Pennate muscle

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Rectus femoris
Rectus femoris.png
Muscles of the iliac and anterior femoral regions. (Rectus femoris highlighted in red.)
Details
Origin Anterior inferior iliac spine and the exterior surface of the bony ridge which forms the iliac portion of the acetabulum
Insertion Inserts into the patellar tendon as one of the four quadriceps muscles
Artery Lateral femoral circumflex artery
Nerve Femoral nerve
Actions Knee extension; hip flexion
Antagonist Hamstring
Identifiers
TA98 A04.0.00.016
TA2 1989
FMA 74993
Anatomical terms of muscle

A pennate or pinnate muscle (also called a penniform muscle) is a type of skeletal muscle with fascicles that attach obliquely (in a slanting position) to its tendon. This type of muscle generally allows higher force production but a smaller range of motion. [1] [2] When a muscle contracts and shortens, the pennation angle increases. [3]

Contents

Etymology

The term "pennate" comes from the Latin pinnātus ("feathered, winged"), from pinna ("feather, wing").

Types of pennate muscle

Figure 1 Pennate muscle fiber arrangements: A, unipennate; B, bipennate; C, multipennate. Blue: anatomical cross-section. Green: physiological cross-section. Fiederung.svg
Figure 1 Pennate muscle fiber arrangements: A, unipennate; B, bipennate; C, multipennate. Blue: anatomical cross-section. Green: physiological cross-section.

In skeletal muscle tissue, 10-100 endomysium-sheathed muscle fibers are organized into perimysium-wrapped bundles known as fascicles. Each muscle is composed of a number of fascicles grouped together by a sleeve of connective tissue, known as an epimysium. In a pennate muscle, aponeuroses run along each side of the muscle and attach to the tendon. The fascicles attach to the aponeuroses and form an angle (the pennation angle) to the load axis of the muscle. If all the fascicles are on the same side of the tendon, the pennate muscle is called unipennate (Fig. 1A). Examples of this include certain muscles in the hand. If there are fascicles on both sides of the central tendon, the pennate muscle is called bipennate (Fig. 1B). The rectus femoris, a large muscle in the quadriceps, is typical. If the central tendon branches within a pennate muscle, the muscle is called multipennate (Fig. 1C), as seen in the deltoid muscle in the shoulder.

Consequences of pennate muscle architecture

Physiological cross sectional area (PCSA)

Pennation angle Pennation angle of fibers in pennate muscle.png
Pennation angle

One advantage of pennate muscles is that more muscle fibers can be packed in parallel, thus allowing the muscle to produce more force, although the fiber angle to the direction of action means that the maximum force in that direction is somewhat less than the maximum force in the fiber direction. [4] [5] The muscle cross sectional area (blue line in figure 1, also known as anatomical cross section area, or ACSA) does not accurately represent the number of muscle fibers in the muscle. A better estimate is provided by the total area of the cross sections perpendicular to the muscle fibers (green lines in figure 1). This measure is known as the physiological cross sectional area (PCSA), and is commonly calculated and defined by the following formula (an alternative definition is provided in the main article): [6] [7] [8]

where ρ is the density of the muscle:

PCSA increases with pennation angle, and with muscle length. In a pennate muscle, PCSA is always larger than ACSA. In a non-pennate muscle, it coincides with ACSA.

Relationship between PCSA and muscle force

The total force exerted by the fibers along their oblique direction is proportional to PCSA. If the specific tension of the muscle fibers is known (force exerted by the fibers per unit of PCSA), it can be computed as follows: [9]

However, only a component of that force can be used to pull the tendon in the desired direction. This component, which is the true muscle force (also called tendon force [8] ), is exerted along the direction of action of the muscle: [8]

The other component, orthogonal to the direction of action of the muscle (Orthogonal force = Total force × sinΦ) is not exerted on the tendon, but simply squeezes the muscle, by pulling its aponeuroses toward each other.

Notice that, although it is practically convenient to compute PCSA based on volume or mass and fiber length, PCSA (and therefore the total fiber force, which is proportional to PCSA) is not proportional to muscle mass or fiber length alone. Namely, the maximum (tetanic) force of a muscle fiber simply depends on its thickness (cross-section area) and type. By no means it depends on its mass or length alone. For instance, when muscle mass increases due to physical development during childhood, this may be only due to an increase in length of the muscle fibers, with no change in fiber thickness (PCSA) or fiber type. In this case, an increase in mass does not produce an increase in force.

