The Mauthner cells are a pair of big and easily identifiable
neuron
A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...
s (one for each half of the body) located in the
rhombomere
In the vertebrate embryo, a rhombomere is a transiently divided segment of the developing neural tube, within the hindbrain region (a neuromere) in the area that will eventually become the rhombencephalon. The rhombomeres appear as a series of sl ...
4 of the
hindbrain
The hindbrain, rhombencephalon (shaped like a rhombus) is a developmental categorization of portions of the central nervous system in vertebrates. It includes the medulla, pons, and cerebellum. Together they support vital bodily processes.
Met ...
in fish and
amphibian
Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s that are responsible for a very fast escape
reflex
In biology, a reflex, or reflex action, is an involuntary, unplanned sequence or action and nearly instantaneous response to a stimulus.
Reflexes are found with varying levels of complexity in organisms with a nervous system. A reflex occurs ...
(in the majority of animals – a so-called C-start response). The cells are also notable for their unusual use of both chemical and
electrical
Electricity is the set of physical phenomena associated with the presence and motion of matter possessing an electric charge. Electricity is related to magnetism, both being part of the phenomenon of electromagnetism, as described by Maxwel ...
synapses
In the nervous system, a synapse is a structure that allows a neuron (or nerve cell) to pass an electrical or chemical signal to another neuron or a target effector cell. Synapses can be classified as either chemical or electrical, depending o ...
.
[
]
Evolutionary history
Mauthner cells first appear in lampreys
Lampreys (sometimes inaccurately called lamprey eels) are a group of Agnatha, jawless fish comprising the order (biology), order Petromyzontiformes , sole order in the Class (biology), class Petromyzontida. The adult lamprey is characterize ...
(being absent in hagfish
Hagfish, of the Class (biology), class Myxini (also known as Hyperotreti) and Order (biology), order Myxiniformes , are eel-shaped Agnatha, jawless fish (occasionally called slime eels). Hagfish are the only known living Animal, animals that h ...
and lancelet
The lancelets ( ), also known as amphioxi (: amphioxus ), consist of 32 described species of somewhat fish-like benthic filter feeding chordates in the subphylum Cephalochordata, class Leptocardii, and family Branchiostomatidae.
Lancelets dive ...
s),[ and are present in virtually all ]teleost
Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
fish, as well as in amphibian
Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s (including postmetamorphic frog
A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order (biology), order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough ski ...
s and toad
Toad (also known as a hoptoad) is a common name for certain frogs, especially of the family Bufonidae, that are characterized by dry, leathery skin, short legs, and large bumps covering the parotoid glands.
In popular culture (folk taxonomy ...
s[). Some fish, such as lumpsuckers, seem to have lost the Mauthner cells however.]
Role in behavior
The C-start
A C-start is a type of a very quick startle or escape reflex that is employed by fish
A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
and amphibian
Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s (including larval
A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
frogs
A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough skin texture due to ...
and toads). There are two sequential stages in the C-start: first, the head rotates about the center of mass
In physics, the center of mass of a distribution of mass in space (sometimes referred to as the barycenter or balance point) is the unique point at any given time where the weight function, weighted relative position (vector), position of the d ...
towards the direction of future escape, and the body of the animal exhibits a curvature that resembles a letter C; then, at the second stage, the animal is propelled forward. The duration of these stages varies from species to species from about 10 to 20 ms for the first stage, and from 20 to 30 ms for the second.[ In fish this forward propulsion does not require contraction of the antagonistic ]muscle
Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...
, but results from the body stiffness and the hydrodynamic
In physics, physical chemistry and engineering, fluid dynamics is a subdiscipline of fluid mechanics that describes the flow of fluids – liquids and gases. It has several subdisciplines, including (the study of air and other gases in moti ...
resistance of the tail
The tail is the elongated section at the rear end of a bilaterian animal's body; in general, the term refers to a distinct, flexible appendage extending backwards from the midline of the torso. In vertebrate animals that evolution, evolved to los ...
. When an antagonistic muscular contraction does occur during stage 2, the fish rotates in the opposite direction, producing a counter-turn, and a directional change.
