Power output from a flight muscle of the bumblebee Bombus terrestris. II. Characterization of the parameters affecting power output

1997 ◽  
Vol 200 (8) ◽  
pp. 1227-1239 ◽  
Author(s):  
R Josephson
Keyword(s):  
Power Output ◽  
Oscillation Frequency ◽  
Flight Muscle ◽  
Bombus Terrestris ◽  
Dynamic Stiffness ◽  
Muscle Length ◽  
Storage Device ◽  
Phase Lag ◽  
Work Output ◽  
Optimum Frequency

1. Length-tension relationships and work output were investigated in the intact, dorso-ventral flight muscle of the bumblebee Bombus terrestris. The muscle is an asynchronous muscle. Like other asynchronous flight muscles, it has high resting stiffness and produces relatively low active force in response to tetanic stimulation. 2. The muscle shows shortening deactivation and stretch activation, properties that result in delayed force changes in response to step changes in length, a phase lag between force and length during imposed sinusoidal strain and, under appropriate conditions, positive work output during oscillatory length change. 3. Work loops were used to quantify work output by the muscle during imposed sinusoidal oscillation. The curves relating net work per cycle with muscle length, oscillatory strain and oscillatory frequency were all roughly bell-shaped. The work-length curve was narrow. The optimum strain for net work per cycle was approximately 3 %, which is probably somewhat greater than the strain experienced by the muscle in an intact, flying bumblebee. The optimum frequency for net work output per cycle was 63 Hz (30 °C). The optimum frequency for power output was 73 Hz, which agrees well with the normal wing stroke frequency if allowance is made for the elevated temperature (approximately 40 °C) in the thorax of a flying bumblebee. The optimal strain for work output was not strongly dependent on oscillation frequency. 4. Resilience (that is the work output during shortening/work input during lengthening) for unstimulated muscle and dynamic stiffness (=stress/strain) for both stimulated and unstimulated muscles were determined using the strain (3 %) and oscillation frequency (64 Hz) which maximized work output in stimulated muscles. Unstimulated muscle is a good energy storage device. Its resilience increased with increasing muscle length (and increasing resting force) to reach values of over 90 %. The dynamic stiffness of both stimulated and unstimulated muscles increased with muscle length, but the increase was relatively greater in unstimulated muscle, and at long muscle lengths the stiffness of unstimulated muscle exceeded that of stimulated muscle. Effectively, dynamic stiffness is reduced by stimulation! This is taken as indicating that part of the stiffness in an unstimulated muscle reflects structures, possibly attached cross bridges, whose properties change upon stimulation.

Power output by an asynchronous flight muscle from a beetle

2000 ◽  
Vol 203 (17) ◽  
pp. 2667-2689 ◽  
Author(s):  
R.K. Josephson ◽  
J.G. Malamud ◽  
D.R. Stokes
Keyword(s):  
Power Output ◽  
Flight Muscle ◽  
Muscle Length ◽  
Mechanical Power ◽  
Effect Of Temperature ◽  
Muscle Temperature ◽  
Work Output ◽  
Stroke Frequency ◽  
Mechanical Power Output ◽  
Wing Stroke

