Name: improving strength, power, muscle aerobic capacity, and glucose tolerance through short-term progressive strength training among elderly people
Uploaded: 2017-07-05T12:30:00
Description: Watch this Scientific Journal Video about Improving Strength, Power, Muscle Aerobic Capacity, and Glucose Tolerance through Short-term Progressive Strength Training Among Elderly People at JoVE.com

The authors are grateful to Andr&#233;e Nienkerk, Dennis Peyron, and Sebastian Skj&#246;ld for supervising the training sessions and several tests; to the subjects participating; to Tim Crosfield for language revision; and to the economic support from The Swedish School of Sport and Health Sciences.

The Regional Ethics Committee of Stockholm, Sweden, approved the design of the investigation.
1. Material
<ol>
	<li>Recruit relatively healthy women and men 65-80 years old who have BMI values between 20 and 30 kg&#183;m-2. Randomize them into two groups. Ensure that the individuals in both groups have relatively low physical activity levels (i.e., moderate daily physical activity and no regular exercise training).</li>
	<li>Exclude beta-blocker users and those with coron...

<ol>
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In this study, a number of techniques have been used to investigate the effects of short-term progressive resistance training on elderly subjects&#39; muscle function/morphology, aerobic capacity, and glucose tolerance. The main finding was that, compared to a control group, many improvements occurred in muscle aerobic capacity, glucose tolerance, strength, power, and muscle quality (i.e., protein involved in cell signaling and muscle fiber composition). An increase was, for example, seen for: static, eccentric,...

Aging is associated with a loss of muscle mass (sarcopenia), strength, and power. Reduced strength, and probably even more importantly, power, results in immobility, an increased risk of injury, and a reduced quality of life. Resistance training is a well-known strategy to counteract sarcopenia and deteriorating muscle function. A rough estimate of muscle strength can be obtained from the load or number of achieved repetitions. However, this study obtained more detailed and accurate information on muscle function using an isokinetic dynamometer to gather information on the torque during isometric, concentric and eccentric contraction, as well as on the kinetics of force development.
Aerobic capacity, both at the whole-body level (VO2max) and in skeletal muscle, is reduced in elderly people. The decline in heart rate with age explains a large part of the decrease in VO2max1, but reduced muscle oxidative capacity, largely related to reduced physical activity2, does contribute. Impaired mitochondrial function may also be involved in the development of sarcopenia and insulin resistance3. The muscle aerobic capacity was assessed in muscle biopsies through biochemical analyses of the contents of mitochondrial enzymes and protein complexes located both in the matrix (i.e., citrate synthase) and the inner mitochondrial membrane. In addition, histochemical techniques were used to measure the effect of resistance training on muscle morphology (i.e., fiber type composition, fiber cross-sectional area, and capillary density). An alternative method to assess muscle aerobic capacity would be to use magnetic resonance spectroscopy to measure the rate of creatine phosphate resynthesis after exercise-induced depletion4. This method provides an estimate of the in vivo muscle aerobic capacity but cannot discriminate between mitochondrial dysfunction and circulatory disorders. Furthermore, the high costs of equipment limit the use of this technique in most laboratories. Aerobic capacity (VO2max and mitochondrial density) can be improved by endurance exercise in both young and old people5,6. However, the effect of resistance training on these parameters has been less investigated, especially in elderly subjects, and the results are conflicting7,8,9,10.
Type 2 diabetes is a widespread disease in the elderly population. Physical inactivity and obesity are major lifestyle-related factors explaining the increased incidence of type 2 diabetes. Low-intensity aerobic exercise is often recommended to subjects with reduced glucose tolerance. However, it is unclear how strength training in the elderly affects glucose tolerance/insulin sensitivity11,12. The most accurate way to measure insulin sensitivity is to use the glucose clamp technique, where the blood glucose is maintained constant by glucose infusion during conditions of elevated insulin13. The disadvantages with this technique are that it is time consuming and invasive (arterial catheterization) and requires special laboratory facilities. In this study, the oral glucose tolerance test, which is common in healthcare units, was used. This method is suitable when several subjects are to be investigated for a limited period of time.
The testing and timeline of the experimental procedure can be summarized as follows. Use three separate days for testing before and after an eight-week period, with the same arrangement and approximate time schedules (&#8805;24 h between each day, Figure 1). On the first test day, measure: anthropometric data, such as height, body mass, fat-free mass (FFM), and upper leg circumference (i.e., 15 cm above the apex patellae in a relaxed supine position); submaximal cycling ability; and knee muscle strength, as described in steps 4 and 5. Take a muscle biopsy from the thigh on the second test day. For further descriptions, see step 6.1. Test oral glucose tolerance (OGTT) on the last testing day. For further descriptions, see step 7.1. Ask all participants to avoid vigorous physical activity for 24 h and to fast overnight prior to each test day. However, ask them to avoid strenuous physical activity for 48 h before the OGTT test day. Ask them to follow their normal everyday physical activity and diet habits. Note that pre- and post-intervention, both groups&#39; self-reported food intake and type of foods were unchanged.
<img alt="Figure 1" class="xfigimg" src="/files/ftp_upload/55518/55518fig1.jpg" /> 
Figure 1: Experimental protocol. Schematic diagram. The timing between the three pre- and post-tests was similar for each subject and was at least 24 h. Further details are given in the text. This figure has been modified from Frank et al. Scand. J. Med. Sci. Sports. 2016: 26, 764-73.28 <a href="http://cloudflare.jove.com/files/ftp_upload/55518/55518fig1large.jpg" target="_blank">Please click here to view a larger version of this figure.</a>
This study sought to investigate the effect of short-term resistance training in elderly people on muscle oxidative capacity and glucose tolerance. The second aim was to examine the effect on strength, power, and muscle qualitative improvements (i.e., proteins involved in cell signaling and muscle fiber type composition).