The role of the Mauthner cell in the C-start behavior
In cases when an abrupt acoustic, tactile or visual
The visual system is the physiological basis of visual perception (the ability to detect and process light). The system detects, transduces and interprets information concerning light within the visible range to construct an image and buil ...
stimulus elicits a single action potential
An action potential (also known as a nerve impulse or "spike" when in a neuron) is a series of quick changes in voltage across a cell membrane. An action potential occurs when the membrane potential of a specific Cell (biology), cell rapidly ri ...
in one M-cell, it always correlates with a contralateral
Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
C-start escape. An extremely quick mutual feedback
Feedback occurs when outputs of a system are routed back as inputs as part of a chain of cause and effect that forms a circuit or loop. The system can then be said to ''feed back'' into itself. The notion of cause-and-effect has to be handle ...
inhibitory
An inhibitory postsynaptic potential (IPSP) is a kind of synaptic potential that makes a Chemical synapse, postsynaptic neuron less likely to generate an action potential.Purves et al. Neuroscience. 4th ed. Sunderland (MA): Sinauer Associates, Inc ...
circuit then assures that only one M-cell reaches spiking threshold—as the C-start has to be unilateral
__NOTOC__
Unilateralism is any doctrine or agenda that supports one-sided action. Such action may be in disregard for other parties, or as an expression of a commitment toward a direction which other parties may find disagreeable. As a word, ''un ...
by definition—and that only one action potential is fired.
The Mauthner cell-mediated C-start reflex is very quick, with about 5-10 ms latency between the acoustic/tactile stimulus and the Mauthner cell discharge, and only about 2 ms between the discharge and the unilateral muscle contraction.[ Mauthner cells are thus the quickest motor neuron to respond to the stimulus. It makes the C-start response behaviorally important as a way to initiate the escape reflex in an ''all or nothing'' fashion, while the direction and speed of the escape can be corrected later through the activity of smaller motor neurons.
In ]larval
A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
zebrafish
The zebrafish (''Danio rerio'') is a species of freshwater ray-finned fish belonging to the family Danionidae of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (an ...
about ~60% of the total population of reticulospinal neurons are also activated by a stimulus that elicits the M-spike and C-start escape. A well-studied group of these reticulospinal neurons are the bilaterally paired M-cell homologues denoted MiD2cm and MiD3cm. These neurons exhibit morphological similarities to the M-cell including a lateral and ventral dendrite. They are located in rhombomere
In the vertebrate embryo, a rhombomere is a transiently divided segment of the developing neural tube, within the hindbrain region (a neuromere) in the area that will eventually become the rhombencephalon. The rhombomeres appear as a series of sl ...
s 5 and 6 of hindbrain
The hindbrain, rhombencephalon (shaped like a rhombus) is a developmental categorization of portions of the central nervous system in vertebrates. It includes the medulla, pons, and cerebellum. Together they support vital bodily processes.
Met ...
respectively, and also receive auditory input in parallel with the M-cell from the pVIIIth nerve. In fish, water jet stimuli that activate these neurons elicit non-mauthner initiated C-starts of a longer latency, compared with M-cell associated ones.
Although the M-cell is often considered the prototype of a command neuron in vertebrate
Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain.
The vertebrates make up the subphylum Vertebra ...
s, this designation may not be fully warranted. Although electrical stimulation of the M-cell is sufficient for eliciting a C-start, this C-start is normally weaker than the one evoked by a sensory stimulus. Moreover, the C-start can be evoked even with the M-cell ablated, although in this case the latency of the response increases. The most widely accepted model of the M-cell system, or brainstem escape network, is that the M-cell initiates a fixed action pattern to the left or right by activating a spinal motor circuit initially described by J. Diamond and colleagues, but the precise trajectory of the escape is encoded by population activity in the other classes of reticulospinal neurons functioning in parallel to the M-cell. This notion is supported by studies using ''in vivo'' calcium imaging in larval zebrafish which show that MiD2cm and MiD3cm are activated along with the M-cell when an offending stimulus is directed towards the head but not the tail, and are correlated with C-starts of a larger initial turn angle.
Another component of the escape response is mediated by cranial relay neurons that are activated by the Mauthner cell spike. These neurons are electrically coupled with motoneurons which innervate extraocular, jaw and opercular muscles and mediate pectoral fin adduction in hatchetfish. This component of the neural circuit was first described by Michael V.L. Bennett and colleagues.