Lower velocity of shortening

In a pennate muscle, as a consequence of their arrangement, fibers are shorter than they would be if they ran from one end of the muscle to the other. This implies that each fiber is composed of a smaller number N of sarcomeres in series. Moreover, the larger the pennation angle is, the shorter are the fibers.

The speed at which a muscle fiber can shorten is partly determined by the length of the muscle fiber (i.e., by N). Thus, a muscle with a large pennation angle will contract more slowly than a similar muscle with a smaller pennation angle.

Figure 2 Architectural gear ratio Gray368.png
Figure 2 Architectural gear ratio

Architectural gear ratio

Architectural gear ratio, also called anatomical gear ratio, (AGR) is a feature of pennate muscle defined by the ratio between the longitudinal strain of the muscle and muscle fiber strain. It is sometimes also defined as the ratio between muscle-shortening velocity and fiber-shortening velocity: [10]

AGR = εxf

where εx = longitudinal strain (or muscle-shortening velocity) and εf is fiber strain (or fiber-shortening velocity). [10]

It was originally thought that the distance between aponeuroses did not change during the contraction of a pennate muscle, [5] thus requiring the fibers to rotate as they shorten. However, recent work has shown this is false, and that the degree of fiber angle change varies under different loading conditions. This dynamic gearing automatically shifts in order to produce either maximal velocity under low loads or maximal force under high loads. [10] [11]

Related Research Articles

<span class="mw-page-title-main">Tendon</span> Type of tissue that connects muscle to bone

A tendon or sinew is a tough band of dense fibrous connective tissue that connects muscle to bone. It sends the mechanical forces of muscle contraction to the skeletal system, while withstanding tension.

<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.

<span class="mw-page-title-main">Striated muscle tissue</span> Muscle tissue with repeating functional units called sarcomeres

Striated muscle tissue is a muscle tissue that features repeating functional units called sarcomeres. The presence of sarcomeres manifests as a series of bands visible along the muscle fibers, which is responsible for the striated appearance observed in microscopic images of this tissue. There are two types of striated muscle:

<span class="mw-page-title-main">Aponeurosis</span> Tissue which connects muscles to other organs

An aponeurosis is a flattened tendon by which muscle attaches to bone or fascia. Aponeuroses exhibit an ordered arrangement of collagen fibres, thus attaining high tensile strength in a particular direction while being vulnerable to tensional or shear forces in other directions. They have a shiny, whitish-silvery color, are histologically similar to tendons, and are very sparingly supplied with blood vessels and nerves. When dissected, aponeuroses are papery and peel off by sections. The primary regions with thick aponeuroses are in the ventral abdominal region, the dorsal lumbar region, the ventriculus in birds, and the palmar (palms) and plantar (soles) regions.

<span class="mw-page-title-main">Muscle contraction</span> Activation of tension-generating sites in muscle

Muscle contraction is the activation of tension-generating sites within muscle cells. In physiology, muscle contraction does not necessarily mean muscle shortening because muscle tension can be produced without changes in muscle length, such as when holding something heavy in the same position. The termination of muscle contraction is followed by muscle relaxation, which is a return of the muscle fibers to their low tension-generating state.

<span class="mw-page-title-main">Muscular hydrostat</span> Body part type that consists mainly of muscles with no skeletal support

A muscular hydrostat is a biological structure found in animals. It is used to manipulate items or to move its host about and consists mainly of muscles with no skeletal support. It performs its hydraulic movement without fluid in a separate compartment, as in a hydrostatic skeleton.

<span class="mw-page-title-main">Soleus muscle</span> Powerful muscle in the back part of the lower leg

In humans and some other mammals, the soleus is a powerful muscle in the back part of the lower leg. It runs from just below the knee to the heel and is involved in standing and walking. It is closely connected to the gastrocnemius muscle, and some anatomists consider this combination to be a single muscle, the triceps surae. Its name is derived from the Latin word "solea", meaning "sandal".

A stretch-shortening cycle (SSC) is an active stretch of a muscle followed by an immediate shortening of that same muscle.

<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.

Preflexes are the latent capacities in the musculoskeletal system that auto-stabilize movements through the use of the nonlinear visco-elastic properties of muscles when they contract. The term "preflex" for such a zero-delay, intrinsic feedback loop was coined by Loeb. Unlike stabilization methods using neurons, such as reflexes and higher brain control, a preflex happens with minimal time delay; however, it only stabilizes the main movements of the musculoskeletal system.

