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Muscle is a soft tissue found in most animals, and is one of the four basic animal tissues, along with nervous tissue, epithelium, and connective tissue. Muscle cells contain protein filaments called myofilaments of actin and myosin that slide past one another, producing a contraction that changes both the length and the shape of the cell. Muscles function to produce force and motion. They are primarily responsible for maintaining and changing posture, locomotion, as well as movement of internal organs, such as the contraction of the heart and the movement of food through the digestive system via peristalsis.

Muscle tissue is derived from the embryonic mesodermal germ layer in a process known as myogenesis. There are three types of muscle, of which skeletal and cardiac muscle are striated and smooth muscle is not. Muscle action can be classified as being either voluntary or involuntary. Cardiac and smooth muscles contract without conscious thought and are termed involuntary, whereas the skeletal muscles contract upon command. Skeletal muscles in turn can be divided into fast and slow twitch fibers.

Muscles are predominantly powered by the oxidation of fats and carbohydrates, but anaerobic chemical reactions are also used, particularly by fast twitch fibers. These chemical reactions produce adenosine triphosphate (ATP) molecules that are used to power the movement of the myosin heads.

The term muscle is derived from the Latin musculus meaning "little mouse" perhaps because of the shape of certain skeletal muscles or because contracting muscles look like mice moving under the skin.

The anatomy of muscles includes gross anatomy, which comprises all the muscles of an organism, and microanatomy, which comprises the structure of a single muscle.

Muscle tissue is a soft tissue, and is one of the four fundamental types of tissue present in animals. There are three types of muscle tissue recognized in vertebrates:

Cardiac and skeletal muscles are "striated" in that they contain sarcomeres that are packed into highly regular arrangements of bundles; the myofibrils of smooth muscle cells are not arranged in sarcomeres and so are not striated. While the sarcomeres in skeletal muscles are arranged in regular, parallel bundles, cardiac muscle sarcomeres connect at branching, irregular angles (called intercalated discs). Striated muscle contracts and relaxes in short, intense bursts, whereas smooth muscle sustains longer or even near-permanent contractions.

The muscle fibers embedded in skeletal muscle are relatively classified into a spectrum of types given their morphological and physiological properties. Given a certain assortment of these properties, muscle fibers are categorized as slow-twitch (low force, slowly fatiguing fibers), fast twitch (high force, rapidly fatiguing fibers), or somewhere in between those two types (i.e. intermediate fibers). Some of the defining morphological and physiological properties used for the categorization of muscle fibers include: the number of mitochondria contained in the fiber, the amount of glycolytic, lipolytic, and other cellular respiration enzymes, M and Z band characteristics, energy source (i.e. glycogen or fat), histology color, and contraction speed and duration. There is no standard procedure for classifying muscle fiber types, the properties chosen for classification depends on the particular muscle. For example, the properties used for distinguishing fast, intermediate, and slow muscle fibers can be different for invertebrate flight and jump muscle. To further complicate this classification scheme, the mitochondria content and other morphological properties within a muscle fiber can change in a tsetse fly with exercise and age.

The density of mammalian skeletal muscle tissue is about 1.06 kg/liter. This can be contrasted with the density of adipose tissue (fat), which is 0.9196 kg/liter. This makes muscle tissue approximately 15% denser than fat tissue.

Skeletal muscles are sheathed by a tough layer of connective tissue called the epimysium. The epimysium anchors muscle tissue to tendons at each end, where the epimysium becomes thicker and collagenous. It also protects muscles from friction against other muscles and bones. Within the epimysium are multiple bundles called fascicles, each of which contains 10 to 100 or more muscle fibers collectively sheathed by a perimysium. Besides surrounding each fascicle, the perimysium is a pathway for nerves and the flow of blood within the muscle. The threadlike muscle fibers are the individual muscle cells, and each cell is encased within its own endomysium of collagen fibers. Thus, the overall muscle consists of fibers (cells) that are bundled into fascicles, which are themselves grouped together to form muscles. At each level of bundling, a collagenous membrane surrounds the bundle, and these membranes support muscle function both by resisting passive stretching of the tissue and by distributing forces applied to the muscle. Scattered throughout the muscles are muscle spindles that provide sensory feedback information to the central nervous system. (This grouping structure is analogous to the organization of nerves which uses epineurium, perineurium, and endoneurium).

This same bundles-within-bundles structure is replicated within the muscle cells. Within the cells of the muscle are myofibrils, which themselves are bundles of protein filaments. The term "myofibril" should not be confused with "myofiber", which is a simply another name for a muscle cell. Myofibrils are complex strands of several kinds of protein filaments organized together into repeating units called sarcomeres. The striated appearance of both skeletal and cardiac muscle results from the regular pattern of sarcomeres within their cells. Although both of these types of muscle contain sarcomeres, the fibers in cardiac muscle are typically branched to form a network. Cardiac muscle cells are interconnected by intercalated discs, giving that tissue the appearance of a syncytium.

The filaments in a sarcomere are composed of actin and myosin.

The gross anatomy of a muscle is the most important indicator of its role in the body. There is an important distinction seen between pennate muscles and other muscles. In most muscles, all the fibers are oriented in the same direction, running in a line from the origin to the insertion. However, In pennate muscles, the individual fibers are oriented at an angle relative to the line of action, attaching to the origin and insertion tendons at each end. Because the contracting fibers are pulling at an angle to the overall action of the muscle, the change in length is smaller, but this same orientation allows for more fibers (thus more force) in a muscle of a given size. Pennate muscles are usually found where their length change is less important than maximum force, such as the rectus femoris.

Skeletal muscle is arranged in discrete muscles, an example of which is the biceps. The tough, fibrous epimysium of skeletal muscle is both connected to and continuous with the tendons. In turn, the tendons connect to the periosteum layer surrounding the bones, permitting the transfer of force from the muscles to the skeleton. Together, these fibrous layers, along with tendons and ligaments, constitute the deep fascia of the body.