Mauthner cells in other types of behavior
Mauthner cells may be involved into behavioral patterns other than the C-start, if these types of behavior also require extremely quick bending movement of the body. Thus in goldfish
The goldfish (''Carassius auratus'') is a freshwater fish in the family Cyprinidae of the order Cypriniformes. It is commonly kept as a pet in indoor aquariums, and is one of the most popular aquarium fish. Goldfish released into the w ...
Mauthner cells are activated during prey capture near the surface of the water, as this type of hunting is dangerous for the fish, and it would benefit from leaving the surface as soon as possible after the prey is captured.
In adult postmetamorphic anurans
A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough skin texture due to ...
(frogs and toads) that do not have a tail, M-cells are nevertheless preserved and their discharges are associated with rapid movement of leg
A leg is a weight-bearing and locomotive anatomical structure, usually having a columnar shape. During locomotion, legs function as "extensible struts". The combination of movements at all joints can be modeled as a single, linear element cap ...
s during an escape. In addition, larval lamprey
Lampreys (sometimes inaccurately called lamprey eels) are a group of Agnatha, jawless fish comprising the order (biology), order Petromyzontiformes , sole order in the Class (biology), class Petromyzontida. The adult lamprey is characterize ...
s (eel-like jawless fish of superclass Cyclostomata) exhibit rapid withdrawal behavior that is correlated with Mauthner cell activity and involves bilateral, posture-dependent muscular contractions along the length of the body. Larval lampreys (ammocoetes) are filter feeders that occupy crescent-shaped burrows in the silt or mud bottoms of freshwater stream beds, with their mouths positioned at, or just above the surface of the mud. Sudden vibration activates both Mauthner neurons in the lamprey brainstem, which causes an accordion-like muscular contraction in the trunk and tail and pulls the head down into the burrow.
Morphology and connections
Inputs to the M-cell: excitation and feed forward inhibition
The M-cell has two primary aspiny (lacking dendritic spine
A dendritic spine (or spine) is a small membrane protrusion from a neuron's dendrite that typically receives input from a single axon at the synapse. Dendritic spines serve as a storage site for synaptic strength and help transmit electrical sign ...
s) dendrite
A dendrite (from Ancient Greek language, Greek δένδρον ''déndron'', "tree") or dendron is a branched cytoplasmic process that extends from a nerve cell that propagates the neurotransmission, electrochemical stimulation received from oth ...
s which receive segregated inputs from various parts of the neural system.[ One dendrite projects laterally and the other projects either in the ventral or medial direction, depending on the species.
The ventral dendrite receives information from the ]optic tectum
In neuroanatomy, the superior colliculus () is a structure lying on the roof of the mammalian midbrain. In non-mammalian vertebrates, the homologous structure is known as the optic tectum or optic lobe. The adjective form '' tectal'' is commonly ...
and spinal cord
The spinal cord is a long, thin, tubular structure made up of nervous tissue that extends from the medulla oblongata in the lower brainstem to the lumbar region of the vertebral column (backbone) of vertebrate animals. The center of the spinal c ...
while the lateral dendrite receives inputs from the octovolateralis systems (the lateral line
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 epithelia ...
, acoustic inputs from the inner ear
The inner ear (internal ear, auris interna) is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the ...
, and inertial information from the statoliths brought by the cranial nerve VIII
The vestibulocochlear nerve or auditory vestibular nerve, also known as the eighth cranial nerve, cranial nerve VIII, or simply CN VIII, is a cranial nerve that transmits sound and equilibrium (balance) information from the inner ear to the bra ...
).[
The fibers from the ]ipsilateral
Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
cranial nerve VIII terminate in excitatory mixed electrical
Electricity is the set of physical phenomena associated with the presence and motion of matter possessing an electric charge. Electricity is related to magnetism, both being part of the phenomenon of electromagnetism, as described by Maxwel ...
and glutamate
Glutamic acid (symbol Glu or E; known as glutamate in its anionic form) is an α-amino acid that is used by almost all living beings in the biosynthesis of proteins. It is a Essential amino acid, non-essential nutrient for humans, meaning that ...
rgic synapses
In the nervous system, a synapse is a structure that allows a neuron (or nerve cell) to pass an electrical or chemical signal to another neuron or a target effector cell. Synapses can be classified as either chemical or electrical, depending o ...