The basalar muscle of the beetle Cotinus mutabilis is a large, fibrillar flight muscle composed of approximately 90 fibers. The paired basalars together make up approximately one-third of the mass of the power muscles of flight. Changes in twitch force with changing stimulus intensity indicated that a basalar muscle is innervated by at least five excitatory axons and at least one inhibitory axon. The muscle is an asynchronous muscle; during normal oscillatory operation there is not a 1:1 relationship between muscle action potentials and contractions. During tethered flight, the wing-stroke frequency was approximately 80 Hz, and the action potential frequency in individual motor units was approximately 20 Hz. As in other asynchronous muscles that have been examined, the basalar is characterized by high passive tension, low tetanic force and long twitch duration. Mechanical power output from the basalar muscle during imposed, sinusoidal strain was measured by the work-loop technique. Work output varied with strain amplitude, strain frequency, the muscle length upon which the strain was superimposed, muscle temperature and stimulation frequency. When other variables were at optimal values, the optimal strain for work per cycle was approximately 5%, the optimal frequency for work per cycle approximately 50 Hz and the optimal frequency for mechanical power output 60–80 Hz. Optimal strain decreased with increasing cycle frequency and increased with muscle temperature. The curve relating work output and strain was narrow. At frequencies approximating those of flight, the width of the work versus strain curve, measured at half-maximal work, was 5% of the resting muscle length. The optimal muscle length for work output was shorter than that at which twitch and tetanic tension were maximal. Optimal muscle length decreased with increasing strain. The curve relating work output and muscle length, like that for work versus strain, was narrow, with a half-width of approximately 3 % at the normal flight frequency. Increasing the frequency with which the muscle was stimulated increased power output up to a plateau, reached at approximately 100 Hz stimulation frequency (at 35 degrees C). The low lift generated by animals during tethered flight is consistent with the low frequency of muscle action potentials in motor units of the wing muscles. The optimal oscillatory frequency for work per cycle increased with muscle temperature over the temperature range tested (25–40 degrees C). When cycle frequency was held constant, the work per cycle rose to an optimum with increasing temperature and then declined. We propose that there is a temperature optimum for work output because increasing temperature increases the shortening velocity of the muscle, which increases the rate of positive work output during shortening, but also decreases the durations of the stretch activation and shortening deactivation that underlie positive work output, the effect of temperature on shortening velocity being dominant at lower temperatures and the effect of temperature on the time course of activation and deactivation being dominant at higher temperatures. The average wing-stroke frequency during free flight was 94 Hz, and the thoracic temperature was 35 degrees C. The mechanical power output at the measured values of wing-stroke frequency and thoracic temperature during flight, and at optimal muscle length and strain, averaged 127 W kg(−1)muscle, with a maximum value of 200 W kg(−1). The power output from this asynchronous flight muscle was approximately twice that measured with similar techniques from synchronous flight muscle of insects, supporting the hypothesis that asynchronous operation has been favored by evolution in flight systems of different insect groups because it allows greater power output at the high contraction frequencies of flight.

Power output from a flight muscle of the bumblebee Bombus terrestris. III. Power during simulated flight

1997 ◽  
Vol 200 (8) ◽  
pp. 1241-1246 ◽  
Author(s):  
R Josephson
Keyword(s):  
Power Output ◽  
Muscle Power ◽  
Flight Muscle ◽  
Bombus Terrestris ◽  
Muscle Length ◽  
Muscle Temperature ◽  
Free Flight ◽  
Mechanical Power Output ◽  
Ventral Muscle ◽  
Work Loop

1. The work loop approach was used to measure mechanical power output from an asynchronous flight muscle, the dorso-ventral muscle of the bumblebee Bombus terrestris. Measurements were made at the optimum muscle length for work output at 30 °C and at a muscle temperature (40 °C) and oscillatory frequency (141­173 Hz, depending on the size of the animal) characteristic of free flight. Oscillatory strain amplitude was adjusted to maximize power output. 2. There was much preparation-to-preparation variability in power output. Power output in the muscles with the highest values was slightly greater than 100 W kg-1. It is argued that there are many experimental factors which might reduce measured power output below that in the living bumblebee, and no obvious factors which might lead to overestimates of muscle power. The conclusion is that flight muscle in the intact bumblebee can produce at least 100 W kg-1.

scholarly journals Efficiency of fast- and slow-twitch muscles of the mouse performing cyclic contractions.

1994 ◽  
Vol 193 (1) ◽  
pp. 65-78 ◽  
Author(s):  
C J Barclay
Keyword(s):  
Heat Production ◽  
Power Output ◽  
Muscle Length ◽  
Mechanical Efficiency ◽  
Maximum Efficiency ◽  
Work Output ◽  
Cycle Frequency ◽  
Length Changes ◽  
Slow Twitch ◽  
Initial Heat