The muscular system consists of all the muscles present in a single body. There are approximately 650 skeletal muscles in the human body, but an exact number is difficult to define. The difficulty lies partly in the fact that different sources group the muscles differently and partly in that some muscles, such as the palmaris longus, are not always present.

A muscular slip is a narrow length of muscle that acts to augment a larger muscle or muscles.

The muscular system is one component of the musculoskeletal system, which includes not only the muscles but also the bones, joints, tendons, and other structures that permit movement.

All muscles are derived from paraxial mesoderm. The paraxial mesoderm is divided along the embryo's length into somites, corresponding to the segmentation of the body (most obviously seen in the vertebral column. Each somite has 3 divisions, sclerotome (which forms vertebrae), dermatome (which forms skin), and myotome (which forms muscle). The myotome is divided into two sections, the epimere and hypomere, which form epaxial and hypaxial muscles, respectively. The only epaxial muscles in humans are the erector spinae and small intervertebral muscles, and are innervated by the dorsal rami of the spinal nerves. All other muscles, including those of the limbs are hypaxial, and innervated by the ventral rami of the spinal nerves.

During development, myoblasts (muscle progenitor cells) either remain in the somite to form muscles associated with the vertebral column or migrate out into the body to form all other muscles. Myoblast migration is preceded by the formation of connective tissue frameworks, usually formed from the somatic lateral plate mesoderm. Myoblasts follow chemical signals to the appropriate locations, where they fuse into elongate skeletal muscle cells.

The three types of muscle (skeletal, cardiac and smooth) have significant differences. However, all three use the movement of actin against myosin to create contraction. In skeletal muscle, contraction is stimulated by electrical impulses transmitted by the nerves, the motor neurons in particular. Cardiac and smooth muscle contractions are stimulated by internal pacemaker cells which regularly contract, and propagate contractions to other muscle cells they are in contact with. All skeletal muscle and many smooth muscle contractions are facilitated by the neurotransmitter acetylcholine.

The action a muscle generates is determined by the origin and insertion locations. The cross-sectional area of a muscle (rather than volume or length) determines the amount of force it can generate by defining the number of sarcomeres which can operate in parallel. Each skeletal muscle contains long units called myofibrils, and each myofibril is a chain of sarcomeres. Since contraction occurs at the same time for all connected sarcomeres in a muscles cell, these chains of sarcomeres shorten together, thus shortening the muscle fiber, resulting in overall length change. The amount of force applied to the external environment is determined by lever mechanics, specifically the ratio of in-lever to out-lever. For example, moving the insertion point of the biceps more distally on the radius (farther from the joint of rotation) would increase the force generated during flexion (and, as a result, the maximum weight lifted in this movement), but decrease the maximum speed of flexion. Moving the insertion point proximally (closer to the joint of rotation) would result in decreased force but increased velocity. This can be most easily seen by comparing the limb of a mole to a horse—in the former, the insertion point is positioned to maximize force (for digging), while in the latter, the insertion point is positioned to maximize speed (for running).

The efferent leg of the peripheral nervous system is responsible for conveying commands to the muscles and glands, and is ultimately responsible for voluntary movement. Nerves move muscles in response to voluntary and autonomic (involuntary) signals from the brain. Deep muscles, superficial muscles, muscles of the face and internal muscles all correspond with dedicated regions in the primary motor cortex of the brain, directly anterior to the central sulcus that divides the frontal and parietal lobes.

In addition, muscles react to reflexive nerve stimuli that do not always send signals all the way to the brain. In this case, the signal from the afferent fiber does not reach the brain, but produces the reflexive movement by direct connections with the efferent nerves in the spine. However, the majority of muscle activity is volitional, and the result of complex interactions between various areas of the brain.

Nerves that control skeletal muscles in mammals correspond with neuron groups along the primary motor cortex of the brain's cerebral cortex. Commands are routed through the basal ganglia and are modified by input from the cerebellum before being relayed through the pyramidal tract to the spinal cord and from there to the motor end plate at the muscles. Along the way, feedback, such as that of the extrapyramidal system contribute signals to influence muscle tone and response.

Deeper muscles such as those involved in posture often are controlled from nuclei in the brain stem and basal ganglia.

In skeletal muscles, muscle spindles convey information about the degree of muscle length and stretch to the central nervous system to assist in maintaining posture and joint position. The sense of where our bodies are in space is called proprioception, the perception of body awareness, the "unconscious" awareness of where the various regions of the body are located at any one time. Several areas in the brain coordinate movement and position with the feedback information gained from proprioception. The cerebellum and red nucleus in particular continuously sample position against movement and make minor corrections to assure smooth motion.[citation needed]

Muscular activity accounts for much of the body's energy consumption. All muscle cells produce adenosine triphosphate (ATP) molecules which are used to power the movement of the myosin heads. Muscles have a short-term store of energy in the form of creatine phosphate which is generated from ATP and can regenerate ATP when needed with creatine kinase. Muscles also keep a storage form of glucose in the form of glycogen. Glycogen can be rapidly converted to glucose when energy is required for sustained, powerful contractions. Within the voluntary skeletal muscles, the glucose molecule can be metabolized anaerobically in a process called glycolysis which produces two ATP and two lactic acid molecules in the process (note that in aerobic conditions, lactate is not formed; instead pyruvate is formed and transmitted through the citric acid cycle). Muscle cells also contain globules of fat, which are used for energy during aerobic exercise. The aerobic energy systems take longer to produce the ATP and reach peak efficiency, and requires many more biochemical steps, but produces significantly more ATP than anaerobic glycolysis. Cardiac muscle on the other hand, can readily consume any of the three macronutrients (protein, glucose and fat) aerobically without a 'warm up' period and always extracts the maximum ATP yield from any molecule involved. The heart, liver and red blood cells will also consume lactic acid produced and excreted by skeletal muscles during exercise.

At rest, skeletal muscle consumes 54.4 kJ/kg (13.0 kcal/kg) per day. This is larger than adipose tissue (fat) at 18.8 kJ/kg (4.5 kcal/kg), and bone at 9.6 kJ/kg (2.3 kcal/kg).