on the M-cell. They also electrically activate glycine
Glycine (symbol Gly or G; ) is an amino acid that has a single hydrogen atom as its side chain. It is the simplest stable amino acid. Glycine is one of the proteinogenic amino acids. It is encoded by all the codons starting with GG (G ...
rgic inhibitory interneurons that terminate on the M-cells. Despite the inhibitory input having one more synapse in its pathway, there is no delay between the excitation and inhibition because the intervening synapse is electrical. It was shown that for weak stimuli the inhibition wins over the excitation, preventing the M-cell from a discharge, while for stronger stimuli excitation becomes dominant. The Inner ear
The inner ear (internal ear, auris interna) is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the ...
afferents also terminate with electrical synapses on a population PHP inhibitory interneurons (see below) to provide an additional level of feed forward inhibition. The Mauthner cell also has GABA
GABA (gamma-aminobutyric acid, γ-aminobutyric acid) is the chief inhibitory neurotransmitter in the developmentally mature mammalian central nervous system. Its principal role is reducing neuronal excitability throughout the nervous system.
GA ...
-, dopamine
Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic chemical of the catecholamine and phenethylamine families. It is an amine synthesized ...
-, serotonin
Serotonin (), also known as 5-hydroxytryptamine (5-HT), is a monoamine neurotransmitter with a wide range of functions in both the central nervous system (CNS) and also peripheral tissues. It is involved in mood, cognition, reward, learning, ...
- and somatostatin
Somatostatin, also known as growth hormone-inhibiting hormone (GHIH) or by #Nomenclature, several other names, is a peptide hormone that regulates the endocrine system and affects neurotransmission and cell proliferation via interaction with G ...
ergic inputs, each restricted to certain dendritic region.[
Inputs from the optic tectum and the lateral line help control which way the C-startle bends by biasing the mauthner cells when there are obstacles in the vicinity. In cases where movement away from the stimulus is blocked, the fish may bend towards the disturbance.][
]
Axon cap
The Mauthner cell axon hillock
The axon hillock is a specialized part of the cell body (or soma) of a neuron that connects to the axon. It can be identified using light microscopy from its appearance and location in a neuron and from its sparse distribution of Nissl substanc ...
is surrounded by a dense formation of neuropil, called the axon cap.[ The high resistance of this axon cap contributes to the typical shape of the Mauthner cell field potential (see below). In its most advanced form the axon cap consists of a core, immediately adjacent to the Mauthner cell axon, and containing a network of very thin ]unmyelinated
Myelin Sheath ( ) is a lipid-rich material that in most vertebrates surrounds the axons of neurons to Insulator (electricity), insulate them and increase the rate at which electrical impulses (called action potentials) pass along the axon. The my ...
fibers, and a peripheral part. This peripheral part contains the large unmyelinated fibers of the PHP neurons (see below) that mediate the inhibitory feedback to the Mauthner cell; the Mauthner cell itself also sends small dendrites from its axon hill to the peripheral part of the axon cap. Finally, the surface of the axon cap is covered with a ''cap wall'' composed of several layers of astrocyte
Astrocytes (from Ancient Greek , , "star" and , , "cavity", "cell"), also known collectively as astroglia, are characteristic star-shaped glial cells in the brain and spinal cord. They perform many functions, including biochemical control of en ...
-like glia
Glia, also called glial cells (gliocytes) or neuroglia, are non-neuronal cells in the central nervous system (the brain and the spinal cord) and in the peripheral nervous system that do not produce electrical impulses. The neuroglia make up ...
l cells. Both glial cells and the unmyelinated fibers are coupled with each other by means of gap junction
Gap junctions are membrane channels between adjacent cells that allow the direct exchange of cytoplasmic substances, such small molecules, substrates, and metabolites.
Gap junctions were first described as ''close appositions'' alongside tight ...
s.[
Evolutionarily, the axon cap is a more recent development than the Mauthner cell itself, so some animals, such as ]lamprey
Lampreys (sometimes inaccurately called lamprey eels) are a group of Agnatha, jawless fish comprising the order (biology), order Petromyzontiformes , sole order in the Class (biology), class Petromyzontida. The adult lamprey is characterize ...
s and eel
Eels are ray-finned fish belonging to the order Anguilliformes (), which consists of eight suborders, 20 families, 164 genera, and about 1000 species. Eels undergo considerable development from the early larval stage to the eventual adult stage ...
s, while having functional Mauthner cells, don't have axon cap at all, while some other animals, such as amphibia
Amphibians are ectothermic, anamniotic, four-limbed vertebrate animals that constitute the class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all tetrapods, but excluding the amniotes (tetrapods with an amniotic ...
and lungfish
Lungfish are freshwater vertebrates belonging to the class Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, inc ...
