Hair plates are a type of proprioceptor found in the folds of insect joints. [1] They consist of a cluster of hairs, in which each hair is innervated by a single mechanosensory neuron. Functionally, hair plates operate as "limit-detectors" by signaling the extremes of joint movement, [2] which then drives reflexive leg movement. [3]
Hair plates consist of a field of cuticular hairs, in which each hair is innervated by a single mechanosensory neuron [1] [4] (Figure 1). Hair plates are positioned within folds of cuticle at joints, and the associated hairs are deflected during joint movement. [5] The number of hairs across and within hair plates can vary [6] [7] and hair plates are found on different body parts, including the legs, [8] [9] [10] [6] [11] neck, [12] [13] and antennae. [14] [15] On the legs of insects, hair plates are found at the proximal joints (i.e. thorax-coxa, coxa-trochanter, and trochanter-femur joints) across the front, middle, and hind legs.
Hair plate neurons project into the ventral nerve cord where the arborize dorsally and around the leg neuromere. [16] Moreover, the neurons of the hair plates located at the thorax-coxa joint project through the ventral, dorsal, and accessory prothoracic nerves, whereas, other hair plate neurons on the leg project through the prothoracic leg nerve. [17] [18] Lastly, the neurons from each hair plate may have distinct axonal morphologies.
Hair-plates are proprioceptors [1] and provide information to the nervous system about the positional and movement state of the body and legs. Hair plate neurons come in two flavors, one that responds phasically (rapidly adapting) and another that responds tonically (slowly adapting) to transient or maintained deflections of the hairs. [11] [19] These encoding properties enable hair plates to signal the position and movement of adjoining body or leg segments. Neurons associated with the longer hairs within a hair plate form direct, mono-synaptic excitatory chemical synapses with motor neurons [6] as well as synapses with non-spiking interneurons that provide inhibitory input onto antagonistic motor neurons. [20] Therefore, hair plates provide rapid feedback to control the movement direction of the leg they are located on. It remains unknown what information the smaller hairs of hair plates signal and what that information is used for in leg motor control. Hair plate neurons are also involved in the presynaptic inhibition to other proprioceptors. [21]
Hair plates located at the leg joints provide sensory feedback for the control of walking. [8] [9] [22] [2] [23] [24] [25] In stick insects and cockroaches, the surgical removal of coxa hair plates alter the extremes of leg movement in such a way that the leg may overstep and collide with an ipsilateral leg. Therefore, hair plates control the transition of leg movement direction as well as the extent to which legs travel during the step cycle. [8] [22] This “limit-detector” function is similar to that of mammalian joint receptors. [26] Therefore, hair plates encode the extremes of joint movement during walking to precisely control the direction of leg movement.
Mechanosensory information from front leg hair plates also contribute to the regulation of feeding behavior in fruit flies, Drosophila melanogaster [27] . Integration of hair plate mechanosensory information with olfactory information from antennal neurons control the proboscis extension reflex (PER) in flies. Thus, the sensory input from hair plates is integrated with the information from other sensory modalities to control behaviors beyond walking.
Hair plates located on the neck (known as the prosternal organ) monitor head position relative to the thorax and provide sensory feedback for the control of head posture. [12] [13] In the blowfly Calliphora , surgical removal of the prosternal organ hairs on one side causes the fly to compensate by rolling the head toward the operated side. [12] These results suggest that the prosternal organ may be involved in gaze stabilization.
Hair plates on the legs have also been shown to be important for resting posture. The ablation of the anterior trochanteral hair plate on a stick insect leg did not alter the coordination between legs, but rather resulted in that leg being held higher. [28] Also, hair plates on the trochanter were shown to control the body height. Overall, in addition to controlling walking kinematics, hair plates are also involved in postural control.
Hair plates located on the proximal segments of the antenna (Figure 2) provide sensory feedback for the control of antennal movement [15] and are thought to play an important role in active sensing, object localization, and targeted reaching movements [14] [29] .
Halteres are a pair of small club-shaped organs on the body of two orders of flying insects that provide information about body rotations during flight. Insects of the large order Diptera (flies) have halteres which evolved from a pair of ancestral hindwings, while males of the much smaller order Strepsiptera (stylops) have halteres which evolved from a pair of ancestral forewings.
The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial cells, known as hair cells, which respond to displacement caused by motion and transduce these signals into electrical impulses via excitatory synapses. Lateral lines play an important role in schooling behavior, predation, and orientation.
A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are located on sensory neurons that convert mechanical pressure into electrical signals that, in animals, are sent to the central nervous system.
Sensory neurons, also known as afferent neurons, are neurons in the nervous system, that convert a specific type of stimulus, via their receptors, into action potentials or graded receptor potentials. This process is called sensory transduction. The cell bodies of the sensory neurons are located in the dorsal root ganglia of the spinal cord.
