Myomere

Last updated

Myomeres are blocks of skeletal muscle tissue arranged in sequence, commonly found in aquatic chordates. Myomeres are separated from adjacent myomeres by connective fascia (myosepta) and most easily seen in larval fishes or in the olm. Myomere counts are sometimes used for identifying specimens, since their number corresponds to the number of vertebrae in the adults. Location varies, with some species containing these only near the tails, while some have them located near the scapular or pelvic girdles. Depending on the species, myomeres could be arranged in an epaxial or hypaxial manner. Hypaxial refers to ventral muscles and related structures while epaxial refers to more dorsal muscles. The horizontal septum divides these two regions in vertebrates from cyclostomes to gnathostomes. In terrestrial chordates, the myomeres become fused as well as indistinct, due to the disappearance of myosepta.

Contents

Shape

Filet of salmon showing the zig-zag of myomeres Myomeres.jpg
Filet of salmon showing the zig-zag of myomeres
Fillet of iridescent shark showing the zig-zag of myomeres Pangasius meat.jpg
Fillet of iridescent shark showing the zig-zag of myomeres

The shape of myomeres varies by species. Myomeres are commonly zig-zag, "V" (lancelets), "W" (fishes), or straight (tetrapods)– shaped muscle fibers. Generally, cyclostome myomeres are arranged in vertical strips while those of jawed fishes are folded in a complex matter due to swimming capability evolution. Specifically, myomeres of elasmobranchs and eels are “W”-shaped. Contrastingly, myomeres of tetrapods run vertically and do not display complex folding. Another species with simply-lain myomeres are mudpuppies. Myomeres overlap each other in succession, meaning myomere activation also allows neighboring myomeres to activate. [1]

Myomeres are made up of myoglobin-rich dark muscle as well as white muscle. Dark muscle, generally, functions as slow-twitch muscle fibers while white muscle is composed of fast-twitch fibers.

Function

Specifically, three types of myomeres in fish-like chordates include amphioxine (lancelet), cyclostomine (jawless fish), and gnathostomine (jawed fish). A common function shared by all of these is that they function to flex the body laterally into concavity to provide force for locomotion. [1]

Since myomeres are composed of multinucleated myofibers (contractile cells), force can be generated via muscle contraction that gets transmitted by the intricate connective tissue (myosepta) network.

Function in fishes

The folded shape of each myomere as "V" or "W" shaped extends over various axial segments, allowing fibers control over a large amount of the body. Specifically, myomeres are overlapping cones bound by connective tissue. Myomeres compose most of the lateral musculature and provide propulsive force to travel along the line of travel. In this sense, they cause flexion to either side in order to produce locomotor force. Myomeres attach to centra of vertebrae, and neural and haemal spines.

Further, myomeres of fish are divided by a horizontal septum into dorsal (epaxial) and ventral (hypaxial) sections as mentioned in previous paragraphs. Further, spinal nerves pass into each myomere. [2]

There are different variations of myomere activation depending on the type of swimming or movement. For example, high loading situations such as fast-starts and turning require almost maximal myomere activation in teleost fish. Further, if swim speeds are lower and movement is in one plane, there is less activation of myomeres. Further, research has discovered that fish are able to spatially restrict axial myomeres during different swimming behaviors. [3] [4]

Some research theorizes that myomeres play additional roles in for the fish beyond force generation for swimming. For example, this microdissection and polarized light microscopy research suggests that anterior myomeres have elongated and reinforced dorsal posterior cones that allow epaxial muscle force to be transmitted to the neurocranium for elevation during suction feeding.