, do have a very simplified version of it.
Feedback network
The main part of the Mauthner cell-associated network is the negative feedback
Feedback occurs when outputs of a system are routed back as inputs as part of a chain of cause and effect that forms a circuit or loop. The system can then be said to ''feed back'' into itself. The notion of cause-and-effect has to be handle ...
network, which assures that only one of the two Mauthner cells fires in response to the stimulus and that, whichever Mauthner cell fires, it does so only once. Both these requirements are quite natural considering that the consequences of a single Mauthner cell discharge are so strong; a failure to comply with these two rules would not only prevent the animal from escaping, but could even physically damage it. The fastest part of this negative feedback network, which is also the one closest to the Mauthner cell, is that of the so-called ''passive hyperpolarizing field potential'' or PHP neurons.[ The fibers of these neurons are located in the axon cap, and they receive inputs from both ]ipsilateral
Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
and contralateral
Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
Mauthner cells. The field potentials of PHP neurons are strongly positive, and form a part of the 'Signature field potential' of the Mauthner cell (see below), with the early (ipsilaterally initiated) component being called the Extracellular Hyperpolarizing Potential (EHP), and the later (contralaterally initiated) component being sometimes addressed in the literature as the Late Collateral Inhibition (LCI).[ The action of PHP neurons onto the Mauthner cells is mediated by electrical, and not chemical effects: the outward currents generated by the ]action potential
An action potential (also known as a nerve impulse or "spike" when in a neuron) is a series of quick changes in voltage across a cell membrane. An action potential occurs when the membrane potential of a specific Cell (biology), cell rapidly ri ...
s in axon cap fibers flow inward across the Mauthner cell axon hillock
The axon hillock is a specialized part of the cell body (or soma) of a neuron that connects to the axon. It can be identified using light microscopy from its appearance and location in a neuron and from its sparse distribution of Nissl substanc ...
and hyperpolarize it.[
]
Outputs
The only axon
An axon (from Greek ἄξων ''áxōn'', axis) or nerve fiber (or nerve fibre: see American and British English spelling differences#-re, -er, spelling differences) is a long, slender cellular extensions, projection of a nerve cell, or neuron, ...
of the Mauthner cell reaches from the cell to the midline of the hindbrain
The hindbrain, rhombencephalon (shaped like a rhombus) is a developmental categorization of portions of the central nervous system in vertebrates. It includes the medulla, pons, and cerebellum. Together they support vital bodily processes.
Met ...
, promptly crosses it to the contralateral side, and then descends caudally along the spinal cord
The spinal cord is a long, thin, tubular structure made up of nervous tissue that extends from the medulla oblongata in the lower brainstem to the lumbar region of the vertebral column (backbone) of vertebrate animals. The center of the spinal c ...
.[ A single discharge of the M-cell achieves a whole set of parallel effects onto the spinal motor networks: 1) it monosynaptically excites large primary ]motoneuron
A motor neuron (or motoneuron), also known as efferent neuron is a neuron whose cell body is located in the motor cortex, brainstem or the spinal cord, and whose axon (fiber) projects to the spinal cord or outside of the spinal cord to directly o ...
s at one side of the body; 2) disynaptically excites smaller motoneurons at the same side of the body; 3) initiates action potentials in inhibitory interneuron
Interneurons (also called internuncial neurons, association neurons, connector neurons, or intermediate neurons) are neurons that are not specifically motor neurons or sensory neurons. Interneurons are the central nodes of neural circuits, enab ...
s electrically coupled to the M-cell axon, and by their means inhibits a) inhibitory interneurons still at the same side of the body (to prevent them from interfering with the C-start), as well as b) motoneurons at the other side of the body. As a result of this pattern of activation the quick muscle
Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...
s at one side of the body contract simultaneously, while the muscles at the other side of the body relax.
