outside. Mechanical stimuli initiate as well as modulate many behaviors of the worm. MRNs allow the worm to respond to light touch, such as stroking with an eyelash as well as harsh touch, such as prodding with a pick.
- Cell migration
Cell movement is an essential cell behavior for metazoan development. When this process is improperly orchestrated it can result in developmental disorders or pathologies such as tumor metastasis. In C. elegans, many cell types including canal associated neurons (CANs), hermaphrodite-specific neurons (HSNs), and Q neuroblasts migrate long distances during embryonic or larval development. Studies in C. elegans have elucidated many of the molecules required for stimulating and guiding the cell. These studies have shown that some directed movement rely on graded chemotactic signaling that is perceived by the cell and transduced to the cell's cytoskeleton. Chemotactic signaling molecules such as UNC-6, an extracellular matrix protein, can act as both an attractant, for cells expressing UNC-5, or as a repellant, for cells expressing UNC-40. Cell migration ultimately requires the regulation of cytoskeletal rearrangements. Studies have demonstrated UNC-73/Trio to be a main activator of Rac signaling in at least some of these migrating cells, which is proposed to drive such intracellular changes.
- Egg laying
C. elegans hermaphrodites exhibit a periodicity in the rate and temporal pattern of egg-laying. Egg laying is modulated by diverse environmental cues. Egg laying behavior has served as an important phenotypic assay for the genetic dissection of neuronal signal transduction mechanisms. Studies in C. elegans have elucidated the roles of specific neurons in the egg-laying motor circuit, which release multiple neurotransmitters affecting distinct parameters of egg-laying muscle activity, and the possible mechanisms for sensory control of egg-laying behavior.
- Foraging
C. elegans exhibits a number of behaviors that influence its ability to find food. The act of foraging proceeds as an alteration of two behavioral states, roaming and dwelling. These states depend on sensory input, as mutations that affect cilia formation alter the time spent in one state or the other. Nose deflection is another characteristic foraging behavior. In this case, the rapid movement of the nose tip from side to side has been shown to require GABAergic RME neurons. Studies of C. elegans isolates also demonstrate a social aspect to foraging, which is due to natural variations in
npr-1, an NPY G-protein-coupled receptor family.
- Muscular system development and organization
The coordinated specification and functional assemblage of cells and tissues into the contractile organ system in the animal. C. elegans muscles are of two types: single sarcomere with focal attachment points at the ends (alimentary system and sex muscles) and obliquely striated muscles with many sarcomeres and no one substantial focal attachment point (body-wall muscles). Components of C. elegans muscles are similar to other animals and include heavy and light-chain myosin, actin, tropomyosin, troponin-like proteins, and paramyosin. Unlike other muscle systems, C. elegans muscles send neuron-like processes to neuropils that contain motor neuron axons rather than motor neurons sending axons to innervate the muscle. Contractile tissue is found throughout C. elegans and is required for locomotion (body wall muscle), eating (pharyngeal muscle), egg laying (vulval and uterine muscles, and gonad sheath), male mating (male tail muscles), and defecation (enteric muscles).
- Defecation
In C. elegans the expulsion of intestinal contents occurs every 45-50 seconds. This cycle is characterized by a pattern of muscle contractions under both muscle and neuronal control. The steps of the defecation cycle are a posterior body contraction (pBoc), an anterior body contraction (aBoc), and the final expulsion step (Exp) where the enteric muscles contract, opening the anus and allowing the intestinal contents to be released. Each step is independently controlled as mutations exist that affect one step but do not alter the timing or occurrence of the other. Further, Ca++ oscillations in the intestine, rather than neuronal stimulation, have been shown to control the initiating pBoc step. The contractions of the enteric muscles are controlled by GABA motor neurons AVL and DVB through an excitatory GABA-gated cation channel. The periodicity of the cycle is influenced by the presence of food, is temperature compensated, and can be reset by mechanosensory input.