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			<td height="21" style="height:21px;width:471px;">Western blot</td>
			<td style="width:479px;"></td>
			<td style="width:148px;"></td>
			<td style="width:167px;"></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Pierce 660nm Protein Assay Kit</td>
			<td>Thermo Scientific, Rockford, IL, USA</td>
			<td align="right">22662</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">SuperSignal West Femto Maximum Sensitivity Substrate&#160;</td>
			<td>Thermo Scientific</td>
			<td align="right">34096</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Halt Protease Inhibitor Cocktail (100X)&#160;</td>
			<td>Thermo Scientific</td>
			<td align="right">78429</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Restore PLUS Western Blot Stripping Buffer</td>
			<td>Thermo Scientific</td>
			<td align="right">46430</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Pierce Reversible Protein Stain Kit for PVDF Membranes</td>
			<td>Thermo Scientific</td>
			<td align="right">24585</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">10 st - 4&#8211;20% Criterion TGX Gel, 18 well, 30 &#181;l&#160;</td>
			<td>Bio-Rad Laboratories, Richmond, CA, USA</td>
			<td style="width:148px;">567-1094</td>
			<td></td>
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		<tr height="21">
			<td height="21" style="height:21px;">Immun-Blot PVDF Membrane&#160;</td>
			<td>Bio-Rad</td>
			<td>162-0177</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Precision Plus Protein Dual Color Standards&#160;</td>
			<td>Bio-Rad</td>
			<td>161-0374</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">2x Laemmli Sample Buffer</td>
			<td>Bio-Rad</td>
			<td>161-0737</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">10x Tris/Glycine</td>
			<td>Bio-Rad</td>
			<td>161-0771</td>
			<td></td>
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		<tr height="21">
			<td height="21" style="height:21px;">2-Mercaptoethanol</td>
			<td>Bio-Rad</td>
			<td>161-0710</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Tween 20</td>
			<td>Bio-Rad</td>
			<td>P1379-250ML</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Band analysis with Quantity One version 4.6.3.software</td>
			<td>Bio-Rad</td>
			<td></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">1% phosphatase inhibitor coctail</td>
			<td style="width:479px;">Sigma-Aldrich, Saint Louis, Missouri, USA</td>
			<td></td>
			<td></td>
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		<tr height="25">
			<td height="25" style="height:25px;width:471px;">Name</td>
			<td style="width:479px;">Company</td>
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			<td>Comments</td>
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			<td height="21" style="height:21px;width:471px;">Antibodies</td>
			<td style="width:479px;"></td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">mTOR (1:1000)</td>
			<td>Cell Signaling, Danvers, Massachusetts, USA</td>
			<td align="right" style="width:148px;">2983</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Akt (1:1000)</td>
			<td>Cell Signaling, Danvers</td>
			<td align="right" style="width:148px;">9272</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Secondary anti-rabbit and anti-mouse HRP-linked (1:10000)</td>
			<td>Cell Signaling, Danvers</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Citrate synthase (CS) (1:1000)</td>
			<td>Gene tex, San Antonio, California, USA</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">OXPHOS (1:1000)</td>
			<td>Abcam, Cambridge, UK</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="26">
			<td height="26" style="height:26px;width:471px;">Name</td>
			<td style="width:479px;">Company</td>
			<td style="width:148px;">Catalog Number</td>
			<td>Comments</td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Equipment - Analysis of muscle samples</td>
			<td></td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Bullet Blender 1.5 for homogenizing</td>
			<td>Next Advance, New York, USA</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Plate reader</td>
			<td>Tecan infinite F200 pro, M&#228;nnedorf, Switzerland</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="38">
			<td height="38" style="height:38px;width:471px;">Name</td>
			<td style="width:479px;">Company</td>
			<td style="width:148px;">Catalog Number</td>
			<td>Comments</td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Histochemistry</td>
			<td style="width:479px;"></td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Mayer hematoxylin</td>
			<td style="width:479px;">HistoLab, V&#228;stra Fr&#246;lunda, Sweden&#160;</td>
			<td align="right" style="width:148px;">1820</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Oil Red o</td>
			<td style="width:479px;">Sigma-Aldrich, Saint Louis, Missouri, USA</td>
			<td style="width:148px;">00625-25y</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">NaCl</td>
			<td style="width:479px;">Sigma-Aldrich</td>
			<td style="width:148px;">793566-2.5 kg</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Cobalt Chloride</td>
			<td style="width:479px;">Sigma-Aldrich</td>
			<td style="width:148px;">60818-50G</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Amylase</td>
			<td style="width:479px;">Sigma-Aldrich</td>
			<td style="width:148px;">A6255-25MG</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">ATP</td>
			<td style="width:479px;">Sigma-Aldrich</td>
			<td style="width:148px;">A2383-5G</td>
			<td></td>
		</tr>
		<tr height="22">
			<td height="22" style="height:22px;width:471px;">Glycine</td>
			<td style="width:479px;">VWR-chemicals / VWR-international, Sp&#229;nga, Sweden</td>
			<td style="width:148px;">101196X</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Calcium Chloride</td>
			<td style="width:479px;">VWR-chemicals / VWR-international</td>
			<td style="width:148px;">22328.262</td>
			<td></td>
		</tr>
		<tr height="22">
			<td height="22" style="height:22px;width:471px;">Iso-pentane</td>
			<td style="width:479px;">VWR-chemicals / VWR-international</td>