Electrophysiology
Ephaptic properties
200px, Ephaptic inhibition at the mauthner axon cap by PHP cells
The inhibition of the M-cell by the PHP cells occurs by ephaptic interactions. The inhibition is brought about without a chemical synapse
Chemical synapses are biological junctions through which neurons' signals can be sent to each other and to non-neuronal cells such as those in muscles or glands. Chemical synapses allow neurons to form circuits within the central nervous syste ...
s or electrical synaptic coupling having low resistance gap junctions
Gap junctions are Membrane channel, membrane channels between adjacent cells that allow the direct exchange of cytoplasmic substances, such small molecules, substrates, and metabolites.
Gap junctions were first described as ''close appositions' ...
joining the cells. When the region of the PHP cell axon outside the axon cap depolarizes, the influx of positive charge into the cell through voltage gated sodium channels is accompanied by a passive outflow of current from the PHP cell axon into the region bound by the axon cap. Due to the high resistance of the surrounding glial cells, the charge does not dissipate and the potential across the M-cell membrane is increased, hyperpolarizing it.
Signature field potential
Because of its size, presence of a quick feedback network, and abundance of electrical
Electricity is the set of physical phenomena associated with the presence and motion of matter possessing an electric charge. Electricity is related to magnetism, both being part of the phenomenon of electromagnetism, as described by Maxwel ...
and quasi-electrical ( ephaptic) synapses, the Mauthner cell has a strong field potential of a very characteristic shape. This field potential starts with a high-amplitude potential sink up to tens of millivolts in amplitude that originates from the Mauthner cell discharge, and which is closely followed by a positive potential, called Extrinsic Hyperpolarizing Potential or EHP, which is associated with the activity of the recurrent feedback network.[
Due to its high amplitude, in some animals the negative part of Mauthner cell field potential can be detected up to several hundred micrometres away from the cell itself.][ The positive components of the field potential are strongest in the axon cap, reaching amplitudes of 45 mV in adult goldfish.][ With a knowledge of these properties of the field potential, it is possible to use field potential monitoring as a way to find the Mauthner cell body ]in vivo
Studies that are ''in vivo'' (Latin for "within the living"; often not italicized in English) are those in which the effects of various biological entities are tested on whole, living organisms or cells, usually animals, including humans, an ...
, or in vitro
''In vitro'' (meaning ''in glass'', or ''in the glass'') Research, studies are performed with Cell (biology), cells or biological molecules outside their normal biological context. Colloquially called "test-tube experiments", these studies in ...
in a whole brain preparation, moving the recording electrode in the hindbrain
The hindbrain, rhombencephalon (shaped like a rhombus) is a developmental categorization of portions of the central nervous system in vertebrates. It includes the medulla, pons, and cerebellum. Together they support vital bodily processes.
Met ...
, while at the same time stimulating the spinal cord
The spinal cord is a long, thin, tubular structure made up of nervous tissue that extends from the medulla oblongata in the lower brainstem to the lumbar region of the vertebral column (backbone) of vertebrate animals. The center of the spinal c ...
, thus evoking antidromic action potentials in the Mauthner cell axon.[
]
Plasticity
Application of serotonin
Serotonin (), also known as 5-hydroxytryptamine (5-HT), is a monoamine neurotransmitter with a wide range of functions in both the central nervous system (CNS) and also peripheral tissues. It is involved in mood, cognition, reward, learning, ...
was shown to increase inhibitory inputs to the M-cell, while application of dopamine
Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic chemical of the catecholamine and phenethylamine families. It is an amine synthesized ...
– to increase the amplitude of both chemical and electrical components of the VIIIth nerve responses via a G protein
G proteins, also known as guanine nucleotide-binding proteins, are a Protein family, family of proteins that act as molecular switches inside cells, and are involved in transmitting signals from a variety of stimuli outside a cell (biology), ...
-mediated activation of postsynaptic D2 receptor
Dopamine receptor D2, also known as D2R, is a protein that, in humans, is encoded by the ''DRD2'' gene. After work from Paul Greengard's lab had suggested that dopamine receptors were the site of action of antipsychotic drugs, several groups, ...
