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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 Organ (anatomy), organs, Cell (biology), cells and cell organelles, using the methods of mechani ...
, Hill's muscle model refers to the 3-element model consisting of a contractile element (CE) in series with a lightly-damped elastic spring element (SE) and in parallel with lightly-damped elastic parallel element (PE). Within this model, the estimated force-velocity relation for the CE element is usually modeled by what is commonly called Hill's equation, which was based on careful experiments involving tetanized
muscle contraction Muscle contraction is the activation of Tension (physics), 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 musc ...
where various muscle loads and associated velocities were measured. They were derived by the famous
physiologist Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a subdiscipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out chemical and ...
Archibald Vivian Hill, who by 1938 when he introduced this model and equation had already won the Nobel Prize for Physiology. He continued to publish in this area through 1970. There are many forms of the basic "Hill-based" or "Hill-type" models, with hundreds of publications having used this model structure for experimental and simulation studies. Most major musculoskeletal simulation packages make use of this model.


AV Hill's force-velocity equation for tetanized muscle

This is a popular state equation applicable to
skeletal muscle Skeletal muscle (commonly referred to as muscle) is one of the three types of vertebrate muscle tissue, the others being cardiac muscle and smooth muscle. They are part of the somatic nervous system, voluntary muscular system and typically are a ...
that has been stimulated to show Tetanic contraction. It relates tension to velocity with regard to the internal
thermodynamics Thermodynamics is a branch of physics that deals with heat, Work (thermodynamics), work, and temperature, and their relation to energy, entropy, and the physical properties of matter and radiation. The behavior of these quantities is governed b ...
. 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 The van der Waals equation is a mathematical formula that describes the behavior of real gases. It is an equation of state that relates the pressure, volume, Avogadro's law, number of molecules, and temperature in a fluid. The equation modifies ...
, the former has units of energy
dissipation In thermodynamics, dissipation is the result of an irreversible process that affects a thermodynamic system. In a dissipative process, energy ( internal, bulk flow kinetic, or system potential) transforms from an initial form to a final form, wh ...
, while the latter has units of
energy Energy () is the physical quantity, quantitative physical property, property that is transferred to a physical body, body or to a physical system, recognizable in the performance of Work (thermodynamics), work and in the form of heat and l ...
. Hill's equation demonstrates that the relationship between F and v is
hyperbolic Hyperbolic may refer to: * of or pertaining to a hyperbola, a type of smooth curve lying in a plane in mathematics ** Hyperbolic geometry, a non-Euclidean geometry ** Hyperbolic functions, analogues of ordinary trigonometric functions, defined u ...
. 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 and can therefore be modeled as a fluid damper .


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 (or a non-linear 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, mathe ...
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 i ...
(SE) and another in parallel (PE). The active
force In physics, a force is an influence that can cause an Physical object, object to change its velocity unless counterbalanced by other forces. In mechanics, force makes ideas like 'pushing' or 'pulling' mathematically precise. Because the Magnitu ...
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 ...
and
myosin Myosins () are a Protein family, family of motor proteins (though most often protein complexes) best known for their roles in muscle contraction and in a wide range of other motility processes in eukaryotes. They are adenosine triphosphate, ATP- ...
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 striated muscle, Skeletal muscles are composed of tubular ...
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, a group of cells that are similar in structure, along with epithelial tissue, muscle tissue, and nervous tissue. It develops mostly from the mesenchyme, derived from the mesod ...
s (
fascia A fascia (; : fasciae or fascias; adjective fascial; ) is a generic term for macroscopic membranous bodily structures. Fasciae are classified as superficial, visceral or deep, and further designated according to their anatomical location. ...
, epimysium, perimysium and endomysium) 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 connective tissue, connects and surrounds or supports internal organs and bones, and includes muscle, tendons, ligaments, Adipose tissue, fat, fibrous tissue, Lymphatic vessel, lymph and blood vessels, fasciae, and synovial membranes.� ...
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 band of fibrous connective tissue, dense fibrous connective tissue that connects skeletal muscle, muscle to bone. It sends the mechanical forces of muscle contraction to the skeletal system, while withstanding tensi ...
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 both shear flow and strain lin ...
, 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 d ...
critically damped 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 * Ex ...
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 (physics), 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 musc ...


References

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