Effects of Aging on Motor-Unit Control Properties

1999 ◽  
Vol 82 (5) ◽  
pp. 2081-2091 ◽  
Author(s):  
Zeynep Erim ◽  
M. Faisal Beg ◽  
David T. Burke ◽  
Carlo J. de Luca
Keyword(s):  
Firing Rate ◽  
Motor Unit ◽  
Motor Units ◽  
Elderly Subjects ◽  
Compensatory Mechanism ◽  
Type I ◽  
Firing Behavior ◽  
Firing Rates ◽  
Age Related ◽  
Young Subjects

It was hypothesized that the age-related alterations in the morphological properties of a motor unit would be accompanied by modifications in the control aspects of the motor unit, as either an adaptive or compensatory mechanism to preserve smooth force production. In specific, the objective of the study was to investigate the age-related alterations in the concurrent firing behavior of multiple motor units in the first dorsal interosseous (FDI) muscle in isometric contractions at 20 and 50% of the subject's voluntary contraction level. Analysis of the data collected from 10 young (24–37 yr of age) and 10 elderly (65–88 yr of age) subjects led to three novel observations regarding the firing behavior of aged motor units. 1) Among elderly subjects, there is a decrease in the common fluctuations that are observed among the firing rates of motor units in the young. 2) The relationship observed between the firing rate and recruitment threshold of young subjects is disturbed in the elderly. Although in young subjects, at any point in a given submaximal contraction, earlier recruited motor units have higher firing rates than later-recruited units; in aged subjects this dependency of firing rate on recruitment rank is compromised. 3) The progressive decrease observed in the firing rates of concurrently active motor units in constant-force contractions in the young is not seen in the aged. In addition to these original findings, this study provided support for earlier reports of 1) decreased average firing rates probably reflecting the slowing of the muscle, 2) a shift in recruitment thresholds toward lower force levels in line with the shift toward type I fibers, and 3) multiphasic action potential shapes indicative of the reinnervation process that takes place during aging. Taken as a whole, these findings indicate significant age-related modifications in the control properties of human motor units.

scholarly journals Blood Flow Restriction Alters Motor Unit Behavior During Resistance Exercise

2019 ◽  
Vol 40 (09) ◽  
pp. 555-562 ◽  
Author(s):  
Pedro Fatela ◽  
Goncalo V. Mendonca ◽  
António Prieto Veloso ◽  
Janne Avela ◽  
Pedro Mil-Homens
Keyword(s):  
Blood Flow ◽  
Firing Rate ◽  
Motor Unit ◽  
Motor Units ◽  
Blood Flow Restriction ◽  
Motor Unit Recruitment ◽  
Recruitment Threshold ◽  
Firing Rates ◽  
Flow Restriction ◽  
Voluntary Contractions

AbstractWe aimed to determine whether blood flow restriction (BFR) alters the characteristics of individual motor units during low-intensity (LI) exercise. Eight men (26.0±3.8 yrs) performed 5 sets of 15 knee extensions at 20% of one-repetition maximum (with and without BFR). Maximal isometric voluntary contractions (MVC) were performed before and after exercise to quantify force decrement. Submaximal isometric voluntary contractions were additionally performed for 18 s, matching trapezoidal target-force trajectories at 40% pre-MVC. EMG activity was recorded from the vastus lateralis muscle. Then, signals were decomposed to extract motor unit recruitment threshold, firing rates and action potential amplitudes (MUAP). Force decrement was only seen after LI BFR exercise (–20.5%; p<0.05). LI BFR exercise also induced greater decrements in the linear slope coefficient of the regression lines between motor unit recruitment threshold and firing rate (BFR: –165.1±120.4 vs. non-BFR: –44.4±33.1%, p<0.05). Finally, there was a notable shift towards higher values of firing rate and MUAP amplitude post-LI BFR exercise. Taken together, our data indicate that LI BFR exercise increases the activity of motor units with higher MUAP amplitude. They also indicate that motor units with similar MUAP amplitudes become activated at higher firing rates post-LI BFR exercise.