The efficiency of human muscle has been measured (in the context of rowing and cycling) at 18% to 26%. The efficiency is defined as the ratio of mechanical work output to the total metabolic cost, as can be calculated from oxygen consumption. This low efficiency is the result of about 40% efficiency of generating ATP from food energy, losses in converting energy from ATP into mechanical work inside the muscle, and mechanical losses inside the body. The latter two losses are dependent on the type of exercise and the type of muscle fibers being used (fast-twitch or slow-twitch). For an overall efficiency of 20 percent, one watt of mechanical power is equivalent to 4.3 kcal per hour. For example, one manufacturer of rowing equipment calibrates its rowing ergometer to count burned calories as equal to four times the actual mechanical work, plus 300 kcal per hour, this amounts to about 20 percent efficiency at 250 watts of mechanical output. The mechanical energy output of a cyclic contraction can depend upon many factors, including activation timing, muscle strain trajectory, and rates of force rise & decay. These can be synthesized experimentally using work loop analysis.

Muscle is a result of three factors that overlap: physiological strength (muscle size, cross sectional area, available crossbridging, responses to training), neurological strength (how strong or weak is the signal that tells the muscle to contract), and mechanical strength (muscle's force angle on the lever, moment arm length, joint capabilities).[citation needed]

Vertebrate muscle typically produces approximately 25–33 N (5.6–7.4 lbf) of force per square centimeter of muscle cross-sectional area when isometric and at optimal length. Some invertebrate muscles, such as in crab claws, have much longer sarcomeres than vertebrates, resulting in many more sites for actin and myosin to bind and thus much greater force per square centimeter at the cost of much slower speed. The force generated by a contraction can be measured non-invasively using either mechanomyography or phonomyography, be measured in vivo using tendon strain (if a prominent tendon is present), or be measured directly using more invasive methods.

The strength of any given muscle, in terms of force exerted on the skeleton, depends upon length, shortening speed, cross sectional area, pennation, sarcomere length, myosin isoforms, and neural activation of motor units. Significant reductions in muscle strength can indicate underlying pathology, with the chart at right used as a guide.

Since three factors affect muscular strength simultaneously and muscles never work individually, it is misleading to compare strength in individual muscles, and state that one is the "strongest". But below are several muscles whose strength is noteworthy for different reasons.

Exercise is often recommended as a means of improving motor skills, fitness, muscle and bone strength, and joint function. Exercise has several effects upon muscles, connective tissue, bone, and the nerves that stimulate the muscles. One such effect is muscle hypertrophy, an increase in size of muscle due to an increase in the number of muscle fibers or cross-sectional area of myofibrils. The degree of hypertrophy and other exercise induced changes in muscle depends on the intensity and duration of exercise.

Generally, there are two types of exercise regimes, aerobic and anaerobic. Aerobic exercise (e.g. marathons) involves low intensity, but long duration activities during which, the muscles used are below their maximal contraction strength. Aerobic activities rely on the aerobic respiration (i.e. citric acid cycle and electron transport chain) for metabolic energy by consuming fat, protein carbohydrates, and oxygen. Muscles involved in aerobic exercises contain a higher percentage of Type I (or slow-twitch) muscle fibers, which primarily contain mitochondrial and oxidation enzymes associated with aerobic respiration. On the contrary, anaerobic exercise is associated with short duration, but high intensity exercise (e.g. sprinting and weight lifting). The anaerobic activities predominately use Type II, fast-twitch, muscle fibers. Type II muscle fibers rely on glucogenesis for energy during anaerobic exercise. During anaerobic exercise, type II fibers consume little oxygen, protein and fat, produces large amounts of lactic acid and are fatigable. Many exercises are partially aerobic and anaerobic; for example, soccer and rock climbing.

The presence of lactic acid has an inhibitory effect on ATP generation within the muscle. It can even stop ATP production if the intracellular concentration becomes too high. However, endurance training mitigates the buildup of lactic acid through increased capillarization and myoglobin. This increases the ability to remove waste products, like lactic acid, out of the muscles in order to not impair muscle function. Once moved out of muscles, lactic acid can be used by other muscles or body tissues as a source of energy, or transported to the liver where it is converted back to pyruvate. In addition to increasing the level of lactic acid, strenuous exercise results in the loss of potassium ions in muscle. This may facilitate the recovery of muscle function by protecting against fatigue .

Delayed onset muscle soreness is pain or discomfort that may be felt one to three days after exercising and generally subsides two to three days after which. Once thought to be caused by lactic acid build-up, a more recent theory is that it is caused by tiny tears in the muscle fibers caused by eccentric contraction, or unaccustomed training levels. Since lactic acid disperses fairly rapidly, it could not explain pain experienced days after exercise.

Independent of strength and performance measures, muscles can be induced to grow larger by a number of factors, including hormone signaling, developmental factors, strength training, and disease. Contrary to popular belief, the number of muscle fibres cannot be increased through exercise. Instead, muscles grow larger through a combination of muscle cell growth as new protein filaments are added along with additional mass provided by undifferentiated satellite cells alongside the existing muscle cells.

Biological factors such as age and hormone levels can affect muscle hypertrophy. During puberty in males, hypertrophy occurs at an accelerated rate as the levels of growth-stimulating hormones produced by the body increase. Natural hypertrophy normally stops at full growth in the late teens. As testosterone is one of the body's major growth hormones, on average, men find hypertrophy much easier to achieve than women. Taking additional testosterone or other anabolic steroids will increase muscular hypertrophy.

Muscular, spinal and neural factors all affect muscle building. Sometimes a person may notice an increase in strength in a given muscle even though only its opposite has been subject to exercise, such as when a bodybuilder finds her left biceps stronger after completing a regimen focusing only on the right biceps. This phenomenon is called cross education.[citation needed]

During ordinary living activities, between 1 and 2 percent of muscle is broken down and rebuilt daily. Inactivity and starvation in mammals lead to atrophy of skeletal muscle, a decrease in muscle mass that may be accompanied by a smaller number and size of the muscle cells as well as lower protein content. Muscle atrophy may also result from the natural aging process or from disease.