Specific taxa

Fossils

Published information on Pikaia gracilens (a well-known Cambrian era fossil) explains evolution of swimming ability in chordates related to myomere shape and function. Specifically, myomeres in this species possessed minimal overlap between successive ones and myosepta dividing them were gently curved. In a biomechanical evaluation, it is presumed that Pikaia were not capable of rapid swimming like in living chordates. Several theories for this idea include lacking fast-twitch muscle fibers, ancestral muscle fiber types more like modern slow-twitch fibers, and less tension on myosepta due to less overlap between successive myomeres. [5]

Larval fish and Amphioxus

Larval fish and amphioxus myomeres are "V"-shaped. They are involved in the specialized notochord of amphioxus. There are muscle cells within myomeres that send, and synapse cytoplasmic extensions of muscle cells with contractile fibrils to the nerve cord surface.

In amphioxus, myomeres run longitudinally along the length of the body in a "V"-shape. As sequential contraction for swimming occurs, force from the myomeres is transmitted via connective tissues to the notochord.

Zebrafish

The tail-bending maneuver generated by myomeres in zebrafish requires innervation from motor neurons for both the hypaxial and epaxial muscle regions. It has been found that timing/intensity of neurons firing in these two regions varies, respectively. This process is mediated by a circuit that controls motor neuron activation during swimming behaviors, which, in turn, affects force generation. Similar to this idea, one study found that hypaxial and epaxial myomere activation did not always correlate with myomeric fibers closer to the horizontal septum itself. [6]

Tetrapods

Myomeres run vertically and do not undergo folding like in bony fishes. Further, in higher order vertebrates, myomeres are fused and run longitudinally. Myosepta that divides myomeres are completely obsolete in amniotes.

Myomeres also play a role in swimming in adult newts. Specifically, epaxial myomeres located opposite to each other at the same longitudinal site alternate rhythmic contraction. During stepping on the ground, the myomeres of the mid-trunk undergo bursts of contraction that are synchronized in contrast to double bursting patterns (in opposite directions) expressed in the anterior and posterior trunks. [7]

In salamanders, hypaxial muscles, myomeres, and myosepta run in a straight line mid-laterally to mid-ventrally. Specifically, the orientation of collagen fibers within these myomeres runs mediolateral. It is also theorized that, in salamanders, myosepta increase the amplification of strain of angled muscle fibers. This controls how myomeres bulge during contraction in what is called the 'bulge control hypothesis'. [8]

Eels

Eel myomeres are "W"-shaped and cover the entire body. Within these is a mucosal-like matrix that is a-cellular. Superficial to these myomeres is an epithelial layer.

Mudpuppy

Salamanders in the genus Necturus (mudpuppies) are a salamander species with simply-lain myomeres, unlike the complex nature of bony fishes. [9]

Chondrichthyes

The myomeres of some Chondrichthyes, specifically sharks, are "W"-shaped. Thus, function in Chondrichthyes is similar to that of bony fish, where myomeres contribute to propulsive force for locomotion.

Leptocephali

Leptocephalus myomeres are "W"-shaped and extend from head all the way to the tail. Distinguishing eels can be done through evaluation of the number of myomeres (European has 112-119 while American has 103–11).

Related Research Articles

<span class="mw-page-title-main">Skeleton</span> Part of the body that forms the supporting structure

A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal frame to which the organs and soft tissues attach; and the hydroskeleton, a flexible internal structure supported by the hydrostatic pressure of body fluids.

<span class="mw-page-title-main">Skeletal muscle</span> One of three major skeletal system types that connect to bones

Skeletal muscles are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscle tissue, and are often known as muscle fibers. The muscle tissue of a skeletal muscle is striated – having a striped appearance due to the arrangement of the sarcomeres.

In biology, a motor unit is made up of a motor neuron and all of the skeletal muscle fibers innervated by the neuron's axon terminals, including the neuromuscular junctions between the neuron and the fibres. Groups of motor units often work together as a motor pool to coordinate the contractions of a single muscle. The concept was proposed by Charles Scott Sherrington.