			<td style="width:148px;">24872.298</td>
			<td></td>
		</tr>
		<tr height="22">
			<td height="22" style="height:22px;width:471px;">Etanol 96%</td>
			<td style="width:479px;">VWR-chemicals / VWR-international</td>
			<td style="width:148px;">20905.296</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">NaOH</td>
			<td style="width:479px;">MERCK, Stockholm, Sweden</td>
			<td style="width:148px;">1.06498.1000</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Na acetate</td>
			<td style="width:479px;">MERCK</td>
			<td style="width:148px;">1.06268.1000</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">KCl</td>
			<td style="width:479px;">MERCK</td>
			<td style="width:148px;">1.04936.1000</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Ammonium Sulphide</td>
			<td style="width:479px;">MERCK</td>
			<td style="width:148px;">U1507042828</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Acetic acid 100%</td>
			<td style="width:479px;">MERCK</td>
			<td style="width:148px;">1.00063.2511</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Schiffs&#180; Reagent</td>
			<td style="width:479px;">MERCK</td>
			<td style="width:148px;">1.09033.0500</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Periodic acid</td>
			<td style="width:479px;">MERCK</td>
			<td style="width:148px;">1.00524.0025</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Chloroform</td>
			<td style="width:479px;">MERCK</td>
			<td style="width:148px;">1.02445.1000</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">pH-meter LANGE</td>
			<td style="width:479px;">HACH LANGE GMBH, Dusseldorf, Germany</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Light microscope</td>
			<td>Olympus BH-2, Olympus, Tokyo, Japan</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Cryostat&#160; Leica CM1950</td>
			<td>Leica Microsystems, Wetzlar, Germany</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Leica software Leica Qwin V3</td>
			<td>Leica Microsystems</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Gel Doc 2000 - Bio-Rad, camera setup</td>
			<td>Bio-Rad Laboratories AB, Solna, Sweden&#160;</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Software program Quantift One - 4.6 (version 4.6.3; Bio Rad)</td>
			<td>Bio-Rad Laboratories AB, Solna, Sweden&#160;</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="23">
			<td height="23" style="height:23px;width:471px;">Name</td>
			<td style="width:479px;">Company</td>
			<td style="width:148px;">Catalog Number</td>
			<td>Comments</td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Oral glucos tolerance test, OGTT</td>
			<td style="width:479px;"></td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Glukos APL 75 g</td>
			<td>APL, Stockholm, Sweden</td>
			<td align="right">323,188</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Automated analyser Biosen 5140</td>
			<td>EKF Diagnostics, Barleben, Germany</td>
			<td></td>
			<td></td>
		</tr>
		<tr height="42">
			<td height="42" style="height:42px;width:471px;">Insulin and C-peptide in plasma kit ELISA</td>
			<td style="width:479px;">Mercodia AB, Uppsala Sweden</td>
			<td style="width:148px;">10-1132-01, 10-1134-01</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Plate reader</td>
			<td>Tecan infinite F200 pro, M&#228;nnedorf, Switzerland</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="23">
			<td height="23" style="height:23px;width:471px;">Name</td>
			<td style="width:479px;">Company</td>
			<td style="width:148px;">Catalog Number</td>
			<td>Comments</td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Further equipment</td>
			<td style="width:479px;"></td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Measures of fat-free-mass</td>
			<td>FFM-Tanita T5896, Tanita, Tokyo, Japan</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Strength training equipment for all training exercises</td>
			<td style="width:479px;">Cybex International Inc., Medway, Massachusetts, USA&#160;</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Cycle ergometer&#160;</td>
			<td>Monark Ergometer 893E, Monark Exercises, Varberg, Sweden&#160;</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Heart rate monitor RS800, Polar</td>
			<td style="width:479px;">Polar Electro OY, Kampele, Finland</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Oxycin-Pro - automatic ergo-spirometric device</td>
			<td>Erich Jaeger GmbH, Hoechberg, Germany</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Isokinetic dynamometer, Isomed 2000, knee muscle strength</td>
			<td style="width:479px;">D&#38;R Ferstl GmbH, Henau, Germany</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">CED 1401 data acquisition system and Signal software</td>
			<td>Cambridge Electronic Design, Cambridge, UK</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Software for muscle strength analysis, Spike 2, version 7</td>
			<td style="width:479px;">Signal Hound, LA Center, WA, USA</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Statistica software for statistical analyses</td>
			<td>Statistica, Stat soft. inc, Tulsa, Oklahoma, USA</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="23">
			<td height="23" style="height:23px;width:471px;">Name</td>
			<td style="width:479px;">Company</td>
			<td style="width:148px;">Catalog Number</td>
			<td>Comments</td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Muscle biopsy equipment</td>
			<td style="width:479px;"></td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Weil Blakesley conchotome</td>
			<td>Wisex, M&#246;lndal, Sweden</td>
			<td style="width:148px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;">Local anesthesia&#160;</td>
			<td>Carbocain, 20 mL, 20 mg/mL; Astra Zeneca, S&#246;dert&#228;lje, Sweden</td>
			<td align="right" style="width:148px;">169,367</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:471px;">Surgical Blade</td>
			<td style="width:479px;">Feather Safety Razor CO, LTD, Osaka, Japan&#160;</td>
			<td align="right" style="width:148px;">11048030</td>
			<td></td>
		</tr>
	</tbody>
</table>