In humans, prolonged periods of immobilization, as in the cases of bed rest or astronauts flying in space, are known to result in muscle weakening and atrophy. Atrophy is of particular interest to the manned spaceflight community, because the weightlessness experienced in spaceflight results is a loss of as much as 30% of mass in some muscles. Such consequences are also noted in small hibernating mammals like the golden-mantled ground squirrels and brown bats.

During aging, there is a gradual decrease in the ability to maintain skeletal muscle function and mass, known as sarcopenia. The exact cause of sarcopenia is unknown, but it may be due to a combination of the gradual failure in the "satellite cells" that help to regenerate skeletal muscle fibers, and a decrease in sensitivity to or the availability of critical secreted growth factors that are necessary to maintain muscle mass and satellite cell survival. Sarcopenia is a normal aspect of aging, and is not actually a disease state yet can be linked to many injuries in the elderly population as well as decreasing quality of life.

There are also many diseases and conditions that cause muscle atrophy. Examples include cancer and AIDS, which induce a body wasting syndrome called cachexia. Other syndromes or conditions that can induce skeletal muscle atrophy are congestive heart disease and some diseases of the liver.

Neuromuscular diseases are those that affect the muscles and/or their nervous control. In general, problems with nervous control can cause spasticity or paralysis, depending on the location and nature of the problem. A large proportion of neurological disorders, ranging from cerebrovascular accident (stroke) and Parkinson's disease to Creutzfeldt–Jakob disease, can lead to problems with movement or motor coordination.

Symptoms of muscle diseases may include weakness, spasticity, myoclonus and myalgia. Diagnostic procedures that may reveal muscular disorders include testing creatine kinase levels in the blood and electromyography (measuring electrical activity in muscles). In some cases, muscle biopsy may be done to identify a myopathy, as well as genetic testing to identify DNA abnormalities associated with specific myopathies and dystrophies.

A non-invasive elastography technique that measures muscle noise is undergoing experimentation to provide a way of monitoring neuromuscular disease. The sound produced by a muscle comes from the shortening of actomyosin filaments along the axis of the muscle. During contraction, the muscle shortens along its longitudinal axis and expands across the transverse axis, producing vibrations at the surface.

The evolutionary origin of muscle cells in animals is highly debated. One view is that muscle cells evolved once and thus all muscles cells have a single common ancestor. Another view is that muscles cells evolved more than once and any morphological or structural similarities are due to convergent evolution and genes that predate the evolution of muscle and even the mesoderm—the germ layer that gives rise to vertebrate muscle cells.

Schmid and Seipel argue that the origin of muscle cells is a monophyletic trait that occurred concurrently with the development of the digestive and nervous systems of all animals and that this origin can be traced to a single metazoan ancestor in which muscle cells are present. They argue that molecular and morphological similarities between the muscles cells in cnidaria and ctenophora are similar enough to those of bilaterians that there would be one ancestor in metazoans from which muscle cells derive. In this case, Schmid and Seipel argue that the last common ancestor of bilateria, ctenophora, and cnidaria was a triploblast or an organism with three germ layers and that diploblasty, meaning an organism with two germ layers, evolved secondarily due to their observation of the lack of mesoderm or muscle found in most cnidarians and ctenophores. By comparing the morphology of cnidarians and ctenophores to bilaterians, Schmid and Seipel were able to conclude that there were myoblast-like structures in the tentacles and gut of some species of cnidarians and in the tentacles of ctenophores. Since this is a structure unique to muscle cells, these scientists determined based on the data collected by their peers that this is a marker for striated muscles similar to that observed in bilaterians. The authors also remark that the muscle cells found in cnidarians and ctenophores are often contests due to the origin of these muscle cells being the ectoderm rather than the mesoderm or mesendoderm. The origin of true muscles cells is argued by others to be the endoderm portion of the mesoderm and the endoderm. However, Schmid and Seipel counter this skepticism about whether or not the muscle cells found in ctenophores and cnidarians are true muscle cells by considering that cnidarians develop through a medusa stage and polyp stage. They observe that in the hydrozoan medusa stage there is a layer of cells that separate from the distal side of the ectoderm to form the striated muscle cells in a way that seems similar to that of the mesoderm and call this third separated layer of cells the ectocodon. They also argue that not all muscle cells are derived from the mesendoderm in bilaterians with key examples being that in both the eye muscles of vertebrates and the muscles of spiralians these cells derive from the ectodermal mesoderm rather than the endodermal mesoderm. Furthermore, Schmid and Seipel argue that since myogenesis does occur in cnidarians with the help of molecular regulatory elements found in the specification of muscles cells in bilaterians that there is evidence for a single origin for striated muscle.

In contrast to this argument for a single origin of muscle cells, Steinmetz et al. argue that molecular markers such as the myosin II protein used to determine this single origin of striated muscle actually predate the formation of muscle cells. This author uses an example of the contractile elements present in the porifera or sponges that do truly lack this striated muscle containing this protein. Furthermore, Steinmetz et al. present evidence for a polyphyletic origin of striated muscle cell development through their analysis of morphological and molecular markers that are present in bilaterians and absent in cnidarians, ctenophores, and bilaterians. Steimetz et al. showed that the traditional morphological and regulatory markers such as actin, the ability to couple myosin side chains phosphorylation to higher concentrations of the positive concentrations of calcium, and other MyHC elements are present in all metazoans not just the organisms that have been shown to have muscle cells. Thus, the usage of any of these structural or regulatory elements in determining whether or not the muscle cells of the cnidarians and ctenophores are similar enough to the muscle cells of the bilaterians to confirm a single lineage is questionable according to Steinmetz et al. Furthermore, Steinmetz et al. explain that the orthologues of the MyHc genes that have been used to hypothesize the origin of striated muscle occurred through a gene duplication event that predates the first true muscle cells (meaning striated muscle), and they show that the MyHc genes are present in the sponges that have contractile elements but no true muscle cells. Furthermore, Steinmetz et all showed that the localization of this duplicated set of genes that serve both the function of facilitating the formation of striated muscle genes and cell regulation and movement genes were already separated into striated myhc and non-muscle myhc. This separation of the duplicated set of genes is shown through the localization of the striated myhc to the contractile vacuole in sponges while the non-muscle myhc was more diffusely expressed during developmental cell shape and change. Steinmetz et al. found a similar pattern of localization in cnidarians with except with the cnidarian N. vectensis having this striated muscle marker present in the smooth muscle of the digestive track. Thus, Steinmetz et al. argue that the pleisiomorphic trait of the separated orthologues of myhc cannot be used to determine the monophylogeny of muscle, and additionally argue that the presence of a striated muscle marker in the smooth muscle of this cnidarian shows a fundamentally different mechanism of muscle cell development and structure in cnidarians.