Activation of type-identified motor units during centrally evoked contractions in the cat medial gastrocnemius muscle. II. Motoneuron firing-rate modulation

1996 ◽  
Vol 75 (1) ◽  
pp. 38-50 ◽  
Author(s):  
K. E. Tansey ◽  
B. R. Botterman
Keyword(s):  
Firing Rate ◽  
Motor Unit ◽  
Muscle Force ◽  
Motor Units ◽  
Medial Gastrocnemius ◽  
Firing Rates ◽  
Rate Modulation ◽  
The Mean ◽  
Fast Twitch ◽  
The Relationship

1. The aim of this study was to examine the nature of motoneuron firing-rate modulation in type-identified motor units during smoothly graded contractions of the cat medial gastrocnemius (MG) muscle evoked by stimulation of the mesencephalic locomotor region (MLR). Motoneuron discharge patterns, firing rates, and the extent of firing-rate modulation in individual units were studied, as was the extent of concomitant changes in firing rates within pairs of simultaneously active units. 2. In 21 pairs of simultaneously active motor units, studied during 41 evoked contractions, the motoneurons' discharge rates and patterns were measured by processing the cells' recorded action potentials through windowing devices and storing their timing in computer memory. Once recruited, most motoneurons increased their firing rates over a limited range of increasing muscle tension and then maintained a fairly constant firing rate as muscle force continued to rise. Most motoneurons also decreased their firing rates over a slightly larger, but still limited, range of declining muscle force before they were derecruited. Although this was the most common discharge pattern recorded, several other interesting patterns were also seen. 3. The mean firing rate for slow twitch (type S) motor units (27.8 imp/s, 5,092 activations) was found to be significantly different from the mean firing rate for fast twitch (type F) motor units (48.4 imp/s, 11,272 activations; Student's t-test, P < 0.001). There was no significant difference between the mean firing rates of fast twitch, fatigue-resistant (type FR) and fast twitch, fatigable (type FF) motor units. When the relationship between motoneuron firing rate and whole-muscle force was analyzed, it was noted that, in general, smaller, lower threshold motor units began firing at lower rates and reached lower peak firing rates than did larger, higher threshold motor units. These results confirm both earlier experimental observations and predictions made by other investigators on the basis of computer simulations of the cat MG motor pool, but are in contrast to motor-unit discharge behavior recorded in some human motor-unit studies. 4. The extent of concomitant changes in firing rate within pairs of simultaneously active motor units was examined to estimate the extent of simultaneous motoneuron firing-rate modulation across the motoneuron pool. A smoothed (5 point sliding average) version of the two motoneurons' instantaneous firing rates was plotted against each other, and the slope and statistical significance of the relationship was determined. In 16 motor-unit pairs, the slope of the motoneurons' firing-rate relationship was significantly distinct from 0. Parallel firing-rate modulation (< 10-fold difference in firing rate change reflected by a slope of > 0.1) was noted only in pairs containing motor units of like physiological type and then only if they were of similar recruitment threshold. 5. Other investigators have demonstrated that changes in a motoneuron's "steady-state" firing rate predictably reflect changes in the amount of effective synaptic current that cell is receiving. The finding in the present study of limited parallel firing-rate modulation between simultaneously active motoneurons would suggest that changes in the synaptic drive to the various motoneurons of the pool is unevenly distributed. This finding, in addition to the findings of orderly motor-unit recruitment and the relationship between motor-unit recruitment threshold and motoneuron firing rate, cannot be adequately accommodated for by the existing models of the synaptic organization in motoneuron pools. Therefore a new model of the synaptic organization within the motoneuron pool has been proposed.

Hand Dominance and Motor Unit Firing Behavior

1998 ◽  
Vol 80 (3) ◽  
pp. 1373-1382 ◽  
Author(s):  
Alexander Adam ◽  
Carlo J. De Luca ◽  
Zeynep Erim
Keyword(s):  
Firing Rate ◽  
Motor Unit ◽  
Cross Correlation ◽  
Dominant Hand ◽  
Hand Dominance ◽  
Myoelectric Signal ◽  
Force Output ◽  
Firing Behavior ◽  
Firing Rates ◽  
Motor Unit Firing