improving strength, power, muscle aerobic capacity, and glucose tolerance through short-term progressive strength training among elderly people

This protocol describes the simultaneous use of a broad span of methods to examine muscle aerobic capacity, glucose tolerance, strength, and power in elderly people performing short-term resistance training (RET). Supervised progressive resistance training for 1 h three times a week over 8 weeks was performed by RET participants (71&#177;1 years, range 65-80). Compared to a control group without training, the RET showed improvements on the measures used to indicate strength, power, glucose tolerance, and several parameters of muscle aerobic capacity. Strength training was performed in a gym with only robust fitness equipment. An isokinetic dynamometer for knee extensor strength permitted the measurement of concentric, eccentric, and static strength, which increased for the RET group (8-12% post- versus pre-test). The power (rate of force development, RFD) at the initial 0-30 ms also showed an increase for the RET group (52%). A glucose tolerance test with frequent blood glucose measurements showed improvements only for the RET group in terms of blood glucose values after 2 h (14%) and the area under the curve (21%). The blood lipid profile also improved (8%). From muscle biopsy samples prepared using histochemistry, the amount of fiber type IIa increased, and a trend towards a decrease in IIx in the RET group reflected a change to a more oxidative profile in terms of fiber composition. Western blot (to determine the protein content related to the signaling for muscle protein synthesis) showed a rise of 69% in both Akt and mTOR in the RET group; this also showed an increase in mitochondrial proteins for OXPHOS complex II and citrate synthase (both ~30%) and for complex IV (90%), in only the RET group. We demonstrate that this type of progressive resistance training offers various improvements (e.g., strength, power, aerobic capacity, glucose tolerance, and plasma lipid profile).

Material
In the study, 21 relatively healthy women and men, 65-80 years old and with BMI values between 20 and 30 kg&#183;m-2 participated and were randomized into two groups. Individuals in both groups had relatively low physical activity levels (i.e., a moderate everyday physical activity level and no regular exercise training). One group (n=12, 6 women and 6 men) performed RET under a trainer for 1 h three ti...

Watch this Scientific Journal Video about Improving Strength, Power, Muscle Aerobic Capacity, and Glucose Tolerance through Short-term Progressive Strength Training Among Elderly People at JoVE.com

Improving Strength, Power, Muscle Aerobic Capacity, and Glucose Tolerance through Short-term Progressive Strength Training Among Elderly People

The effect of short-term resistance training on elderly people was investigated through the simultaneous use of several methods. Compared to a control group, many improvements were seen, including on muscle aerobic capacity, glucose tolerance, strength, power, and muscle quality (i.e., protein involved in cell signaling and muscle fiber type composition).

This protocol describes the simultaneous use of a broad span of methods to examine muscle aerobic capacity, glucose tolerance, strength, and power in ...