Steinmetz et al. continue to argue for multiple origins of striated muscle in the metazoans by explaining that a key set of genes used to form the troponin complex for muscle regulation and formation in bilaterians is missing from the cnidarians and ctenophores, and of 47 structural and regulatory proteins observed, Steinmetz et al. were not able to find even on unique striated muscle cell protein that was expressed in both cnidarians and bilaterians. Furthermore, the Z-disc seemed to have evolved differently even within bilaterians and there is a great deal diversity of proteins developed even between this clade, showing a large degree of radiation for muscle cells. Through this divergence of the Z-disc, Steimetz et al. argue that there are only four common protein components that were present in all bilaterians muscle ancestors and that of these for necessary Z-disc components only an actin protein that they have already argued is an uninformative marker through its pleisiomorphic state is present in cnidarians. Through further molecular marker testing, Steinmetz et al. observe that non-bilaterians lack many regulatory and structural components necessary for bilaterians muscle formation and do not find any unique set of proteins to both bilaterians and cnidarians and ctenophores that are not present in earlier, more primitive animals such as the sponges and amoebozoans. Through this analysis the authors conclude that due to the lack of elements that bilaterians muscles are dependent on for structure and usage, nonbilaterian muscles must be of a different origin with a different set regulatory and structural proteins.

In another take on the argument, Andrikou and Arnone use the newly available data on gene regulatory networks to look at how the hierarchy of genes and morphogens and other mechanism of tissue specification diverge and are similar among early deuterostomes and protostomes. By understanding not only what genes are present in all bilaterians but also the time and place of deployment of these genes, Andrikou and Arnone discuss a deeper understanding of the evolution of myogenesis.

In their paper Andrikou and Arnone argue that to truly understand the evolution of muscle cells the function of transcriptional regulators must be understood in the context of other external and internal interactions. Through their analysis, Andrikou and Arnone found that there were conserved orthologues of the gene regulatory network in both invertebrate bilaterians and in cnidarians. They argue that having this common, general regulatory circuit allowed for a high degree of divergence from a single well functioning network. Andrikou and Arnone found that the orthologues of genes found in vertebrates had been changed through different types of structural mutations in the invertebrate deuterostomes and protostomes, and they argue that these structural changes in the genes allowed for a large divergence of muscle function and muscle formation in these species. Andrikou and Arnone were able to recognize not only any difference due to mutation in the genes found in vertebrates and invertebrates but also the integration of species specific genes that could also cause divergence from the original gene regulatory network function. Thus, although a common muscle patterning system has been determined, they argue that this could be due to a more ancestral gene regulatory network being coopted several times across lineages with additional genes and mutations causing very divergent development of muscles. Thus it seems that myogenic patterning framework may be an ancestral trait. However, Andrikou and Arnone explain that the basic muscle patterning structure must also be considered in combination with the cis regulatory elements present at different times during development. In contrast with the high level of gene family apparatuses structure, Andrikou and Arnone found that the cis regulatory elements were not well conserved both in time and place in the network which could show a large degree of divergence in the formation of muscle cells. Through this analysis, it seems that the myogenic GRN is an ancestral GRN with actual changes in myogenic function and structure possibly being linked to later coopts of genes at different times and places.

Evolutionarily, specialized forms of skeletal and cardiac muscles predated the divergence of the vertebrate/arthropod evolutionary line. This indicates that these types of muscle developed in a common ancestor sometime before 700 million years ago (mya). Vertebrate smooth muscle was found to have evolved independently from the skeletal and cardiac muscle types.