A hydrostatic skeleton or hydroskeleton is a type of skeleton supported by hydrostatic fluid pressure, common among soft-bodied invertebrate animals colloquially referred to as "worms". While more advanced organisms can be considered hydrostatic, they are sometimes referred to as hydrostatic for their possession of a hydrostatic organ instead of a hydrostatic skeleton, where the two may have the same capabilities but are not the same. As the prefix hydro- meaning "water", being hydrostatic means being fluid-filled.

<span class="mw-page-title-main">Notochord</span> Flexible rod-shaped structure in all chordates

In zoology and developmental anatomy, the notochord is an elastic rod-like structure found in many deuterostomal animals. Any species that has a notochord at any stage of its life cycle is, by definition, a chordate.

<i>Pikaia</i> Extinct genus of primitive chordates

Pikaia gracilens is an extinct species of primitive chordate animal known from the Middle Cambrian Burgess Shale of British Columbia. Described in 1911 by Charles Doolittle Walcott as an annelid, and in 1979 by Harry B. Whittington and Simon Conway Morris as a chordate, it became "one of the most famous early chordate fossils," or "famously known as the earliest described Cambrian chordate". It is estimated to have lived during the latter period of the Cambrian explosion. Since its initial discovery, more than a hundred specimens have been recovered.

<span class="mw-page-title-main">Cephalochordate</span> Subphylum of lancelets

A cephalochordate is an animal in the chordate subphylum Cephalochordata. Cephalochordates are commonly called lancelets, and possess 5 synapomorphies, or primary characteristics, that all chordates have at some point during their larval or adulthood stages. These 5 synapomorphies are a notochord, dorsal hollow nerve cord, endostyle, pharyngeal slits, and a post-anal tail. The fine structure of the cephalochordate notochord is best known for the Bahamas lancelet, Asymmetron lucayanum. Cephalochordates are represented in modern oceans by the Amphioxiformes and are commonly found in warm temperate and tropical seas worldwide. With the presence of a notochord, adult amphioxus are able to swim and tolerate the tides of coastal environments, but they are most likely to be found within the sediment of these communities.

<span class="mw-page-title-main">Lancelet</span> Order of chordates

The lancelets, also known as amphioxi, consist of some 30 to 35 species of "fish-like" benthic filter feeding chordates in the order Amphioxiformes. They are modern representatives of the subphylum Cephalochordata. Lancelets closely resemble 530-million-year-old Pikaia, fossils of which are known from the Burgess Shale. However, according to phylogenetic analysis, the lancelet group itself probably evolved around the Cretaceous, 97.7 million years ago for Pacific species and 112 million years ago for Atlantic species. Palaeobranchiostoma from the Permian may be part of the fossil record of lancelets; however, due to poor preservation, some doubt about its nature remains. Zoologists are interested in them because they provide evolutionary insight into the origins of vertebrates. Lancelets contain many organs and organ systems that are closely related to those of modern fish, but in a more primitive form. Therefore, they provide a number of examples of possible evolutionary exaptation. For example, the gill-slits of lancelets are used for feeding only, and not for respiration. The circulatory system carries food throughout their body, but does not have red blood cells or hemoglobin for transporting oxygen. Lancelet genomes hold clues about the early evolution of vertebrates: by comparing genes from lancelets with the same genes in vertebrates, changes in gene expression, function and number as vertebrates evolved can be discovered. The genome of a few species in the genus Branchiostoma have been sequenced: B. floridae,B. belcheri, and B. lanceolatum.

<span class="mw-page-title-main">Motor unit recruitment</span> Additional activation of motor units to increase contractile strength

Motor unit recruitment is the activation of additional motor units to accomplish an increase in contractile strength in a muscle. A motor unit consists of one motor neuron and all of the muscle fibers it stimulates. All muscles consist of a number of motor units and the fibers belonging to a motor unit are dispersed and intermingle amongst fibers of other units. The muscle fibers belonging to one motor unit can be spread throughout part, or most of the entire muscle, depending on the number of fibers and size of the muscle. When a motor neuron is activated, all of the muscle fibers innervated by the motor neuron are stimulated and contract. The activation of one motor neuron will result in a weak but distributed muscle contraction. The activation of more motor neurons will result in more muscle fibers being activated, and therefore a stronger muscle contraction. Motor unit recruitment is a measure of how many motor neurons are activated in a particular muscle, and therefore is a measure of how many muscle fibers of that muscle are activated. The higher the recruitment the stronger the muscle contraction will be. Motor units are generally recruited in order of smallest to largest as contraction increases. This is known as Henneman's size principle.