.[ An activity-dependent LTP can be evoked in M-cells by a high-frequency stimulation of the VIIIth nerve. Surprisingly, this LTP is ]electrical synapse
An electrical synapse, or gap junction, is a mechanical and electrically conductive synapse, a functional junction between two neighboring neurons. The synapse is formed at a narrow gap between the pre- and postsynaptic neurons known as a gap junc ...
-mediated, and is presumed to involve modification of the gap junction
Gap junctions are membrane channels between adjacent cells that allow the direct exchange of cytoplasmic substances, such small molecules, substrates, and metabolites.
Gap junctions were first described as ''close appositions'' alongside tight ...
channels.[ A possibility of LTP induction by sensory stimuli ]in vivo
Studies that are ''in vivo'' (Latin for "within the living"; often not italicized in English) are those in which the effects of various biological entities are tested on whole, living organisms or cells, usually animals, including humans, an ...
,[ and the evidence for the LTP of inhibitory inputs to M-cells][ were also demonstrated.
Spontaneous preference in turn direction in young goldfish is correlated with one of the Mauthner cells being bigger than the other one. It is possible to change the preference of fish by raising them in conditions facilitating turns in a specific direction; this shift is accompanied by a correspondent change in M-cell sizes.]
History of research
The Mauthner cell was first identified by the Viennese ophthalmologist Ludwig Mauthner in the teleost
Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
fish for its associated neural circuit which mediates an escape response called the C-start or C- startle to direct the fish away from a predator.
The M-cell is a model system in the field of Neuroethology
Neuroethology is the evolutionary and comparative approach to the study of animal behavior and its underlying mechanistic control by the nervous system. It is an interdisciplinary science that combines both neuroscience (study of the nervous s ...
. The M-cell system has served for detailed neurophysiological
Neurophysiology is a branch of physiology and neuroscience concerned with the functions of the nervous system and their mechanisms. The term ''neurophysiology'' originates from the Greek word ''νεῦρον'' ("nerve") and ''physiology'' (whic ...
and histological
Histology,
also known as microscopic anatomy or microanatomy, is the branch of biology that studies the microscopic anatomy of biological tissue (biology), tissues. Histology is the microscopic counterpart to gross anatomy, which looks at large ...
investigations of synaptic transmission
Neurotransmission (Latin: ''transmissio'' "passage, crossing" from ''transmittere'' "send, let through") is the process by which signaling molecules called neurotransmitters are released by the axon terminal of a neuron (the presynaptic neuron) ...
and synaptic plasticity
In neuroscience, synaptic plasticity is the ability of synapses to Chemical synapse#Synaptic strength, strengthen or weaken over time, in response to increases or decreases in their activity. Since memory, memories are postulated to be represent ...
.[ Studies by Donald Faber and Henri Korn helped to establish the one vesicle hypothesis of ]synaptic transmission
Neurotransmission (Latin: ''transmissio'' "passage, crossing" from ''transmittere'' "send, let through") is the process by which signaling molecules called neurotransmitters are released by the axon terminal of a neuron (the presynaptic neuron) ...
in the CNS. Other important research topics that have been investigated in the M-cell system include studies by Yoichi Oda and colleagues on inhibitory long-term potentiation
In neuroscience, long-term potentiation (LTP) is a persistent strengthening of synapses based on recent patterns of activity. These are patterns of synaptic activity that produce a long-lasting increase in signal transmission between two neuron ...
and auditory conditioning Conditioning may refer to:
Science, computing, and technology
* Air conditioning, the removal of heat from indoor air for thermal comfort
** Automobile air conditioning, air conditioning in a vehicle
** Ice storage air conditioning, air conditio ...
of the startle response, and studies by Alberto Pereda and colleagues on plasticity of electrical synapse
An electrical synapse, or gap junction, is a mechanical and electrically conductive synapse, a functional junction between two neighboring neurons. The synapse is formed at a narrow gap between the pre- and postsynaptic neurons known as a gap junc ...
s. Other research topics investigated in the M-cell system include studies of spinal neural networks and neural regeneration by Joe Fetcho and colleagues, as well as underwater sound localization
Sound localization is a listener's ability to identify the location or origin of a detected sound in direction and distance.
The sound localization mechanisms of the mammalian auditory system have been extensively studied. The auditory system u ...
, and the biophysics of computation in single neurons.
References
Further reading
*
*
*
*
*
*
*
*
*
{{Neuroethology
Neuroethology
Fish nervous system