Hill's Muscle Model
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In
biomechanics Biomechanics is the study of the structure, function and motion of the mechanical aspects of biological systems, at any level from whole organisms to organs, cells and cell organelles, using the methods of mechanics. Biomechanics is a branch of ...
, Hill's muscle model refers to either Hill's equations for tetanized
muscle contraction 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 ...
or to the 3-element model. They were derived by the famous
physiologist Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical a ...
Archibald Vivian Hill.


Equation to tetanized muscle

This is a popular
state equation In physics, chemistry, and thermodynamics, an equation of state is a thermodynamic equation relating state variables, which describe the state of matter under a given set of physical conditions, such as pressure, volume, temperature, or interna ...
applicable to
skeletal muscle Skeletal muscles (commonly referred to as 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 muscl ...
that has been stimulated to show
Tetanic contraction A tetanic contraction (also called tetanized state, tetanus, or physiologic tetanus, the latter to differentiate from the disease called tetanus) is a sustained muscle contraction evoked when the motor nerve that innervates a skeletal muscle emits ...
. It relates
tension Tension may refer to: Science * Psychological stress * Tension (physics), a force related to the stretching of an object (the opposite of compression) * Tension (geology), a stress which stretches rocks in two opposite directions * Voltage or el ...
to velocity with regard to the internal
thermodynamics Thermodynamics is a branch of physics that deals with heat, work, and temperature, and their relation to energy, entropy, and the physical properties of matter and radiation. The behavior of these quantities is governed by the four laws of the ...
. The equation is :\left(v+b\right)(F+a) = b(F_0+a), \qquad (1) where * F is the tension (or load) in the muscle * v is the velocity of contraction * F_0 is the maximum isometric tension (or load) generated in the muscle * a coefficient of shortening heat * b=a\cdot v_0/F_0 * v_0 is the maximum velocity, when F=0 Although Hill's equation looks very much like the
van der Waals equation In chemistry and thermodynamics, the Van der Waals equation (or Van der Waals equation of state) is an equation of state which extends the ideal gas law to include the effects of interaction between molecules of a gas, as well as accounting for ...
, the former has units of energy
dissipation In thermodynamics, dissipation is the result of an irreversible process that takes place in homogeneous thermodynamic systems. In a dissipative process, energy (internal, bulk flow kinetic, or system potential) transforms from an initial form to a ...
, while the latter has units of
energy In physics, energy (from Ancient Greek: ἐνέργεια, ''enérgeia'', “activity”) is the quantitative property that is transferred to a body or to a physical system, recognizable in the performance of work and in the form of heat a ...
. Hill's equation demonstrates that the relationship between F and v is
hyperbolic Hyperbolic is an adjective describing something that resembles or pertains to a hyperbola (a curve), to hyperbole (an overstatement or exaggeration), or to hyperbolic geometry. The following phenomena are described as ''hyperbolic'' because they ...
. Therefore, the higher the load applied to the muscle, the lower the contraction velocity. Similarly, the higher the contraction velocity, the lower the tension in the muscle. This hyperbolic form has been found to fit the empirical constant only during isotonic contractions near resting length. The muscle tension decreases as the shortening velocity increases. This feature has been attributed to two main causes. The major appears to be the loss in tension as the cross bridges in the contractile element and then reform in a shortened condition. The second cause appears to be the fluid viscosity in both the contractile element and the connective tissue. Whichever the cause of loss of tension, it is a
viscous friction The viscosity of a fluid is a measure of its resistance to deformation at a given rate. For liquids, it corresponds to the informal concept of "thickness": for example, syrup has a higher viscosity than water. Viscosity quantifies the inter ...
and can therefore be modeled as a fluid
damper A damper is a device that deadens, restrains, or depresses. It may refer to: Music * Damper pedal, a device that mutes musical tones, particularly in stringed instruments * A mute for various brass instruments Structure * Damper (flow), a mechan ...
.