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RT @_DIONSTARR_: 👑 @Uncut_Muscle 🍆💦
33
RT @MuscleBulls: Sometimes it is time to milk the Big Bull's massive cock and play with his huge balls 🐷🐷🐷🐷 ✅ #mu…
87
RT @thealphasmen: Retweet if you like huge muscle hunks
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@erebonbon maybe if he's a super fit muscle moo all of that Big Huge went right to his dong
0
RT @oppaisquad: By tight I mean the muscle won’t stop flexing please grandma tsunade heal it!
140
RT @NippleMuscleCum: Mmm muscle jap daddy pt.2
150
RT @baitomist: " เมษาแล้วยังไง เมใจให้กันก็พอแล้ว " ว่ะ 5555555 🤣 N : How cool you can be, brooo Z : I'm looking forward to see your body…
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RT @timtales_com: Chinese/German hardfucker @TianTaoXXX meets Indonesian muscle bottom @FabioTobaXXX Don't miss the epic cum shot. It's o…
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RT @jenniferatank98: You don't need the gym to get ripped! This amazing push-up board gives you the muscle definition that you desire!
13
RT @DatabaseVip: Achil get fucked by parkingman. Tan skin & muscle top from indonesia. So hot video! Full duration 44 minutes. Join and get…
322
@nobugard he has a lot lean muscle in his chest but his limbs are more gentle imo
0
RT @rifaijju_09: Ngentot Chinese muscle power Bottom. New account,account lama di hack. Minat bisa order. 👉🏽wa +6282261066396 👉🏽line rifaij…
674
@nobiodu @TreasuredWretch @ikeamadi @Elohor_T @tetoda @daeetoni @benny_obasi @Eliaswal @oremeyi @TheLAfamily He did…
0
BREAKING: Eden Hazard has suffered another muscle injury and is a doubt for Real Madrid's Champions League clash wi…
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Real Madrid confirm Eden Hazard has suffered a muscle injury. He is expected to be out for at least four weeks, pe…
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⚪️ Real Madrid confirm that Eden Hazard has suffered a muscle injury in training. #UCL
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RT @brfootball: Real Madrid confirm Eden Hazard has suffered a muscle injury. He is expected to be out for at least four weeks, per @HLNin…
1284
Stop calling your car a muscle car if it’s not V8
0
#AmberDeLuca #bodybuilder #fbb #ripped #bicep #vascular #abs #athlete #tan #strongwoman #strongwomen #photoshoot…
0
is Josh King alright? Barely moved a muscle since he came on. Christ.
0
RT @liangjjgz: 173:我的大雞吧被肌肉小受緊致的菊花夾射 My big dick cummed with a muscle guy's flower tightness infibulating. 我穿著短白襪操他 邊乾邊玩他敏感乳頭 再用腳撫摸著他 沒充血…
477
( ˘ω˘)クソネミ 毎日毎日こんなだけど なんで夜中の3時に目が覚めてしまったわけ? そこで目覚めなくて4時半くらいに目覚めてればこんな眠くならなかったはずだ
0
RT @jenniferatank98: You don't need the gym to get ripped! This amazing push-up board gives you the muscle definition that you desire!
9
RT @PoojaLRao1: "It is hard to believe that this tiny, fist-sized, two-hundred-and-fifty-gram muscle is the core of human existence. Nearly…
6
RT @ChaudhuryAlok: @Breathedreamgo Please read d farm laws. These middlemen were exploitators of real farmers causing hundreds of suicides…
16
I want a muscle car
0
RT @Manyu_adult: 야생의 가로막구리가 나타났다! #Obstagoon #furry #pokemon #muscle
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RT @goal: BREAKING: Eden Hazard has suffered another muscle injury and is a doubt for Real Madrid's Champions League clash with Atalanta, G…
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RT @blancoslive: ℹ Hazard had a similar injury in August 2019 in the same muscle which kept him out for 3 games. #rmlive 🇧🇪 🔴 — COPE
3
@ManLikeFrane1 @itsAliChour @Dannn777 Milik is still a shadow of a player he was before he got that ACL injury.. T…
0
RT @jakedanieloffic: Full Muscle Worship! What would you do to be @caseycooperxxx ? 😈 Come see the full version NOW🔥
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RT @theQueerPig: Hot muscle man @ManuelSkyexxx fucks @FelixODairxxx 's twink hole raw in @dadcreep 's new video! 🍆 + GET 50% OFF for LIFE…
8
$1 $NCT...SOON!🚀
0
RT @bloominanna: @cbrennansports @usatodaysports Males who identify as trans MUST be excluded from women’s & girls sport because they are M…
7
RT @seegasmworlds: (273S) Slim Muscle Straight Guy 2 (=267S)(4/4) Why was he wiggling his toes whenever I looked away! While I was editing…
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RT @Bricklayer1001: I only enjoy taking cock from older muscle daddies, dunno why, but it does it for me
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@sakakenblog おはけん✨ 今日もぶち上げ!! 燃えてくぜ🔥
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RT @seegasmworlds: (273S) Slim Muscle Straight Guy 2 (=267S) (*2/4) We took turns sucking each other's cock while I stimulated his prostat…
414
RT @cryptofrog202: I was thinking $AAVE eventual listing of $AMPL would be boolish but HOLY FUQ, reading what $YLD
1
RT @terongkeras: anjir.. pagi cerah lgsg isepin konto1 mas muscle di semak2 Download app video LIVE streaming lokal disini
131
@eminor3rd @RuudyEB @ACLU There is no data. what’s the average height and weight of a women; and a man What’s the…
0
Checkout latest News post : Does muscle weigh more than fat? No, building muscle can actually help you lose weight…
0
RT @suamiganteng: lagi sange minta di sodok nih laki muscle jadi ngac3ng hehe Download app video kumpulan lokal JOIN playstore disini
71
China Priddy the new Worlds Great Superpower unless The US shows its Military muscle.
0
RT @Beautiful_Won: @6godwolo his body, his choice. As if we didn't go through this shit with him when he was with MX. Remember the muscle…
2
@NOBUO_muscle のぶさん💫✨おはにょん ダンペル💪💪💪💪💪 のぶさんには欠かせないですもんね😁 今日も楽しんでいきましょう❣️
0
RT @Rubysole628: Do you like petite girls with a bit of muscle? :)
326
RT @emas_online: Secondary analysis of a randomized trial (TULIP study) finds unsupervised behavioural and pelvic floor muscle training pro…
2
Quem é karol conká? Para o crossfiter, o muscle up
0