Adam, Alexander, Carlo J. De Luca, and Zeynep Erim. Hand dominance and motor unit firing behavior. J. Neurophysiol. 80: 1373–1382, 1998. Daily preferential use was shown to alter physiological and mechanical properties of skeletal muscle. This study was aimed at revealing differences in the control strategy of muscle pairs in humans who show a clear preference for one hand. We compared the motor unit (MU) recruitment and firing behavior in the first dorsal interosseous (FDI) muscle of both hands in eight male volunteers whose hand preference was evaluated with the use of a standard questionnaire. Myoelectric signals were recorded while subjects isometrically abducted the index finger at 30% of the maximal voluntary contraction (MVC) force. A myoelectric signal decomposition technique was used to accurately identify MU firing times from the myoelectric signal. In MUs of the dominant hand, mean values for recruitment threshold, initial firing rate, average firing rate at target force, and discharge variability were lower when compared with the nondominant hand. Analysis of the cross-correlation between mean firing rate and muscle force revealed cross-correlation peaks of longer latency in the dominant hand than in the nondominant side. This lag of the force output with respect to fluctuations in the firing behavior of MUs is indicative of a greater mechanical delay in the dominant FDI muscle. MVC force was not significantly different across muscle pairs, but the variability of force at the submaximal target level was higher in the nondominant side. The presence of lower average firing rates, lower recruitment thresholds, and greater firing rate/force delay in the dominant hand is consistent with the notion of an increased percentage of slow twitch fibers in the preferentially used muscle, allowing twitch fusion and force buildup to occur at lower firing rates. It is suggested that a lifetime of preferred use may cause adaptations in the fiber composition of the dominant muscle such that the mechanical effectiveness of its MUs increased.

Doublet Discharges in Motoneurons of Young and Older Adults

2006 ◽  
Vol 95 (5) ◽  
pp. 2787-2795 ◽  
Author(s):  
Anita Christie ◽  
Gary Kamen
Keyword(s):  
Firing Rate ◽  
Motor Unit ◽  
Force Production ◽  
Motor Units ◽  
Voluntary Contraction ◽  
Older Individuals ◽  
Motor Unit Firing ◽  
Older Subjects ◽  
Young Subjects ◽  
Ramp Contractions

The purpose of this study was to investigate the occurrence of motor unit doublet discharges in young and older individuals at different rates of increasing force. Participants included eight young (21.9 ± 3.56 yr) and eight older (74.1 ± 8.79 yr) individuals, with equal numbers of males and females in each group. Motor unit activity was recorded from the tibialis anterior during isometric dorsiflexion using a four-wire needle electrode. Subjects performed three ramp contractions from zero to 50% maximal voluntary contraction (MVC) force at each of three rates: 10, 30, and 50% MVC/s. Overall, the occurrence of doublets was significantly higher in the young than in the older individuals. However, neither group showed differences in the occurrence of doublets across the three rates of force production. Doublet firings were observed in 45.6 (young) and 35.1% (old) of motor units at 10% MVC/s; 48.6 (young) and 22.5% (old) of motor units at 30% MVC/s; and 48.4 (young) and 31.4% (old) at 50% MVC/s. The maximal firing rate was significantly higher and the force at which the motor units were recruited was significantly lower for those units that fired doublets than those that did not. The force at which doublets occurred ranged from 3.42 to 50% MVC in the young subjects and from 0 (force onset) to 50% MVC in the older subjects. The results of this study suggest that the occurrence of doublets is dependent on both motor unit firing rate and force level. The lower incidence of doublets in older individuals may be attributable to changes in the intrinsic properties of the motoneurons with aging, which appear to play a role in doublet discharges.

scholarly journals Hierarchical control of motor units in voluntary contractions

2012 ◽  
Vol 107 (1) ◽  
pp. 178-195 ◽  
Author(s):  
Carlo J. De Luca ◽  
Paola Contessa
Keyword(s):  
Firing Rate ◽  
Motor Unit ◽  
Hierarchical Control ◽  
Motor Units ◽  
Exponential Functions ◽  
Size Principle ◽  
Recruitment Threshold ◽  
Firing Rates ◽  
Control Scheme ◽  
Voluntary Contractions

For the past five decades there has been wide acceptance of a relationship between the firing rate of motor units and the afterhyperpolarization of motoneurons. It has been promulgated that the higher-threshold, larger-soma, motoneurons fire faster than the lower-threshold, smaller-soma, motor units. This relationship was based on studies on anesthetized cats with electrically stimulated motoneurons. We questioned its applicability to motor unit control during voluntary contractions in humans. We found that during linearly force-increasing contractions, firing rates increased as exponential functions. At any time and force level, including at recruitment, the firing rate values were inversely related to the recruitment threshold of the motor unit. The time constants of the exponential functions were directly related to the recruitment threshold. From the Henneman size principle it follows that the characteristics of the firing rates are also related to the size of the soma. The “firing rate spectrum” presents a beautifully simple control scheme in which, at any given time or force, the firing rate value of earlier-recruited motor units is greater than that of later-recruited motor units. This hierarchical control scheme describes a mechanism that provides an effective economy of force generation for the earlier-recruited lower force-twitch motor units, and reduces the fatigue of later-recruited higher force-twitch motor units—both characteristics being well suited for generating and sustaining force during the fight-or-flight response.