The mechanical efficiency of mouse fast- and slow-twitch muscle was determined during contractions involving sinusoidal length changes. Measurements were made of muscle length, force production and initial heat output from bundles of muscle fibres in vitro at 31 degrees C. Power output was calculated as the product of the net work output per sinusoidal length cycle and the cycle frequency. The initial mechanical efficiency was defined as power output/(rate of initial heat production+power output). Both power output and rate of initial heat production were averaged over a full cycle of length change. The amplitude of length changes was +/- 5% of muscle length. Stimulus phase and duration were adjusted to maximise net work output at each cycle frequency used. The maximum initial mechanical efficiency of slow-twitch soleus muscle was 0.52 +/- 0.01 (mean +/- 1 S.E.M. N = 4) and occurred at a cycle frequency of 3 Hz. Efficiency was not significantly different from this at cycle frequencies of 1.5-4 Hz, but was significantly lower at cycle frequencies of 0.5 and 1 Hz. The maximum efficiency of fast-twitch extensor digitorum longus muscle was 0.34 +/- 0.03 (N = 4) and was relatively constant (0.32-0.34) over a broad range of frequencies (4-12 Hz). A comparison of these results with those from previous studies of the mechanical efficiency of mammalian muscles indicates that efficiency depends markedly on contraction protocol.

POWER OUTPUT OF FISH MUSCLE FIBRES PERFORMING OSCILLATORY WORK: EFFECTS OF ACUTE AND SEASONAL TEMPERATURE CHANGE

1991 ◽  
Vol 157 (1) ◽  
pp. 409-423 ◽  
Author(s):  
TIMOTHY P. JOHNSON ◽  
IAN A. JOHNSTON
Keyword(s):  
Strain Amplitude ◽  
Power Output ◽  
Muscle Length ◽  
Fast Muscle ◽  
Muscle Fibres ◽  
Work Output ◽  
Cycle Frequency ◽  
Maximum Power Output ◽  
Resting Muscle ◽  
Positive Work

Fast muscle fibres were isolated from the abdominal myotomes of the shorthorned sculpin Myoxocephalus scorpius L. Sinusoidal length changes were imposed about resting muscle length and fibres were stimulated at a selected phase during the strain cycle. The work output per cycle was calculated from the area of the resulting force-position loops. The strain amplitude required for maximum work per cycle had a distinct optimum at ±5 % of resting length, which was independent of temperature. Maximum positive work loops were obtained by retarding the stimulus relative to the start of the length-change cycle by 30° (full cycle=360°). The maximum negative work output was obtained with a 210° stimulus phase shift. At intermediate stimulus phase shifts, work loops became complex with both positive (anticlockwise) and negative (clockwise) components. The number and timing of stimuli were adjusted, at constant strain amplitude (±5% of resting muscle length), to optimize net positive work output over a range of cycle frequencies. The cycle frequency required for maximum power output (work per cycle times cycle frequency) increased from around 5–7 Hz at 4°C to 9–13 Hz at 15°C. The maximum tension generated per cycle at 15°C was around two times higher at all cycle frequencies in summer-relative to winter-acclimatized fish. Fast muscle fibres from summer fish produced consistently higher tensions at 4°C, but the differences were only significant at 15 Hz. Acclimatization also modified the relationship between peak length and peak force at 4°C and 15°C. The maximum power output of muscle fibres showed little seasonal variation at 4°C and was in the range 20–25 W kg−1. In contrast, at 15°C, maximum muscle power output increased from 9 W kg−1 in the winter- to 30 W kg−1 in the summeracclimatized fish

scholarly journals The Mechanical Power Output of a Crab Respiratory Muscle

1988 ◽  
Vol 140 (1) ◽  
pp. 287-299 ◽  
Author(s):  
DARRELL R. STOKES ◽  
ROBERT K. JOSEPHSON
Keyword(s):  
Power Output ◽  
Muscle Power ◽  
Beat Frequency ◽  
Muscle Length ◽  
Metabolic Energy ◽  
Mechanical Power ◽  
Work Output ◽  
Hydraulic Power ◽  
Cycle Frequency ◽  
Mechanical Power Output

The mechanical power output was measured from scaphognathite (SG = gill bailer) muscle L2B of the crab Carcinus maenas (L.). The work was determined from the area of the loop formed by plotting muscle length against force when the muscle was subjected to sinusoidal length change (strain) and phasic stimulation in the length cycle. The stimulation pattern (10 stimuli per burst, burst length = 20% of cycle length) mimicked that which has been recorded from muscle L2B in intact animals. Work output was measured at cycle frequencies ranging from 0.5 to 5 Hz. The work output at optimum strain and stimulus phase increased with increasing cycle frequency to a maximum at 2–3 Hz and declined thereafter. The maximum work per cycle was 2.7 J kg−1 (15 °C). The power output reached a maximum (8.8 W kg−1) at 4 Hz. Both optimum strain and optimum stimulus phase were relatively constant over the range of burst frequencies examined. Based on the fraction of the total SG musculature represented by muscle L2B (18%) and literature values for the oxygen consumption associated with ventilation in C. maenas and for the hydraulic power output from an SG, we estimate that at a beat frequency of 2 Hz the SG muscle is about 10% efficient in converting metabolic energy to muscle power, and about 19% efficient in converting muscle power to hydraulic power.