Range fractionation is a term used in biology to describe the way by which a group of sensory neurons are able to encode varying magnitudes of a stimulus. Sense organs are usually composed of many sensory receptors measuring the same property. These sensory receptors show a limited degree of precision due to an upper limit in firing rate. If the receptors are endowed with distinct transfer functions in such a way that the points of highest sensitivity are scattered along the axis of the quality being measured, the precision of the sense organ as a whole can be increased.
Johnston's organ is a collection of sensory cells found in the pedicel of the antennae in the class Insecta. Johnston's organ detects motion in the flagellum. It consists of scolopidia arrayed in a bowl shape, each of which contains a mechanosensory chordotonal neuron. The number of scolopidia varies between species. In homopterans, the Johnston's organs contain 25–79 scolopidia. The presence of Johnston's organ is a defining characteristic which separates the class Insecta from the other hexapods belonging to the group Entognatha. Johnston's organ was named after the physician Christopher Johnston (1822-1891) father of the physician and Assyriologist Christopher Johnston.
Escape response, escape reaction, or escape behavior is a mechanism by which animals avoid potential predation. It consists of a rapid sequence of movements, or lack of movement, that position the animal in such a way that allows it to hide, freeze, or flee from the supposed predator. Often, an animal's escape response is representative of an instinctual defensive mechanism, though there is evidence that these escape responses may be learned or influenced by experience.
Campaniform sensilla are a class of mechanoreceptors found in insects, which respond to local stress and strain within the animal's cuticle. Campaniform sensilla function as proprioceptors that detect mechanical load as resistance to muscle contraction, similar to mammalian Golgi tendon organs. Sensory feedback from campaniform sensilla is integrated in the control of posture and locomotion.
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa, trochanter, femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus, patella.
Chordotonal organs are stretch receptor organs found only in insects and crustaceans. They are located at most joints and are made up of clusters of scolopidia that either directly or indirectly connect two joints and sense their movements relative to one another. They can have both extero- and proprioceptive functions, for example sensing auditory stimuli or leg movement. The word was coined by Vitus Graber in 1882, though he interpreted them as being stretched between two points like a string, sensing vibrations through resonance.
A sensillum is an arthropod sensory organ protruding from the cuticle of exoskeleton, or sometimes lying within or beneath it. Sensilla appear as small hairs or pegs over an individual's body. Inside each sensillum there are two to four sensory neurons. These neurons, or receptors, gather information about environment the arthropod is in:
Proprioception is the sense of self-movement, force, and body position.
Ultrasound avoidance is an escape or avoidance reflex displayed by certain animal species that are preyed upon by echolocating predators. Ultrasound avoidance is known for several groups of insects that have independently evolved mechanisms for ultrasonic hearing. Insects have evolved a variety of ultrasound-sensitive ears based upon a vibrating tympanic membrane tuned to sense the bat's echolocating calls. The ultrasonic hearing is coupled to a motor response that causes evasion of the bat during flight.
A scolopidium is the fundamental unit of a mechanoreceptor organ in insects. It is a composition of three cells: a scolopale cap cell which caps the scolopale cell, and a bipolar sensory nerve cell.
Haller's organ is a complex sensory organ possessed by hard and soft ticks. Not found outside of Acari, it is proposed to function like the chemosensation of insect antennae, but is structurally different. Ticks, being obligate parasites, must find a host in order to survive. Bloodmeals are necessary for completion of the life cycle, including reproduction and ontogenetic development. First described in 1881, it was named for its discoverer, Haller. While Haller initially proposed it was involved in auditory sensation, this was rejected in favor of olfactory sensation by 1905. This theory was supported by Lee's behavioral studies as early as 1948.
The subgenual organ is an organ in insects that is involved in the perception of sound. The name refers to the location of the organ just below the knee in the tibia of all legs in most insects.
Insect olfaction refers to the function of chemical receptors that enable insects to detect and identify volatile compounds for foraging, predator avoidance, finding mating partners and locating oviposition habitats. Thus, it is the most important sensation for insects. Most important insect behaviors must be timed perfectly which is dependent on what they smell and when they smell it. For example, olfaction is essential for locating host plants and hunting prey in many species of insects, such as the moth Deilephila elpenor and the wasp Polybia sericea, respectively.
The study of animal locomotion is a branch of biology that investigates and quantifies how animals move.
The femoral chordotonal organ is a group of mechanosensory neurons found in an insect leg that detects the movements and the position of the femur/tibia joint. It is thought to function as a proprioceptor that is critical for precise control of leg position by sending the information regarding the femur/tibia joint to the motor circuits in the ventral nerve cord and the brain
Bristle sensilla are a class of mechanoreceptors found in insects and other arthropods that respond to mechanical stimuli generated by the external world. As a result, they are considered exteroceptors. Bristle sensilla can be divided into two main types, macrochaete and microchaete, based on their size and physiology. The larger macrochaete are thicker and stouter than the smaller microchaete. Macrochaete are also more consistent in their number and distribution across individuals of the same species. Between species, the organization of macrochaete is more conserved among closely related species, whereas the organization of microchaete is more variable and less correlated with phylogenetic relatedness.