The overall goal of this experiment with a broad span of methods is to examine muscle aerobic capacity, muscle protein expression, glucose tolerance, strength, and power in elderly people performing short term progressive resistance training. The methods used in this study, they are not unique but by combining them with a broad spectrum of methods and applying them to studies of strength training in elderly, we were able to show very important reasons not only concerning improvements in strength, which was expected, but also in muscle aerobic capacity, in insulin sensitivity as well as other aspects in muscle tissue. The value of the methods we have used is great regarding therapy with strength training for elderly people who are physically inactive.And the methods we use are, for example, following strength, power, glucose tolerance, and protein expressions. These methods, you can use them into rehabilitation, strength training for old people, among the elderlies. But in addition to this, you can even use them in other kind of context such as sports physiology.Generally individuals new to these methods will struggle from the multi-step process in examining protein expression in the muscle. Also for the elderly subjects will struggle from obtaining real maximum value for strength and power. These methods are developed in order to follow the changes in the muscle with power, energy, concentration, in science concentration, and other factors of importance with strength training.Begin by asking the subject to sit on the bench of an isokinetic dynamometer. Fix the subject's trunk with straps over the shoulders and hips. And securely strap the subject's shank to the dynamometer shaft with two straps, one below the knee and one above the ankle.Then align the knee joint axis with the rotational center of the dynamometer shaft. Set the range of motion between 90 degrees and 15 degrees. After the subject is secured, ask them to perform a total of eight trials alternating between four maximal voluntary eccentric extensions and four maximal voluntary concentric knee extensions with the right leg at a constant angular velocity of 30 degrees per second.Specifically for the eccentric task, ask the subject to resist the dynamometer shaft with maximal effort through the whole movement from the 15 degree to 90 degree knee angle. While in the concentric task, ask the subject to press the lower leg in the dynamometer shaft in a knee extension as hard as possible throughout the whole motion range. Next, allow a four-minute rest after the dynamic recordings.Then, assess the static maximal voluntary contraction torque four times at a 65-degree knee angle by asking the subject to kick as fast and as hard as they can against the dynamometer shaft which should be fixed at 65 degrees. In the software program for muscle strength analysis register the highest peak torque in the eccentric and concentric recordings respectively for each subject and the highest strength value among the four static trials. Following the eccentric, concentric, and static measurements, record the highest value obtained from each task for each subject.Then, move the cursor to the 7.5 newton metre value on the Y scale to obtain the position for zero milliseconds in order to measure the rate of forced development during zero to 30 milliseconds and zero to 200 milliseconds in the highest value found among the static trials. Finally, for the pretest assessment, set the cursor on the 30 millisecond value and write down the value showing the raise in newton meters at 30 milliseconds. To take a muscle biopsy on test day two, start by injecting one to two milliliters of local anesthesia subcutaneously and into the fascia.After a couple of minutes, make and incision with a small scalpel through the skin and fascia approximately one third of the distance from the patella to the anterior superior iliac spine. Then, take a muscle biopsy from the middle portion of the thigh muscle, vastus lateralis, using a conchotome. On test day three perform the oral glucose tolerance test or OGTT, on subjects who have fasted overnight and have not done any strenuous exercise on the testing day or the day before.For the test, have the subject lie in a supine position. Take four milliliter blood samples via venous cannula in the antecubital vein 15 minutes before and just prior to the intake of glucose followed by 15, 30, 60, 90, and 120 minutes after the ingestion of the glucose. Use the samples to perform standard glucose level tests.Begin by freeze drying the muscle sample in a Lifelizer at a pressure below 10 to the minus one millibar for 12 hours. Dissect it using a needle and forceps under a light microscope so that it is free of blood and connective tissue. And store it at minus 80 degrees celsius.Then, place a scoop of 0.5 millimeter zirconium oxide beads in each tube with the muscle. Add 80 microliters per milligram of ice cold buffer. And homogenize for two times one minute at speed step seven to eight and four degrees celsius.Centrifuge the homogenate for 10 minutes at 10, 000 times G.Transfer the remaining supernatant to new tubes and discard the pellet containing the structural proteins. Spectrophotometrically determine the protein concentration in the supernatant with a commercially available kit using a plate reader at 660 nanometers. Then dilute the samples with 2x Laemmli sample buffer and homogenizing buffer to a final protein concentration of 1.5 micrograms per microliter.Heat them to 95 degrees celsius for five minutes to denature the proteins. Store the diluted samples at minus 20 degrees celsius prior to analysis. For native polyacrylamide gel electrophoresis, of PAGE, load 30 micrograms of protein from each sample into 18 well precast gradient cells.And perform electrophoresis at 300 volts for 30 minutes on ice. Next, equilibrate the gel and transfer buffer for 30 minutes at four degrees celsius. Transfer proteins to polyvinylidene fluoride membranes with 0.2 micron pore sizes at a constant current of 300 milliamps for three hours at four degrees celsius.To confirm equal loading and transfer stain the membranes with a total protein stain. The membrane is separated for each target protein. Then, block the membrane for one hour at room temperature in Tris buffered saline containing 5%nonfat milk.Add primary antibodies diluted in TBS containing 2.5%nonfat milk and supplemented with 0.1%Tween-20. And then incubate the membranes overnight. Following primary antibody incubation wash the membranes with TBSTM, and incubate with secondary antibodies conjugated with horseradish peroxidase for one hour at room temperature.Wash again with TBSTM followed by four additional five-minute washes with TBS. Finally, apply six to 12 mL of chemiluminescent substrate to the membrane for five minutes. Place the membrane between two transparent plastic sheets.Then in front of a CCD camera blocking external light take serial exposures using a chemiluminescent camera filter. Compared to the CON group, the RET group showed improvement in static, eccentric, and concentric strength with eight to 12%Further, the dynamometer measurements also showed for the rate of forced development an increase of 52%at the initial zero to 30 milliseconds in the RET group. From the muscle biopsy samples, histochemistry indicated that the amount of fiber type IIa increased, and there was a trend to a decrease in 2x for the RET group from pre to post.More from the muscle biopsy samples, protein content related to the signaling of muscle protein synthesis showed a rise of 69%for both Akt and mTOR in the RET group. Analyses also proved among mitochondrial proteins an increase of about 30%for OXPHOS Complex II and citrate synthase and of 90%for complex IV only in the RET group. Further, the RET group's results from the glucose tolerance test showed improved blood glucose, both in blood values after two hours and for the area under the curve.The measurements done in this study were done on three different separate days for each subject and with one day's rest in between. That means the testing could be done in one week altogether for one subject. In addition to that, we have the muscle analysis, and Western Blot analysis, which takes another two days.While attempting, in these kind of procedures or projects, it's important to remember that you should have a day of rest between each test day. And for the subjects, of course, it's very important to know that they shouldn't do any kind of rigorous exercise the same day and the day before the test day, or each test day. By using the methods we have in the project one could also measure the maximum strength in the abdominal, in the back, in the arm muscles and see how maximal strength may improve, as well, in those groups of muscles by short term resistance training.Don't forget that when working with elderly people in strength training progress one should maintain or monitor their health status and also progressively increase the chain load in a manner that they will tolerate and also to keep their wellbeing.

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Anatomy and Physiology

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Unit 1

Muscle Tissue

SCIENTISTS IN ACTION

Measuring the Strength of Mice

Kondziela7 devised the inverted screen test and published it in 1964. It is a test of muscle strength using all four limbs. Most normal mice easily score maximum on this task; it is a quick but insensitive gross screen, and the weights test described in this article will provide a finer measure of muscular strength.
There are also several strain gauge-based pieces of apparatus available commercially that will provide more graded data than the inverted screen test, but their cost may put them beyond the reach of many laboratories which do not specialize in strength testing. Hence in 2000 a cheap and simple apparatus was devised by the author. It consists of a series of chain links of increasing length, attached to a "fur collector" a ball of fine wire mesh sold for preventing limescale build up in hard water areas. An accidental observation revealed that mice could grip these very tightly, so they proved ideal as a grip point for a weight-lifting apparatus. A common fault with commercial strength meters is that the bar or other grip feature is not thin enough for mice to exert a maximum grip. As a general rule, the thinner the wire or bar, the better a mouse can grip with its small claws.
This is a pure test of strength, although as for any test motivational factors could potentially play a role. The use of scale collectors, however, seems to minimize motivational problems as the motivation appears to be very high for most normal young adult mice.