scholarly journals EFFECTS OF OPERATING FREQUENCY AND TEMPERATURE ON MECHANICAL POWER OUTPUT FROM MOTH FLIGHT MUSCLE

1990 ◽  
Vol 149 (1) ◽  
pp. 61-78 ◽  
Author(s):  
R. D. STEVENSON ◽  
ROBERT K. JOSEPHSON
Keyword(s):  
Power Output ◽  
Flight Muscle ◽  
Beat Frequency ◽  
Length Change ◽  
Mechanical Power ◽  
Work Output ◽  
Optimal Frequency ◽  
Hovering Flight ◽  
Cycle Frequency ◽  
Mechanical Power Output

1. Mechanical work output was determined for an indirect flight muscle, the first dorsoventral, of the tobacco hawkmoth Manduca sexta. Work output per cycle was calculated from the area of force-position loops obtained during phasic electrical stimulation (1 stimulus cycle−1) and imposed sinusoidal length change. There was an optimal stimulus phase and an optimal length change (strain) that maximized work output (loop area) at constant cycle frequency and temperature. 2. When cycle frequency was increased at constant temperature, work output first increased and then decreased. It was always possible to find a frequency that maximized work output. There also always existed a higher frequency (termed the ‘optimal’ frequency in this paper) that maximized the mechanical power output, which is the product of the cycle frequency (s−1) and the work per cycle (J). 3. As temperature increased from 20 to 40°C, the mean maximum power output increased from about 20 to about 90 W kg−1 of muscle (Q10=2.09). There was a corresponding increase in optimal frequency from 12.7 to 28.3 Hz, in the work per cycle at optimal frequency from 1.6 to 3.2Jkg−1 muscle and in mean optimal strain from 5.9 to 7.9%. 4. Two electrical stimuli per cycle cannot increase power output at flight frequencies, but if frequency is reduced then power output can be increased with multiple stimulation. 5. Comparison of mechanical power output from muscle and published values of energy expenditure during free hovering flight of Manduca suggests that mechanical efficiency is about 10%. 6. In the tobacco hawkmoth there is a good correspondence between, on the one hand, the conditions of temperature (35–40°C) and cycle frequency (28–32 Hz) that produce maximal mechanical power output in the muscle preparation and, on the other hand, the thoracic temperature (35–42°C) and wing beat frequency (24–32 Hz) observed during hovering flight.

scholarly journals Alterations in flight muscle ultrastructure and function in Drosophila tropomyosin mutants.

1996 ◽  
Vol 135 (3) ◽  
pp. 673-687 ◽  
Author(s):  
A J Kreuz ◽  
A Simcox ◽  
D Maughan
Keyword(s):  
Amino Acid ◽  
Power Output ◽  
Flight Muscle ◽  
Structure And Function ◽  
Wild Type ◽  
Muscle Tropomyosin ◽  
Ultrastructure And Function ◽  
And Function ◽  
Net Power Output ◽  
Current Report

Drosophila indirect flight muscle (IFM) contains two different types of tropomyosin: a standard 284-amino acid muscle tropomyosin, Ifm-TmI, encoded by the TmI gene, and two > 400 amino acid tropomyosins, TnH-33 and TnH-34, encoded by TmII. The two IFM-specific TnH isoforms are unique tropomyosins with a COOH-terminal extension of approximately 200 residues which is hydrophobic and rich in prolines. Previous analysis of a hypomorphic TmI mutant, Ifm(3)3, demonstrated that Ifm-TmI is necessary for proper myofibrillar assembly, but no null TmI mutant or TmII mutant which affects the TnH isoforms have been reported. In the current report, we show that four flightless mutants (Warmke et al., 1989) are alleles of TmI, and characterize a deficiency which deletes both TmI and TmII. We find that haploidy of TmI causes myofibrillar disruptions and flightless behavior, but that haploidy of TmII causes neither. Single fiber mechanics demonstrates that power output is much lower in the TmI haploid line (32% of wild-type) than in the TmII haploid line (73% of wild-type). In myofibers nearly depleted of Ifm-TmI, net power output is virtually abolished (< 1% of wild-type) despite the presence of an organized fibrillar core (approximately 20% of wild-type). The results suggest Ifm-TmI (the standard tropomyosin) plays a key role in fiber structure, power production, and flight, with reduced Ifm-TmI expression producing corresponding changes of IFM structure and function. In contrast, reduced expression of the TnH isoforms has an unexpectedly mild effect on IFM structure and function.