Three-element model

The three-element Hill muscle model is a representation of the muscle mechanical response. The model is constituted by a contractile element (CE) and two
non-linear In mathematics and science, a nonlinear system is a system in which the change of the output is not proportional to the change of the input. Nonlinear problems are of interest to engineers, biologists, physicists, mathematicians, and many other ...
spring elements, one in
series Series may refer to: People with the name * Caroline Series (born 1951), English mathematician, daughter of George Series * George Series (1920–1995), English physicist Arts, entertainment, and media Music * Series, the ordered sets used in ...
(SE) and another in parallel (PE). The active
force In physics, a force is an influence that can change the motion of an object. A force can cause an object with mass to change its velocity (e.g. moving from a state of rest), i.e., to accelerate. Force can also be described intuitively as a p ...
of the contractile element comes from the force generated by the
actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of over ...
and
myosin Myosins () are a superfamily of motor proteins best known for their roles in muscle contraction and in a wide range of other motility processes in eukaryotes. They are ATP-dependent and responsible for actin-based motility. The first myosin ...
cross-bridges at the
sarcomere A sarcomere (Greek σάρξ ''sarx'' "flesh", μέρος ''meros'' "part") is the smallest functional unit of striated muscle tissue. It is the repeating unit between two Z-lines. Skeletal muscles are composed of tubular muscle cells (called musc ...
level. It is fully extensible when inactive but capable of shortening when activated. The
connective tissue Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tiss ...
s (
fascia A fascia (; plural fasciae or fascias; adjective fascial; from Latin: "band") is a band or sheet of connective tissue, primarily collagen, beneath the skin that attaches to, stabilizes, encloses, and separates muscles and other internal organs. ...
,
epimysium Epimysium (plural ''epimysia'') (Greek ''epi-'' for on, upon, or above + Greek ''mys'' for muscle) is the fibrous tissue envelope that surrounds skeletal muscle. It is a layer of dense irregular connective tissue which ensheaths the entire muscle ...
,
perimysium Perimysium is a sheath of connective tissue that groups muscle fibers into bundles (anywhere between 10 and 100 or more) or fascicles. Studies of muscle physiology suggest that the perimysium plays a role in transmitting lateral contractile ...
and
endomysium The endomysium, meaning ''within the muscle'', is a wispy layer of areolar connective tissue that ensheaths each individual Skeletal muscle#Skeletal muscle fibers, muscle fiber, or muscle cell. It also contains capillaries and nerves. It overlies ...
) that surround the contractile element influences the muscle's force-length curve. The parallel element represents the passive force of these connective tissues and has a
soft tissue Soft tissue is all the tissue in the body that is not hardened by the processes of ossification or calcification such as bones and teeth. Soft tissue connects, surrounds or supports internal organs and bones, and includes muscle, tendons, ligam ...
mechanical behavior. The parallel element is responsible for the muscle passive behavior when it is stretched, even when the contractile element is not activated. The series element represents the
tendon A tendon or sinew is a tough, high-tensile-strength band of dense fibrous connective tissue that connects muscle to bone. It is able to transmit the mechanical forces of muscle contraction to the skeletal system without sacrificing its ability ...
and the intrinsic elasticity of the myofilaments. It also has a soft tissue response and provides energy storing mechanism. The net force-length characteristics of a muscle is a combination of the force-length characteristics of both active and passive elements. The forces in the contractile element, in the series element and in the parallel element, F^, F^ and F^, respectively, satisfy :F = F^+F^, \qquad F^=F^, \qquad (2) On the other hand, the muscle length L and the lengths L^, L^ and L^ of those elements satisfy :L = L^, \qquad L = L^+L^, \qquad (3) During isometric contractions the series elastic component is under tension and therefore is stretched a finite amount. Because the overall length of the muscle is kept constant, the stretching of the series element can only occur if there is an equal shortening of the contractile element itself. The forces in the parallel, series and contractile elements are defined by:F^(\lambda_f) = F_f^(\lambda_f), \qquad F^(\lambda^,\lambda^) = F_0f^(\lambda^,\lambda^), \qquad F^(\lambda^,\dot^,a) = F_0f_^(\lambda^)f_^(\dot^)a, \qquad (4)where \lambda_f, \lambda_, \lambda_ are strain measures for the different elements defined by:\lambda_ = \frac, \quad \lambda^ = \frac, \quad \lambda^ = \frac, \qquad (5)where L is the deformed muscle length and L^ is the deformed muscle length due to motion of the contractile element, both from equation (3). L_0 is the rest length of the muscle. \lambda_ can be split as \lambda_ = \lambda^\lambda^. The force term, F_, is the peak isometric muscle force and the functions f^, f^, f_L^, f_V^ are given by:\begin f^(\lambda_f) = \begin 2cA(\lambda_f-1)e^, & \lambda_f>1 \\ \text, & \text \end, & (6) \\ ptf^(\lambda^,\lambda^) = \begin 0.1(e^-1), & \lambda^\geq1 \\ \text, & \text \end, & (7) \\ ptf_L^(\lambda^) = \begin -4(\lambda^-1)^2+1, & 0.5\leq\lambda^\leq1.5 \\ \text, & \text \end, & (8) \\ ptf_V^(\dot^) = \begin \text, & \dot^<-10s^ \\ -\frac\arctan(-0.5\dot^)+1, & -10s^\leq\dot^\leq2s^\\ \frac+1, & \dot^>2s^ \end, & (9) \end where c, A are empirical constants. The function a(t) from equation (4) represents the muscle activation. It is defined based on the ordinary differential equation:\frac = \frac(1-a(t)u(t)+\frac(a_-a(t))(1-u(t))), \qquad (10)where \tau_, \tau_ are time constants related to rise and decay for muscle activation and a_ is a minimum bound, all determined from experiments. u(t) is the neural excitation that leads to muscle contraction.


Viscoelasticity

Muscles present
viscoelasticity In materials science and continuum mechanics, viscoelasticity is the property of materials that exhibit both viscous and elastic characteristics when undergoing deformation. Viscous materials, like water, resist shear flow and strain linearly wi ...
, therefore a viscous damper may be included in the model, when the dynamics of the
second-order Second-order may refer to: Mathematics * Second order approximation, an approximation that includes quadratic terms * Second-order arithmetic, an axiomatization allowing quantification of sets of numbers * Second-order differential equation, a di ...
critically damped Damping is an influence within or upon an oscillatory system that has the effect of reducing or preventing its oscillation. In physical systems, damping is produced by processes that dissipate the energy stored in the oscillation. Examples in ...
twitch is regarded. One common model for muscular viscosity is an
exponential Exponential may refer to any of several mathematical topics related to exponentiation, including: *Exponential function, also: **Matrix exponential, the matrix analogue to the above * Exponential decay, decrease at a rate proportional to value *Exp ...
form damper, where :F_ = k(\dot_)^a, \qquad (11) is added to the model's global equation, whose k and a are constants.


See also

*
Muscle contraction 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 ...


References

{{DEFAULTSORT:Hill's Muscle Model Biomechanics Equations Exercise physiology