scholarly journals Neural control of muscle force: indications from a simulation model

2013 ◽  
Vol 109 (6) ◽  
pp. 1548-1570 ◽  
Author(s):  
Paola Contessa ◽  
Carlo J. De Luca
Keyword(s):  
Motor Unit ◽  
Neural Control ◽  
Muscle Force ◽  
Force Production ◽  
Motor Units ◽  
Isometric Contractions ◽  
Alternative Mechanism ◽  
Firing Behavior ◽  
Firing Rates ◽  
Muscle Force Production

We developed a model to investigate the influence of the muscle force twitch on the simulated firing behavior of motoneurons and muscle force production during voluntary isometric contractions. The input consists of an excitatory signal common to all the motor units in the pool of a muscle, consistent with the “common drive” property. Motor units respond with a hierarchically structured firing behavior wherein at any time and force, firing rates are inversely proportional to recruitment threshold, as described by the “onion skin” property. Time- and force-dependent changes in muscle force production are introduced by varying the motor unit force twitches as a function of time or by varying the number of active motor units. A force feedback adjusts the input excitation, maintaining the simulated force at a target level. The simulations replicate motor unit behavior characteristics similar to those reported in previous empirical studies of sustained contractions: 1) the initial decrease and subsequent increase of firing rates, 2) the derecruitment and recruitment of motor units throughout sustained contractions, and 3) the continual increase in the force fluctuation caused by the progressive recruitment of larger motor units. The model cautions the use of motor unit behavior at recruitment and derecruitment without consideration of changes in the muscle force generation capacity. It describes an alternative mechanism for the reserve capacity of motor units to generate extraordinary force. It supports the hypothesis that the control of motoneurons remains invariant during force-varying and sustained isometric contractions.

scholarly journals Contribution from motor unit firing adaptations and muscle coactivation during fatigue

2018 ◽  
Vol 119 (6) ◽  
pp. 2186-2193 ◽  
Author(s):  
Paola Contessa ◽  
John Letizi ◽  
Gianluca De Luca ◽  
Joshua C. Kline
Keyword(s):  
Motor Unit ◽  
Late Phase ◽  
Motor Units ◽  
Force Output ◽  
Firing Behavior ◽  
Firing Rates ◽  
Pronator Teres ◽  
Motor Unit Firing ◽  
The Central Nervous System ◽  
Extensor Carpi Radialis

The control of motor unit firing behavior during fatigue is still debated in the literature. Most studies agree that the central nervous system increases the excitation to the motoneuron pool to compensate for decreased force contributions of individual motor units and sustain muscle force output during fatigue. However, some studies claim that motor units may decrease their firing rates despite increased excitation, contradicting the direct relationship between firing rates and excitation that governs the voluntary control of motor units. To investigate whether the control of motor units in fact changes with fatigue, we measured motor unit firing behavior during repeated contractions of the first dorsal interosseous (FDI) muscle while concurrently monitoring the activation of surrounding muscles, including the flexor carpi radialis, extensor carpi radialis, and pronator teres. Across all subjects, we observed an overall increase in FDI activation and motor unit firing rates by the end of the fatigue task. However, in some subjects we observed increases in FDI activation and motor unit firing rates only during the initial phase of the fatigue task, followed by subsequent decreases during the late phase of the fatigue task while the coactivation of surrounding muscles increased. These findings indicate that the strategy for sustaining force output may occasionally change, leading to increases in the relative activation of surrounding muscles while the excitation to the fatiguing muscle decreases. Importantly, irrespective of changes in the strategy for sustaining force output, the control properties regulating motor unit firing behavior remain unchanged during fatigue. NEW & NOTEWORTHY This work addresses sources of debate surrounding the manner in which motor unit firing behavior is controlled during fatigue. We found that decreases in the motor unit firing rates of the fatiguing muscle may occasionally be observed when the contribution of coactive muscles increases. Despite changes in the strategy employed to sustain the force output, the underlying control properties regulating motor unit firing behavior remain unchanged during muscle fatigue.