Deficits in muscular strength occur in many clinical conditions such as motor neuron disease. The inverted screen and weight lifting tests described here measure strength in mice almost exclusively, with minimal influence of factors such as coordination.

Kondziela7 devised the inverted screen test and published it in 1964. It is a test of muscle strength using all four limbs. Most normal mice easily score ...

Manual Muscle Testing:  A Method of Measuring Extremity Muscle Strength Applied to Critically Ill Patients

Survivors of acute respiratory distress syndrome (ARDS) and other causes of critical illness often have generalized weakness, reduced exercise tolerance, and persistent nerve and muscle impairments after hospital discharge.1-6 Using an explicit protocol with a structured approach to training and quality assurance of research staff, manual muscle testing (MMT) is a highly reliable method for assessing strength, using a standardized clinical examination, for patients following ARDS, and can be completed with mechanically ventilated patients who can tolerate sitting upright in bed and are able to follow two-step commands. 7, 8 
This video demonstrates a protocol for MMT, which has been taught to &ge;43 research staff who have performed &gt;800 assessments on &gt;280 ARDS survivors. Modifications for the bedridden patient are included. Each muscle is tested with specific techniques for positioning, stabilization, resistance, and palpation for each score of the 6-point ordinal Medical Research Council scale.7,9-11 Three upper and three lower extremity muscles are graded in this protocol: shoulder abduction, elbow flexion, wrist extension, hip flexion, knee extension, and ankle dorsiflexion. These muscles were chosen based on the standard approach for evaluating patients for ICU-acquired weakness used in prior publications. 1,2.

Survivors of acute respiratory distress syndrome (ARDS) and critical illness frequently develop long-lasting muscle weakness. Manual muscle testing (MMT) is a standardized clinical examination commonly used to measure strength of peripheral skeletal muscle groups. This video demonstrates MMT using the 6-point Medical Research Council scale.

Survivors of acute respiratory distress syndrome (ARDS) and other causes of critical illness often have generalized weakness, reduced exercise tolerance, ...

A Murine Model of Muscle Training by Neuromuscular Electrical Stimulation

Neuromuscular electrical stimulation (NMES) is a common clinical modality that is widely used to restore1, maintain2 or enhance3-5 muscle functional capacity. Transcutaneous surface stimulation of skeletal muscle involves a current flow between a cathode and an anode, thereby inducing excitement of the motor unit and the surrounding muscle fibers. 
NMES is an attractive modality to evaluate skeletal muscle adaptive responses for several reasons. First, it provides a reproducible experimental model in which physiological adaptations, such as myofiber hypertophy and muscle strengthening6, angiogenesis7-9, growth factor secretion9-11, and muscle precursor cell activation12 are well documented. Such physiological responses may be carefully titrated using different parameters of stimulation (for Cochrane review, see 13). In addition, NMES recruits motor units non-selectively, and in a spatially fixed and temporally synchronous manner14, offering the advantage of exerting a treatment effect on all fibers, regardless of fiber type. Although there are specified contraindications to NMES in clinical populations, including peripheral venous disorders or malignancy, for example, NMES is safe and feasible, even for those who are ill and/or bedridden and for populations in which rigorous exercise may be challenging. 
Here, we demonstrate the protocol for adapting commercially available electrodes and performing a NMES protocol using a murine model. This animal model has the advantage of utilizing a clinically available device and providing instant feedback regarding positioning of the electrode to elicit the desired muscle contractile effect. For the purpose of this manuscript, we will describe the protocol for muscle stimulation of the anterior compartment muscles of a mouse hindlimb.

A murine model of neuromuscular electrical stimulation (NMES), a safe and inexpensive clinical modality, to the anterior compartment muscles is described. This model has the advantage of modifying a readily available clinical device for the purpose of eliciting targeted and specific muscle contractions in mice.

Neuromuscular electrical stimulation (NMES) is a common clinical modality that is widely used to restore1, maintain2 or enhance3-5 muscle functional ...

Characterization of Metabolic Status in Nonhuman Primates with the Intravenous Glucose Tolerance Test

The intravenous glucose tolerance test (IVGTT) plays a key role in the characterization of glucose homeostasis. When taken together with serum biochemical profiles, inclusive of blood glucose levels in both the fed and fasted state, HbA1c, insulin levels, clinical history of diet, body composition, and body weight status, an assessment of normal and abnormal glycemic control can be made. Interpretation of an IVGTT is done through measurement of changes in glucose and insulin levels over time in relation to the dextrose challenge. Critical components to be considered are: peak glucose and insulin levels reached in relation to T0 (end of glucose infusion), the glucose clearance rate K derived from the slope of rapid glucose clearance in the first 20 min (T1 to T20), the time to return to glucose baseline, and the area under the curve (AUC). These IVGTT measures will show characteristic changes as glucose homeostasis moves from a healthy to a diseased metabolic state5. Herein we will describe the characterization of nonhuman primates (Rhesus and Cynomolgus macaques), which are the most relevant animal model of Type II diabetes (T2D) in humans and the IVGTT and clinical profiles of these animals from a lean healthy, to obese dysmetabolic, and T2D state 8, 10, 11.