<span class="mw-page-title-main">Myogenesis</span> Formation of muscular tissue, particularly during embryonic development

Myogenesis is the formation of skeletal muscular tissue, particularly during embryonic development.

<span class="mw-page-title-main">Shark anatomy</span>

Shark anatomy differs from that of bony fish in a variety of ways. Variation observed within shark anatomy is a potential result of speciation and habitat variation.

<i>Cathaymyrus</i> Extinct genus of lancelets

Cathaymyrus is a genus of Early Cambrian cephalochordate known from the Chengjiang locality in Yunnan Province, China. Both species have a long segmented body with no distinctive head. The segments resemble the v-shaped muscle blocks found in similar cephalochordates such as Amphioxus. A long linear impression runs along the "back" of the body looking something like a chordate notochord. Cathaymyrus is related to Pikaia, another cephalochordate from the Burgess Shale.

<span class="mw-page-title-main">Muscle</span> Basic biological tissue present in animals

Muscle is a soft tissue, one of the four basic types of animal tissue. Muscle tissue gives skeletal muscles the ability to contract. Muscle is formed during embryonic development, in a process known as myogenesis. Muscle tissue contains special contractile proteins called actin and myosin which interact to cause movement. Among many other muscle proteins present are two regulatory proteins, troponin and tropomyosin.

<i>Metaspriggina</i> Cambrian fossil genus of chordate

Metaspriggina is a genus of chordate initially known from two specimens in the Middle Cambrian Burgess Shale and 44 specimens found in 2012 at the Marble Canyon bed in Kootenay National Park.

<span class="mw-page-title-main">Undulatory locomotion</span>

Undulatory locomotion is the type of motion characterized by wave-like movement patterns that act to propel an animal forward. Examples of this type of gait include crawling in snakes, or swimming in the lamprey. Although this is typically the type of gait utilized by limbless animals, some creatures with limbs, such as the salamander, forgo use of their legs in certain environments and exhibit undulatory locomotion. In robotics this movement strategy is studied in order to create novel robotic devices capable of traversing a variety of environments.

<span class="mw-page-title-main">Epaxial and hypaxial muscles</span> Trunk muscles

In adult vertebrates, trunk muscles can be broadly divided into hypaxial muscles, which lie ventral to the horizontal septum of the vertebrae and epaxial muscles, which lie dorsal to the septum. Hypaxial muscles include some vertebral muscles, the diaphragm, the abdominal muscles, and all limb muscles. The serratus posterior inferior and serratus posterior superior are innervated by the ventral primary ramus and are hypaxial muscles. Epaxial muscles include other (dorsal) muscles associated with the vertebrae, ribs, and base of the skull. In humans, the erector spinae, the transversospinales, the splenius and suboccipital muscles are the only epaxial muscles.

<span class="mw-page-title-main">Work loop</span>

The work loop technique is used in muscle physiology to evaluate the mechanical work and power output of skeletal or cardiac muscle contractions via in vitro muscle testing of whole muscles, fiber bundles or single muscle fibers. This technique is primarily used for cyclical contractions such as cockroach walking., the rhythmic flapping of bird wings or the beating of heart ventricular muscle.

<span class="mw-page-title-main">Fish physiology</span> Scientific study of how the component parts of fish function together in the living fish

Fish physiology is the scientific study of how the component parts of fish function together in the living fish. It can be contrasted with fish anatomy, which is the study of the form or morphology of fishes. In practice, fish anatomy and physiology complement each other, the former dealing with the structure of a fish, its organs or component parts and how they are put together, such as might be observed on the dissecting table or under the microscope, and the later dealing with how those components function together in the living fish. For this, at first we need to know about their intestinal morphology.