Motor unit control properties in constant-force isometric contractions

1996 ◽  
Vol 76 (3) ◽  
pp. 1503-1516 ◽  
Author(s):  
C. J. de Luca ◽  
P. J. Foley ◽  
Z. Erim
Keyword(s):  
Firing Rate ◽  
Motor Unit ◽  
Motor Units ◽  
Motor Unit Recruitment ◽  
Constant Force ◽  
Force Output ◽  
Myoelectric Signals ◽  
Firing Rates ◽  
Twitch Potentiation ◽  
Maximal Effort

1. The purpose of this study was 1) to characterize the decrease observed in mean firing rates of motor units in the first 8-15 s of isometric constant-force contractions and 2) to investigate possible mechanisms that could account for the ability to maintain force output in the presence of decreasing motor unit firing rates. 2. The decrease in mean firing rates was characterized by investigating myoelectric signals detected with a specialized quadrifilar needle electrode from the first dorsal interosseus (FDI) and the tibialis anterior (TA) muscles of 19 healthy subjects during a total of 85 constant-force isometric contractions at 30, 50, or 80% of maximal effort. The firing times of motor units were obtained from the myoelectric signals with the use of computer algorithms to decompose the signal into the constituent motor unit action potentials. Time-varying mean firing rates and recruitment thresholds were also calculated. 3. Motor units detected from the TA muscle were found to have a continual decrease in their mean firing rates in 36 of 44 trials performed during isometric ankle dorsiflexion at force values ranging from 30 to 80% of maximal effort and a duration of 8-15 s. Likewise, motor units detected in the FDI muscle displayed a decrease in firing rate in 32 of 41 trials performed during constant-force isometric index finger abduction for contractions ranging from 30 to 80% of maximal effort. In 14 contractions (16% of total), firing rates were essentially constant, whereas in 3 contractions (4%), firing rates appeared to increase. 4. Motor units with the higher recruitment thresholds and lower firing rates tended to display the greater decreases in firing rate over the constant-force interval, whereas motor units with lower recruitment thresholds and higher firing rates had lesser rates of decrease. Furthermore, increasing contraction levels tended to intensify the decrease in the motor unit firing rates. 5. Three possible mechanisms were considered as factors responsible for the maintaining of force output while motor units decreased their firing rates: motor unit recruitment, agonist/antagonist interaction, and twitch potentiation. Of these, motor unit recruitment was discarded first because none was observed during the 8-15 s duration of any of the 85 contractions. Furthermore, contractions outside the physiological range of motor unit recruitment (at 80% of maximal effort) revealed the same decreasing trend in firing rates, ruling out recruitment as the means of sustaining force output. 6. The role of agonist or antagonist muscle interaction was investigated with the use of the muscles controlling the wrist joint. Myoelectric signals were recorded with quadrifilar needle electrodes from the wrist extensor muscles while myoelectric activity in the wrist flexor muscles was concurrently monitored with surface electrodes during constant-force isometric wrist extension at 50% of maximal effort. Firing rates of the motor units in the wrist extensor muscles simultaneously decreased while the flexor muscles were determined to be inactive. 7. All the findings of this study regarding the behavior of the firing rates could be well explained by the reported characteristics of twitch potentiation that have been previously documented in animals and humans. 8. The results of this study, combined with the results of other investigators, provide the following scenario to explain how a constant-force isometric contraction is sustained. As the contraction progresses, the twitch force of the muscle fibers undergoes a potentiation followed by a decrease. Simultaneously, the "late adaptation" property of the motoneuron decreases the firing rate of the motor unit. Findings of this study suggest that voluntary reduction in firing rates also cannot be ruled out as a means to augment the adaptation in motoneurons. (ABSTRACT TRUNCATED)

scholarly journals Bioengineered model of the human motor unit with physiologically functional neuromuscular junctions

2021 ◽  
Vol 11 (1) ◽  
Author(s):  
Rowan P. Rimington ◽  
Jacob W. Fleming ◽  
Andrew J. Capel ◽  
Patrick C. Wheeler ◽  
Mark P. Lewis
Keyword(s):  
Neuromuscular Junction ◽  
Motor Neuron ◽  
Motor Unit ◽  
Motor Nerve ◽  
Motor Units ◽  
Extracellular Matrices ◽  
Synaptic Membrane ◽  
Type I ◽  
Dependent Manner ◽  
Human Motor