EFFICIENCY OF ENERGY CONVERSION DURING SINUSOIDAL MOVEMENT OF WHITE MUSCLE FIBRES FROM THE DOGFISH SCYLIORHINUS CANICULA

1993 ◽  
Vol 183 (1) ◽  
pp. 137-147 ◽  
Author(s):  
N. A. Curtin ◽  
R. C. Woledge
Keyword(s):  
Power Output ◽  
High Efficiency ◽  
Fibre Length ◽  
Dry Mass ◽  
Energy Output ◽  
Maximum Efficiency ◽  
Muscle Fibres ◽  
Work Output ◽  
Sinusoidal Movement ◽  
Myotomal Muscle

Net work output and heat production of white myotomal muscle fibres from the dogfish were measured during complete cycles of sinusoidal movement at 12°C. The peak-to-peak movement was about 9 % of the muscle fibre length; three stimuli at 32 ms intervals were given in each mechanical cycle. The frequency of movement and the timing of the stimulation were varied for each preparation to find the optimal conditions for power output and those optimal for efficiency (the ratio of net work output to total energy output as heat+work). To achieve either maximum power or maximum efficiency, the tetanus must start while the muscle fibres are being stretched, before the beginning of the shortening part of the mechanical cycle. The highest power output, averaged over one cycle, was 0.23+/−0.014 W g-1 dry mass (+/−s.e.m., N=9, 46.9+/−2.8 mW g-1 wet mass) and was produced during movement at 3.5 Hz. The highest efficiency, 0.41+/−0.02 (+/−s.e.m., N=13), occurred during movements at 2.0-2.5 Hz. This value is higher than the efficiency previously measured during isovelocity shortening of these fibres. The implications of the high efficiency for crossbridge models of muscle contraction are discussed.

Dynamics of in vivo power output and efficiency of Nasonia asynchronous flight muscle

2006 ◽  
Vol 124 (1) ◽  
pp. 93-107 ◽  
Author(s):  
Fritz-Olaf Lehmann ◽  
Nicole Heymann
Keyword(s):  
Power Output ◽  
Flight Muscle

scholarly journals Influence of muscle length on work from trabecular muscle of frog atrium and ventricle.

1995 ◽  
Vol 198 (10) ◽  
pp. 2221-2227 ◽  
Author(s):  
D A Syme ◽  
R K Josephson
Keyword(s):  
Isometric Force ◽  
Muscle Length ◽  
Work Capacity ◽  
Linear Increase ◽  
Ventricular Muscle ◽  
Work Output ◽  
Loop Technique ◽  
Work Done ◽  
The Relationship ◽  
Work Loop

The work capacity of segments of atrial and ventricular muscle from the frog Rana pipiens was measured as a function of muscle length using the work loop technique. Both the work done during shortening and the work required to re-lengthen the muscle after shortening increased with muscle length. Net work increased with length up to a maximum, beyond which work declined. The optimum sarcomere length for work output was 2.5-2.6 microns for both atrial and ventricular muscle. Isometric force increased with muscle length to lengths well beyond the optimum for work output. Thus, the decline in work at long lengths is not simply a consequence of a reduction in the capacity of heart muscle to generate force. It is proposed that it is the non-linear increase in work required to re-lengthen muscle with increasing muscle length which limits net work output and leads to a maximum in the relationship between net work and muscle length. Extension of the results from muscle strips to intact hearts suggests that the work required to fill the ventricle exceeds that available from atrial muscle at all but rather short ventricular muscle lengths.

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