More speculative risk factors: decreased mental chronometry and speed and repetitive eccentric contraction of muscle.
0
As China gains more muscle, we’re tied down with division at home. It’s time to step it up.
76
That is a damn fine automobile... I love that I’ve been able to spend 15+ years in ‘60’s muscle-cars. #walker
618
BJP govt failed in getting India's financial defaulters like Nirav Modi, Lalit Modi & Mallya extradited but is fili…
124
Flexing so hard I might pull a muscle but I just keep stunning
0
@buttonalltheway Packed on muscle too
0
What would you do if you had my ass in your hands? Lick it 👅 or spank it 🏓? Follow & sub for🆓on OnlyFans! 🆕content…
0
RT @BdfJjl: For #FossilFriday it is Tail-less week here, so an excellent opportunity to post the 2.74 metre high Leedsichthys tail NHMUK P.…
6
RT @wartraktor: My bodybuilder friend @SamanthaTheMilf sucks dick like a boss! #milf #muscle #blowjob #hardcore #sex
2
RT @EveryDayBulge: 🍌 @MrOfficialRAY #bulge #bigbulge @vegas_muscle @maspalo75 @ethan_reyes @estebaneneko @gayladzUK
9
@EveryDayBulge @MrOfficialRAY @vegas_muscle @maspalo75 @ethan_reyes @EstebanEneko @GayLadzUK Wooooooooow negrote 💋🍆🍼🍆💋
0
RT @TheDedicated_1: Philly boxer who I been sucking for 2 years about 23 years old finally let me record, he hmu on ig for some head. You k…
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Je crois que je me suis froissé un muscle je fais quoi
0
RT @PriaaBerotot: == OPEN MEMBERSHIP == VIDEO MUSCLE AND BODYBUILDER LOKAL (1000++ VIDEOS) + ACCESS TO DOWNLOAD VIDEO FROM HUNK-CH / TRANC…
2
RT @MensHealthUK: S&C coach Rock Gullickson talks MH through his tips and tricks for big lifts that will get your body on the offensive htt…
1
Time to hustle for that muscle 💪🏽🏋🏻‍♀️💦🔥#FitnessFriday #Fitness
0
RT @markwulfgar: A nice change in the usual stuff people commission me. @VelRyder and @shadowwolf0590 (featured in this) always request me…
21
RT @nsfw_horizon: Well you heard him #nsfw #nsfwart #muscle #gayart #sketch
2
@Ryukun_active おはりゅう✨ 良さそう!寝付き悪いんでやってみます!やるぞ🔥
0
RT @rj_coli: 🔥 BABANG MUSCLE 🔥 PAPA MUDA MENGGODA Ayo download versi full ➡️ ⬅️ . . . . .
38
Into The Woods - #YoshiKawasakiXXX - #gaysex #Asian #Bareback #Fitlads #Jerking #Muscle #PublicSex #Rimming…
0
Senate impeachment trial now in recess because Democrats are trying to figure out a way out of the predicament they…
1362
According to a 15-year study, wearing bras can accelerate breast sagging as they prevent supportive muscle from gro…
330
Nigerians are being killed by organized ethnic violence in Oyo, but the government’s security muscle is flexing at unarmed protesters.
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ゆうじくん可愛いすぎて……我が子?…孫???箱推しすぎて供給過多
0
RT @MusclesRuth: #onlyfans #femalemuscle #femaebodybuiding #amazona #amazing #muscle #sexymuscle #bigmuscle #huge #hugemuscle #
1
RT @teddy_bryce: ✨🥺 I’m really sorry about my phat muscle pussy 🥺✨ 💖50% OFF ONLYFANS ALL FEBRUARY💖
245
RT @EditiEffiong: Nigerians are being killed by organized ethnic violence in Oyo, but the government’s security muscle is flexing at unarme…
615
【#筋肉娘】逞しく鍛え抜かれた #女の子 の体、エロすぎてヤバい【三次元】 #筋肉美女 #筋肉 #筋骨隆々 #逞しい女 #ゴツい女 #太もも #ムチムチ #musclegirl #musculargirl #muscle
1
@frmstbz keknya muscle injury
0
Penat tu tak sangat but badan aku dah sampai limit. Muscle soreness. Kena rest
0
RT @buttsohot: MINUM PEJU brondong MUSCLE part 2, SEXY ABIS 🔥bangun TIDUR langsung NGEWE BOOLku💦 Mau ORDER KOLEKSIku? Fast respon bisa C…
484
RT @Napalm_femuscle: 【#筋肉娘】逞しく鍛え抜かれた #女の子 の体、エロすぎてヤバい【三次元】 #筋肉美女 #筋肉 #筋骨隆々 #逞しい女 #ゴツい女 #太もも #ムチムチ #musclegirl #mu…
1
マン汁で金玉の裏までびしょびしょ侍
0
大阪から帰ります!半年振りに来れて楽しかった。今回は大阪住みの友人とりんくうタウン→泉南イオン→鳳→西成→天王寺→心斎橋を巡りました🚶‍♀️ また夏来ます!次は大阪住みのFFさんとも会えるといいな
0
RT @EditiEffiong: Nigerians are being killed by organized ethnic violence in Oyo, but the government’s security muscle is flexing at unarme…
615
RT @sallylepage: If you are an adult aged between 25 and 70, there is a 1 in 4 chance you will develop long Covid. Let me say that again.…
26
@femoral_muscle ㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋㅋ아패라서그럼
0
Behind da Scenes of His Majesty, Sire_Shon Vday night! Link in Bio! 💦💕🍑💯 #fans #onlyfans #model #wet #muscle…
0
RT @ElphusMk: Feel better than you have in years with herbalife products 💚 Gain weight 💚 Build muscle 💚 Get fit/boost energy levels 🚚Deli…
2
@badendido Not too bad. Been stuck in bed for 4 days. Pulled a muscle in my back. But my best mate has just dropped…
0
RT @oshitono: HANDSOME ASIAN MUSCLE
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@lalaleni_a ralat deh, triple muscle idiots + kak taiki 🥺
0
RT @A_mounds: And on the next episode of “It’s Gonna Hurt” ft FullMetalTwunk @StarryTwunk 🍠🤕🥲 #bareback #bigdick #raw #bubblebutt #black #…
477
So I fell asleep on the couch last night and tweaked the muscle in my hip/ass and I been walking funny all day and…
0
Dudes will grow one (1) muscle and make it their whole personality
0
RT @muscleflexxx: Who’s ready for a shoot with @lewissurv and his monster tool! Support your favourite muscle guys and alphas .... >>>>>…
70
RT @fetishkink25: Yes, Sir: Jon Darra Submits to Muscle-Stud, Max Konnor Muscle bound Max Konnor has Jon Darra at his mercy. Blindfolded w…
45
RT @oshitono: CUTIE KOREAN MUSCLE
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@HardNotice What if we're more curve than muscle?
0
RT @BenedictDunant: The army has made intimidating displays of muscle at several major protest sites in Yangon, but many demonstrators aren…
67
止まない雨はないし生えない陰毛もない
0
RT @scallysex: Bane and Seth buck naked football fun in the California sun. 🎥▶️ … #NudeBeach #NakedMuscleMenTweet…
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RT @LustRashad: Big Muscle Mood OF: JFF:
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@ashleypeeeps @soundguyeric Healthy relationships are abnormal to modern gals. You'll save yourself a world of hear…
0
RT @bustforme: Anon muscle guys breeding my hole just hits different 😈
332
Lets drool over american muscle cars from the 60’s