AbstractInvestigations of the human neuromuscular junction (NMJ) have predominately utilised experimental animals, model organisms, or monolayer cell cultures that fail to represent the physiological complexity of the synapse. Consequently, there remains a paucity of data regarding the development of the human NMJ and a lack of systems that enable investigation of the motor unit. This work addresses this need, providing the methodologies to bioengineer 3D models of the human motor unit. Spheroid culture of iPSC derived motor neuron progenitors augmented the transcription of OLIG2, ISLET1 and SMI32 motor neuron mRNAs ~ 400, ~ 150 and ~ 200-fold respectively compared to monolayer equivalents. Axon projections of adhered spheroids exceeded 1000 μm in monolayer, with transcription of SMI32 and VACHT mRNAs further enhanced by addition to 3D extracellular matrices in a type I collagen concentration dependent manner. Bioengineered skeletal muscles produced functional tetanic and twitch profiles, demonstrated increased acetylcholine receptor (AChR) clustering and transcription of MUSK and LRP4 mRNAs, indicating enhanced organisation of the post-synaptic membrane. The number of motor neuron spheroids, or motor pool, required to functionally innervate 3D muscle tissues was then determined, generating functional human NMJs that evidence pre- and post-synaptic membrane and motor nerve axon co-localisation. Spontaneous firing was significantly elevated in 3D motor units, confirmed to be driven by the motor nerve via antagonistic inhibition of the AChR. Functional analysis outlined decreased time to peak twitch and half relaxation times, indicating enhanced physiology of excitation contraction coupling in innervated motor units. Our findings provide the methods to maximise the maturity of both iPSC motor neurons and primary human skeletal muscle, utilising cell type specific extracellular matrices and developmental timelines to bioengineer the human motor unit for the study of neuromuscular junction physiology.

scholarly journals Extra Virgin Olive Oil Prevents the Age-Related Shifts of the Distribution of HDL Subclasses and Improves Their Functionality

Nutrients ◽  
2021 ◽  
Vol 13 (7) ◽  
pp. 2235
Author(s):  
Alyann Otrante ◽  
Amal Trigui ◽  
Roua Walha ◽  
Hicham Berrougui ◽  
Tamas Fulop ◽  
...  
Keyword(s):  
Olive Oil ◽  
Virgin Olive Oil ◽  
Density Lipoprotein ◽  
Extra Virgin Olive Oil ◽  
The Elderly ◽  
High Density ◽  
Lipoprotein Cholesterol ◽  
Elderly Subjects ◽  
Age Related ◽  
Young Subjects

High-density lipoproteins (HDL) maintain cholesterol homeostasis through the role they play in regulating reverse cholesterol transport (RCT), a process by which excess cholesterol is transported back to the liver for elimination. However, RCT can be altered in the presence of cardiovascular risk factors, such as aging, which contributes to the increase in the incidence of cardiovascular diseases (CVD). The present study was aimed at investigating the effect of extra virgin olive oil (EVOO) intake on the cholesterol efflux capacity (CEC) of HDL, and to elucidate on the mechanisms by which EVOO intake improves the anti-atherogenic activity of HDL. A total of 84 healthy women and men were enrolled and were distributed, according to age, into two groups: 27 young (31.81 ± 6.79 years) and 57 elderly (70.72 ± 5.6 years) subjects. The subjects in both groups were given 25 mL/d of extra virgin olive oil (EVOO) for 12 weeks. CEC was measured using J774 macrophages radiolabeled with tritiated cholesterol ((3H) cholesterol). HDL subclass distributions were analyzed using the Quantimetrix Lipoprint® system. The HDL from the elderly subjects exhibited a lower level of CEC, at 11.12% (p < 0.0001), than the HDL from the young subjects. The CEC of the elderly subjects returned to normal levels following 12 weeks of EVOO intake. An analysis of the distribution of HDL subclasses showed that HDL from the elderly subjects were composed of lower levels of large HDL (L-HDL) (p < 0.03) and higher levels of small HDL (S-HDL) (p < 0.002) compared to HDL from the young subjects. A multiple linear regression analysis revealed a positive correlation between CEC and L-HDL levels (r = 0.35 and p < 0.001) as well as an inverse correlation between CEC and S-HDL levels (r = −0.27 and p < 0.01). This correlation remained significant even when several variables, including age, sex, and BMI as well as low-density lipoprotein cholesterol (LDL-C), high-density lipoprotein cholesterol (HDL-C), and glucose levels (β = 0.28, p < 0.002, and β = 0.24, p = 0.01) were accounted for. Consuming EVOO for 12 weeks modulated the age-related difference in the distribution of HDL subclasses by reducing the level of S-HDL and increasing the level of intermediate-HDL/large-HDL (I-HDL/L-HDL) in the elderly subjects. The age-related alteration of the CEC of HDL was due, in part, to an alteration in the distribution of HDL subclasses. A diet enriched in EVOO improved the functionality of HDL through an increase in I-HDL/L-HDL and a decrease in S-HDL.

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