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Le fibre muscolari:

classificazioni e proprietà
funzionali

Marsilio Saccavini
UOC Degenza di Medicina Riabilitativa
TREVISO
Structural organisation of skeletal muscle
Fibre type classification
Myosin molecule
Essential features of MHCs

• Molecular weight: ~200 kD


• Two MHCs for each myosin molecule with
tails coiled up to form an -helix
• Each MHC comprises a globular head (“the
motor region”) containing ATPase.
• Exists in at least 7 different variants (isoforms)
mATPase-BASED FIBRE TYPES
• (Engel 1962): fibres can be classified into type I &II
(Dubowitz & Pearse nomenclature, 1960) according to the
pH sensitivity of myosinAPSase
– Slow (type I) fibres: show low mATPase activity in
alkaline conditions (pH 10.4) but high activity in acid
conditions (pH 4.3 and 4.6)  slow fibres are alkali
labile and acid stable and stain poorly in alkaline pH
– Fast (type II) fibres: alkali stable and acid labile, stain
strongly in alkaline pH (10.4 and 9.7). Subtypes: IIA,
IIB, IIC. C fibres are precursors of A and B (Brooke and
Kaiser, 1970,74), other recent subtypes: IIAB, IIAC, IC.
• Total of 7 fibre types: I, IC, IIC, IIAC, IIA, IIAB, IIB
Myosin ATPase-based
fibre classification
MHC-BASED FIBRE TYPES
• Antigenic differences between myosins enables
immunohistochemical differenciation of muscle fibres
(Billiter et al. 1980)
• mATPase-based fibre classification correlates with myosin
composition (Billiter 1980):
– Type I fibres contain only slow myosin
– Type IIA and IIB fibres contain only fast myosins
– Type C fibres contain both
• MHC portion of myosin contains the site for ATP
hydrolysis (Wagner and Giniger, 1981)
MHC expression & fibre type
Immunoistochemical and electrophoretic techniques (Staron, 1991) proved
that the 7 mATPase-based fibre types correspond to a specific MHC profile:

FIBRE TYPE MHC expression


Type I MHC-I (one type MHC)

Type IIA MHC-IIa ,,


Type IIB* MHC-IIb ,,
Type IIAB MHC-IIa & MHC-IIb (two types MHC)

Type C (IC,IIC, IIAC) MHC-I & MHC-IIa ,,


( present in 0-5% of most adult muscles)
* (MHC similar to that of rat type IIX fibres )
Agreement between mATPase activity and
MHC expressions (Sant’Ana Pereira et al. 1994)
• mATPase activity and MHC expression exist in a continuum
(Klitgaard et al. 1990; Sant’Ana Pereira et al. 1995)
Essential features of MLCs
MLC: Molecular mass 17-23 kDa

• Attached to each MHC are two MLCs


– One regulatory isoform (MLC2s and MLC2f)
– One alkali (MLC1s, MLC1f, MLC3f)
– So, altogether there are 5 MLCs
• Fast skeletal muscle contains three isoforms:
– MLC1f, MLC3f and MLC2f
• Slow skeletal muscle contains two isoforms:
– MLC1s and MLC2s
Functional role of MLCs
Contractile properties of human muscle fibres
F-V properties of slow and fast fibres

FACTS
Vo type 1 is ~1/10 Vo type 2B
Vo type 2A is ~1/3 Vo type 2B
Force, velocity and muscle power
Power-velocity features of
human slow and fast fibres
FACTS
• Wmax type 1 is ~1/10
of type 2B fibres and
1/4 of type 2A
• Vopt type 1 is ~1/4 of
type 2B and ~1/2 of
type 2A fibres
Optimal velocity of MHC isoforms
Optimum velocity in vivo
• Slow and fast fibres shorten at different velocities,
but both shorten at their optimum velocity i.e. at
the velocity at which, in vitro, they develop
maximum power
• Muscles are selectively recruited to allow
movement in the whole range of physiological
speeds to optimise power and efficiency of
movement
(L.C. Rome et al., Nature 335:824-827, 1988)
Spontaneous pedalling rate
Power-Velocity characteristics of slow,
fast fibres and mixed muscle
Cycling power in relation to
type I and II fibre power
Maximum muscle power vs.
performance duration
Sustainable Power and Energy Sources

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