0
RT @sangaytsheltrim: Age is just a number @BeingSalmanKhan. Very excited to start my countdown , 13 weeks out for #radhe #eid2021 #salmankh…
161
The Ultimate Forearm Workout: The 5 Best Forearm Exercises for Popeye Arms | If you want to know the best forearm workout for increasing muscle mass and grip strength, then you want to read this article.
0
Testosterone is a sex hormone that plays important roles in the body. In men, it’s thought to regulate sex drive (libido), bone mass, fat distribution, muscle mass and strength, and the production of red blood cells and sperm. A small amount of circulating testosterone is converted to estradiol, a form of oestrogen. As men age, they often make less testosterone, and so they produce less estradiol as well.
1
İlgili kaslar: Yan karın kasları. Dik durun ve üst bedeninizi yavaşça sağa doğru bükün. Aynı hareketi diğer tarafta tekrarlayın.
197
Sayısız egzersiz programı bulunuyor ama bunlardan en etkililerinden biri "Tibet'in 5 İncisi" adı verilen set. Bu eski yöntem sadece tüm kasları çalıştırmakla kalmıyor vücut aktivitelerini de normal hale getiriyor. 10 Dakikada Tüm Kasları Çalıştıran 5 Tibet Egzersizi Sadece
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DEVE DURUŞU
2
OMUZ GERME
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BOYUN ÖN KASLARI GERME
1
SIRT KASLARI
1
Weight lifting schedule for 12 weeks for building muscle for women. fitness motivation, #healthy #fitness #fitspo
1
Find out how to gain muscle mass naturally.
0
Weight lifting schedule for 12 weeks for building muscle for women. fitness motivation, #healthy #fitness #fitspo
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The Ultimate Forearm Workout: The 5 Best Forearm Exercises for Popeye Arms | If you want to know the best forearm workout for increasing muscle mass and grip strength, then you want to read this article.
1
Testosterone is a sex hormone that plays important roles in the body. In men, it’s thought to regulate sex drive (libido), bone mass, fat distribution, muscle mass and strength, and the production of red blood cells and sperm. A small amount of circulating testosterone is converted to estradiol, a form of oestrogen. As men age, they often make less testosterone, and so they produce less estradiol as well.
0
muscle Pinterest Ideas
  • muscle fitness
  • muscle men
  • muscle cars
  • muscle growth
  • muscle building workouts
  • muscle anatomy
  • muscle building foods
  • muscle building women
Feb 13, 2021 16:05
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Still wear my dad’s old AirForce PT shirt, Minus the sleeves, RIP ...
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Feb 12, 2021 00:46
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채색이 제일 어려워 ㅠ#illustration #drawing #muscle #muscleman #beard #underwater #man #일러스트 #드로잉 #그림 #만화 #근육남 #수염 #속옷 #남자 #肌肉 #肌肉男 #插图
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#man #illustration #muscleman #beard #muscle
Feb 13, 2021 16:05
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Feb 13, 2021 16:05
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RTJ 🔥___intro into the gym🎧
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Feb 13, 2021 16:05
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Jun 21, 2020 06:21
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練習を任せて頂けてるので 毎日練習メニューを考えています!  こっちが適当になれば、やる側も適当になると思っているので心を込めて考えています  #仙女 #センダイガールズプロレスリング #sendaigirlspro #女子プロレス #プロレス #prowrestling #prowrestler #筋肉女子 #ぎゅん #道場 #training #warmingup #muscle #musclegirls
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#muscle #warmingup #prowrestler #training #musclegirls
Jun 21, 2020 06:20
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Dessa creatina que ele cheirou eu não chego perto nem da tampa kkkkk . . . . @toguro @mansaomaromba_ @scarpelly_oficial @lilian.lemos . . . #maromba #bodybuilder #sup #humor #fitness #bodybuilding #standup #memes #treino #standupcomedy #meme #gym #standuppaddle #humorlatino #esmagaquecresce #workout #standuppaddleboard #funny #vemmonstro #muscle #comedy #humornegro #nopainnogain #fitnessmotivation #standupjetski #lol #13memo #bodybuilderlifestyle #standupcomedian #humorous
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#memes #fitness #bodybuilderlifestyle #fitnessmotivation #esmagaquecresce
Jun 21, 2020 06:19
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Can u solve this? 👆#challenge #challenger #fitness #challengeaccepted #motivation #dodge #mopar #workout #fit #challenges #challengeyourself #hardwork #goals #challengerfam #hananeelkhader #model #healthy #follow #determination #challenger_brigade #challenge_hananeelkhader #inspiration #moparornocar #srt #challengeď #bodybuilding #challenged #muscle #train #physique
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#dodge #motivation #physique #challenge #challenger
Jun 21, 2020 06:19
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The hungrier you are the faster you win 💪🏼☠ . . . . . . . . . . . . .#a6000 #fitness #bodybuilding #bodystransformation #homeworkout #nogymneeded #muscle #musclebuilding #fitnessmotivation #summerbody #workout #portrait #portraitphotography #selfportrait #sonyalpha #photo #photography #selfie #bicepworkout #shoulderworkout #me #instagram
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#musclebuilding #photo #portrait #bicepworkout #instagram
Jun 21, 2020 06:20
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@badmatchup #gymlife #vegasfitfam #weightloss #fatburner #planetfitness #personaltrainer #eastlasvegas #clarkcountynv #vegasculture #calisthenicmovement #vegasfit #lasvegassun #calisthenics #barstarzz #lvbarstarzz #barlife #barfit #stevia #gymrat #squats #increasemuscle #vgk #thebigguy #muscle #musclegoals #fitlife #healthy #lifetimefitness #yeezy #buildmuscle
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#calisthenicmovement #vgk #gymlife #personaltrainer #squats