The goal of this protocol is to present a standard method to perform intravenous glucose tolerance tests (IVGTTs) to assess glycemic control in nonhuman primates and assess their metabolic status from healthy to dysmetabolic.

The intravenous glucose tolerance test (IVGTT) plays a key role in the characterization of glucose homeostasis. When taken together with serum biochemical ...

Adapted Resistance Training Improves Strength in Eight Weeks in Individuals with Multiple Sclerosis

Hip weakness is a common symptom affecting walking ability in people with multiple sclerosis (MS). It is known that resistance strength training (RST) can improve strength in individuals with MS, however; it remains unclear the duration of RST that is needed to make strength gains and how to adapt hip strengthening exercises for individuals of varying strength using only resistance bands. This paper describes the methodology to set up and implement an adapted resistance strength training program, using resistance bands, for individuals with MS. Directions for pre- and post-strength tests to evaluate efficacy of the strength-training program are included. Safety features and detailed instructions outline the weekly program content and progression. Current evidence is presented showing that significant strength gains can be made within 8 weeks of starting a RST program. Evidence is also presented showing that resistance strength training can be successfully adapted for individuals with MS of varying strength with little equipment.

Hip weakness is a common symptom affecting walking ability in people with multiple sclerosis. Isolated muscle strengthening is a useful method to target specific weaknesses. This protocol describes a progressive resistance-training program using exercise bands to increase hip muscle strength.

Hip weakness is a common symptom affecting walking ability in people with multiple sclerosis (MS). It is known that resistance strength training (RST) can ...

Proximal Cadaveric Femur Preparation for Fracture Strength Testing and Quantitative CT-based Finite Element Analysis

Cadaveric fracture testing is routinely used to understand factors that affect proximal femur strength. Because ex vivo biological tissues are prone to lose their mechanical properties over time, specimen preparation for experimental testing must be performed carefully to obtain reliable results that represent in vivo conditions. For that reason, we designed a protocol and a set of fixtures to prepare the femoral specimens such that their mechanical properties experienced minimal changes. The femora were kept in a frozen state except during preparation steps and mechanical testing. The relevant clinical measures of total hip and femoral neck bone mineral density (BMD) were obtained with a clinical dual X-ray absorptiometry (DXA) bone densitometer, and the 3D geometry and distribution of bone mineral were obtained using CT with a calibration phantom for quantitative estimations based on the greyscale values. Any possible bone disease, fracture, or the presence of implants or artifacts affecting the bone structure, was ruled out with X-ray scans. For preparation, all bones were carefully cleaned of excess soft tissue, and were cut and potted at the internal rotation angle of interest. A cutting fixture allowed the distal end of the bone to be cut off leaving the proximal femur at a desired length. To allow positioning of the femoral neck at prescribed angles during later CT scanning and mechanical testing, the proximal femoral shafts were potted in polymethylmethacrylate (PMMA) using a fixture designed specifically for desired orientations. The data collected from our experiments were then used for validation of quantitative computed tomography (QCT)-based finite element analysis (FEA), as described in a different protocol. In this manuscript, we present the protocol for the precise bone preparation for mechanical testing and subsequent QCT/FEA modeling. The current protocol was successfully applied to prepare about 200 cadaveric femora over a 6-year time period.

We present a robust protocol on how to carefully preserve and prepare cadaveric femora for fracture testing and quantitative computed tomography imaging. The method provides precise control over input conditions for the purpose of determining relationships between bone mineral density, fracture strength, and defining finite element model geometry and properties.

Cadaveric fracture testing is routinely used to understand factors that affect proximal femur strength.  Because ex vivo biological tissues are prone to ...

Phosphorus-31 Magnetic Resonance Spectroscopy: A Tool for Measuring In Vivo Mitochondrial Oxidative Phosphorylation Capacity in Human Skeletal Muscle

Skeletal muscle mitochondrial oxidative phosphorylation (OXPHOS) capacity, which is critically important in health and disease, can be measured in vivo and noninvasively in humans via phosphorus-31 magnetic resonance spectroscopy (31PMRS). However, the approach has not been widely adopted in translational and clinical research, with variations in methodology and limited guidance from the literature. Increased optimization, standardization, and dissemination of methods for in vivo 31PMRS would facilitate the development of targeted therapies to improve OXPHOS capacity and could ultimately favorably impact cardiovascular health. 31PMRS produces a noninvasive, in vivo measure of OXPHOS capacity in human skeletal muscle, as opposed to alternative measures obtained from explanted and potentially altered mitochondria via muscle biopsy. It relies upon only modest additional instrumentation beyond what is already in place on magnetic resonance scanners available for clinical and translational research at most institutions. In this work, we outline a method to measure in vivo skeletal muscle OXPHOS. The technique is demonstrated using a 1.5 Tesla whole-body MR scanner equipped with the suitable hardware and software for 31PMRS, and we explain a simple and robust protocol for in-magnet resistive exercise to rapidly fatigue the quadriceps muscle. Reproducibility and feasibility are demonstrated in volunteers as well as subjects over a wide range of functional capacities.

Phosphorus-31 Magnetic Resonance Spectroscopy: A Tool for Measuring In Vivo Mitochondrial Oxidative Phosphorylation Capacity in Human Skeletal Muscle

This work demonstrates the feasibility of an in vivo phosphorus-31 magnetic resonance spectroscopy (31PMRS) technique to quantify mitochondrial oxidative phosphorylation (OXPHOS) capacity in human skeletal muscle.