<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.

Role of skin in locomotion describes how the integumentary system is involved in locomotion. Typically the integumentary system can be thought of as skin, however the integumentary system also includes the segmented exoskeleton in arthropods and feathers of birds. The primary role of the integumentary system is to provide protection for the body. However, the structure of the skin has evolved to aid animals in their different modes of locomotion. Soft bodied animals such as starfish rely on the arrangement of the fibers in their tube feet for movement. Eels, snakes, and fish use their skin like an external tendon to generate the propulsive forces need for undulatory locomotion. Vertebrates that fly, glide, and parachute also have a characteristic fiber arrangements of their flight membranes that allows for the skin to maintain its structural integrity during the stress and strain experienced during flight.

<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">Physiological cross-sectional area</span> Area perpendicular to fiber direction

In muscle physiology, physiological cross-sectional area (PCSA) is the area of the cross section of a muscle perpendicular to its fibers, generally at its largest point. It is typically used to describe the contraction properties of pennate muscles. It is not the same as the anatomical cross-sectional area (ACSA), which is the area of the crossection of a muscle perpendicular to its longitudinal axis. In a non-pennate muscle the fibers are parallel to the longitudinal axis, and therefore PCSA and ACSA coincide.

In vitro muscle testing is a method used to characterize properties of living muscle tissue after removing it from an organism, which allows more extensive and precise quantification of its properties than in vivo testing. In vitro muscle testing has provided the bulk of scientific knowledge of muscle structure and physiology, and how both relate to organismal performance. Stem cell research relies on in vitro muscle testing to establish sole muscle cell function and its individual behavior apart from muscle cells in the presence of nonmuscle cells seen in in vitro studies.

<span class="mw-page-title-main">Lateral force transmission in skeletal muscle</span>

A key component in lateral force transmission in skeletal muscle is the extracellular matrix (ECM). Skeletal muscle is a complex biological material that is composed of muscle fibers and an ECM consisting of the epimysium, perimysium, and endomysium. It can be described as a collagen fiber-reinforced composite. The ECM has at least three functions: (1) to provide a framework binding muscle fibers together and ensure their proper alignment, (2) to transmit the forces, either from active muscle contraction or ones passively imposed on it, and (3) providing lubricated surfaces between muscle fibers and bundles enabling the muscle to change shape. The mechanical properties of skeletal muscle depend on both the properties of muscle fibers and the ECM, and the interaction between the two. Contractile forces are transmitted laterally within intramuscular connective tissue to the epimysium and then to the tendon. Due to the nature of skeletal muscle, direct measurements are not possible, but many indirect studies and analyses have shown that the ECM is an important part of force transmission during muscle contraction.

<span class="mw-page-title-main">Architectural gear ratio</span> Ratio between muscle-shortening velocity and fiber-shortening velocity

Architectural gear ratio, also called anatomical gear ratio (AGR) is a feature of pennate muscle defined by the ratio between the longitudinal strain of the muscle and muscle fiber strain. It is sometimes also defined as the ratio between muscle-shortening velocity and fiber-shortening velocity.

<span class="mw-page-title-main">Limitations of animal running speed</span> Factors determining maximum running speed in animals

Limitations of animal running speed provides an overview of how various factors determine the maximum running speed. Some terrestrial animals are built for achieving extremely high speeds, such as the cheetah, pronghorn, race horse and greyhound, while humans can train to achieve high sprint speeds. There is no single determinant of maximum running speed: however, certain factors stand out against others and have been investigated in both animals and humans. These factors include: Muscle moment arms, foot morphology, muscle architecture, and muscle fiber type. Each factor contributes to the ground reaction force (GRF) and foot contact time of which the changes to increase maximal speed are not well understood across all species.

Muscle architecture is the physical arrangement of muscle fibers at the macroscopic level that determines a muscle's mechanical function. There are several different muscle architecture types including: parallel, pennate and hydrostats. Force production and gearing vary depending on the different muscle parameters such as muscle length, fiber length, pennation angle, and the physiological cross-sectional area (PCSA).

<span class="mw-page-title-main">Anatomical terms of muscle</span> Muscles terminology

Anatomical terminology is used to uniquely describe aspects of skeletal muscle, cardiac muscle, and smooth muscle such as their actions, structure, size, and location.

References

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