<span class="mw-page-title-main">Webbed foot</span> Animal feet with non-pathogenic interdigital webbing

The webbed foot is a specialized limb with interdigital membranes (webbings) that aids in aquatic locomotion, present in a variety of tetrapod vertebrates. This adaptation is primarily found in semiaquatic species, and has convergently evolved many times across vertebrate taxa.

The Cambrian chordates are an extinct group of animals belonging to the phylum Chordata that lived during the Cambrian, between 485 and 538 million years ago. The first Cambrian chordate known is Pikaia gracilens, a lancelet-like animal from the Burgess Shale in British Columbia, Canada. The discoverer, Charles Doolittle Walcott, described it as a kind of worm (annelid) in 1911, but it was later identified as a chordate. Subsequent discoveries of other Cambrian fossils from the Burgess Shale in 1991, and from the Chengjiang biota of China in 1991, which were later found to be of chordates, several Cambrian chordates are known, with some fossils considered as putative chordates.

References

  1. 1 2 Nursall, J. R. (1956). "The Lateral Musculature and the Swimming of Fish". Proceedings of the Zoological Society of London. 126 (1): 127–144. doi:10.1111/j.1096-3642.1956.tb00429.x. ISSN   1469-7998.
  2. Walker, Warren F.; Noback, Charles R. (2021). "Muscular system". Access Science. doi:10.1036/1097-8542.440200.
  3. Van Leeuwen, J. L. (1999-12-01). "A mechanical analysis of myomere shape in fish". The Journal of Experimental Biology. 202 (Pt 23): 3405–3414. doi:10.1242/jeb.202.23.3405. ISSN   0022-0949. PMID   10562523.
  4. Flammang, B. E.; Lauder, G. V. (2009-01-15). "Caudal fin shape modulation and control during acceleration, braking and backing maneuvers in bluegill sunfish, Lepomis macrochirus". Journal of Experimental Biology. 212 (2): 277–286. doi: 10.1242/jeb.021360 . ISSN   0022-0949. PMID   19112147. S2CID   14529276.
  5. Lacalli, Thurston (2012-07-06). "The Middle Cambrian fossil Pikaia and the evolution of chordate swimming". EvoDevo. 3 (1): 12. doi: 10.1186/2041-9139-3-12 . ISSN   2041-9139. PMC   3390900 . PMID   22695332.
  6. Nair, Arjun; Azatian, Grigor; McHenry, Matthew J. (2015-12-01). "The kinematics of directional control in the fast start of zebrafish larvae". Journal of Experimental Biology. 218 (24): 3996–4004. doi: 10.1242/jeb.126292 . ISSN   0022-0949. PMID   26519511. S2CID   15224608.
  7. Delvolvé, I.; Bem, T.; Cabelguen, J. M. (August 1997). "Epaxial and limb muscle activity during swimming and terrestrial stepping in the adult newt, Pleurodeles waltl". Journal of Neurophysiology. 78 (2): 638–650. doi:10.1152/jn.1997.78.2.638. ISSN   0022-3077. PMID   9307101.
  8. Azizi, Emanuel; Gillis, Gary B.; Brainerd, Elizabeth L. (December 2002). "Morphology and mechanics of myosepta in a swimming salamander (Siren lacertina)". Comparative Biochemistry and Physiology. Part A, Molecular & Integrative Physiology. 133 (4): 967–978. doi:10.1016/s1095-6433(02)00223-4. ISSN   1095-6433. PMID   12485686.
  9. Gerardo De Iuliis, PhD (8 November 2010). The Dissection of Vertebrates | ScienceDirect. Elsevier Science. ISBN   9780123750600 . Retrieved 2021-11-22.