Skeletal muscle mitochondrial oxidative phosphorylation (OXPHOS) capacity, which is critically important in health and disease, can be measured in vivo ...

Study of In Vivo Glucose Metabolism in High-fat Diet-fed Mice Using Oral Glucose Tolerance Test (OGTT) and Insulin Tolerance Test (ITT)

Obesity represents the most important single risk factor in the pathogenesis of type 2 diabetes, a disease which is characterized by a resistance to insulin-stimulated glucose uptake and a gross decompensation of systemic glucose metabolism. Despite considerable progress in the understanding of glucose metabolism, the molecular mechanisms of its regulation in health and disease remain under-investigated, while novel approaches to prevent and treat diabetes are urgently needed. Diet derived glucose stimulates the pancreatic secretion of insulin, which serves as the principal regulator of cellular anabolic processes during the fed-state and thus balances blood glucose levels to maintain systemic energy status. Chronic overfeeding triggers meta-inflammation, which leads to alterations in peripheral insulin receptor-associated signaling and thus reduces the sensitivity to insulin-mediated glucose disposal. These events ultimately result in elevated fasting glucose and insulin levels as well as a reduction in glucose tolerance, which in turn serve as important indicators of insulin resistance. Here, we present a protocol for the generation and metabolic characterization of high-fat diet (HFD)-fed mice as a frequently used model of diet-induced insulin resistance. We illustrate in detail the oral glucose tolerance test (OGTT), which monitors the peripheral disposal of an orally administered glucose load and insulin secretion over time. Additionally, we present a protocol for the insulin tolerance test (ITT) to monitor whole-body insulin action. Together, these methods and their downstream applications represent powerful tools to characterize the general metabolic phenotype of mice as well as to specifically assess alterations in glucose metabolism. They may be especially useful in the broad research field of insulin resistance, diabetes and obesity to provide a better understanding of pathogenesis as well as to test the effects of therapeutic interventions.

Study of In Vivo Glucose Metabolism in High-fat Diet-fed Mice Using Oral Glucose Tolerance Test OGTT and Insulin Tolerance Test ITT

The current article describes the generation and metabolic characterization of high-fat diet-fed mice as a model of diet-induced insulin resistance and obesity. It further features detailed protocols to perform the oral glucose tolerance test and the insulin tolerance test, monitoring whole-body alterations of glucose metabolism in vivo.

Obesity represents the most important single risk factor in the pathogenesis of type 2 diabetes, a disease which is characterized by a resistance to ...

Skeletal Muscle Neurovascular Coupling, Oxidative Capacity, and Microvascular Function with 'One Stop Shop' Near-infrared Spectroscopy

Exercise represents a major hemodynamic stress that demands a highly coordinated neurovascular response in order to match oxygen delivery to metabolic demand. Reactive hyperemia (in response to a brief period of tissue ischemia) is an independent predictor of cardiovascular events and provides important insight into vascular health and vasodilatory capacity. Skeletal muscle oxidative capacity is equally important in health and disease, as it determines the energy supply for myocellular processes. Here, we describe a simple, non-invasive approach using near-infrared spectroscopy to assess each of these major clinical endpoints (reactive hyperemia, neurovascular coupling, and muscle oxidative capacity) during a single clinic or laboratory visit. Unlike Doppler ultrasound, magnetic resonance images/spectroscopy, or invasive catheter-based flow measurements or muscle biopsies, our approach is less operator-dependent, low-cost, and completely non-invasive. Representative data from our lab taken together with summary data from previously published literature illustrate the utility of each of these end-points. Once this technique is mastered, application to clinical populations will provide important mechanistic insight into exercise intolerance and cardiovascular dysfunction.

Here, we describe a simple, non-invasive approach using near-infrared spectroscopy to assess reactive hyperemia, neurovascular coupling and skeletal muscle oxidative capacity in a single clinic or laboratory visit.

Exercise represents a major hemodynamic stress that demands a highly coordinated neurovascular response in order to match oxygen delivery to metabolic ...

Muscle Imbalances: Testing and Training Functional Eccentric Hamstring Strength in Athletic Populations

Many hamstring injuries that occur during physical activity occur while the muscles are lengthening, during eccentric hamstring muscle actions. Opposite of these eccentric hamstring actions are concentric quadriceps actions, where the larger and likely stronger quadriceps straighten the knee. Therefore, to stabilize the lower limbs during movement, the hamstrings must eccentrically combat against the strong knee-straightening torque of the quadriceps. As such, eccentric hamstring strength expressed relative to concentric quadricep strength is commonly referred to as the &#34;functional ratio&#34; as most movements in sports require simultaneous concentric knee extension and eccentric knee flexion. To increase the strength, resiliency, and functional performance of the hamstrings, it is necessary to test and train the hamstrings at different eccentric speeds. The main purpose of this work is to provide instructions for measuring and interpreting eccentric hamstring strength. Techniques for measuring the functional ratio using isokinetic dynamometry are provided and sample data will be compared. Additionally, we briefly describe how to address hamstring strength deficiencies or unilateral strength differences using exercises that specifically focus on increasing eccentric hamstring strength.

The hamstrings are a group of muscles that are sometimes problematic for athletes, resulting in soft tissue injury in the lower limbs. To prevent such injuries, functional training of the hamstrings requires intensive eccentric contractions. Additionally, hamstring function should be tested in relation to quadricep function at different contraction speeds.

Many hamstring injuries that occur during physical activity occur while the muscles are lengthening, during eccentric hamstring muscle actions. Opposite ...