Predicted threshold against forward and backward loss of balance for perturbed walking

The biomechanical mechanisms of loss of balance have been studied before for slip condition but have not been investigated for arbitrary perturbation profiles under non-slip conditions in sagittal plane. This study aimed to determine the thresholds of center of mass (COM) velocity and position relative to the base of support (BOS) that predict forward and backward loss of balance during walking with a range of BOS perturbations. Perturbations were modeled as sinusoidal BOS motions in the vertical or anterior-posterior direction or as sagittal rotation. The human body was modeled using a seven-link model. Forward dynamics alongside with dynamic optimization were used to find the thresholds of initial COM velocity for each initial COM position that would predict forward or backward loss of balance. The effects of perturbation frequency and amplitude on these thresholds were modeled based on the simulation data. Experimental data were collected from 15 able-bodied individuals and three individuals with disability during perturbed walking. The simulation results showed similarity with the stability region reported for slip and non-slip conditions. The feasible stability region shrank when the perturbation frequency and amplitude increased, especially for larger initial COM velocities. 89.5% (70.9%) and 82.4% (68.2%) of the measured COM position and velocity combinations during low (high) perturbations were located inside the simulated limits of the stability region, for able-bodied and disabled individuals, respectively. The simulation results demonstrated the effects of different perturbation levels on the stability region. The obtained stability region can be used for developing rehabilitative programs in interactive environments.     

The independent effects of speed and propulsive force on joint power generation in walking

Walking speed is modulated using propulsive forces (F_P) during push-off and both preferred speed and F_P decrease with aging. However, even prior to walking slower, reduced F_P may be accompanied by potentially unfavorable changes in joint power generation. For example, compared to young adults, older adults exhibit a redistribution of mechanical power generation from the propulsive plantarflexor muscles to more proximal muscles acting across the knee and hip. Here, we used visual biofeedback based on real-time F_P measurements to decouple and investigate the interaction between joint-level coordination, whole-body F_P, and walking speed. 12 healthy young subjects walked on a dual-belt instrumented treadmill at a range of speeds (0.9–1.3 m/s). We immediately calculated the average F_P from each speed. Subjects then walked at 1.3 m/s while completing a series of biofeedback trials with instructions to match their instantaneous F_P to their averaged F_P from slower speeds. Walking slower decreased F_P and total positive joint work with little effect on relative joint-level contributions. Conversely, subjects walked at a constant speed with reduced F_P, not by reducing total positive joint work, but by redistributing the mechanical demands of each step from the plantarflexor muscles during push-off to more proximal leg muscles during single support. Interestingly, these naturally emergent joint- and limb-level biomechanical changes, in the absence of neuromuscular constraints, resemble those due to aging. Our findings provide important reference data to understand the presumably complex interactions between joint power generation, whole-body F_P, and walking speed in our aging population.     

A comparison of muscle energy models for simulating human walking in three dimensions

The popular Hill model for muscle activation and contractile dynamics has been extended with several different formulations for predicting the metabolic energy expenditure of human muscle actions. These extended models differ considerably in their approach to computing energy expenditure, particularly in their treatment of active lengthening and eccentric work, but their predictive abilities have never been compared. In this study, we compared the predictions of five different Hill-based muscle energy models in 3D forward dynamics simulations of normal human walking. In a data-tracking simulation that minimized muscle fatigue, the energy models predicted metabolic costs that varied over a three-fold range (2.45–7.15 J/m/kg), with the distinction arising from whether or not eccentric work was subtracted from the net heat rate in the calculation of the muscle metabolic rate. In predictive simulations that optimized neuromuscular control to minimize the metabolic cost, all five models predicted similar speeds, step lengths, and stance phase durations. However, some of the models predicted a hip circumduction strategy to minimize metabolic cost, while others did not, and the accuracy of the predicted knee and ankle angles and ground reaction forces also depended on the energy model used. The results highlights the need to clarify how eccentric work should be treated when calculating muscle energy expenditure, the difficulty in predicting realistic metabolic costs in simulated walking even with a detailed 3D musculoskeletal model, the potential for using such models to predict energetically-optimal gait modifications, and the room for improvement in existing muscle energy models and locomotion simulation frameworks.     

The effect of speed and influence of individual muscles on hamstring mechanics during the swing phase of sprinting

The purpose of this study was to characterize the effect of speed and influence of individual muscles on hamstring stretch, loading, and work during the swing phase of sprinting. We measured three-dimensional kinematics and electromyography (EMG) activities of 19 athletes sprinting on a treadmill at speeds ranging from 80% to 100% of maximum speed. We then generated muscle-actuated forward dynamic simulations of swing and double float phases of the sprinting gait cycle. Simulated lower extremity joint angles and model predicted excitations were similar to measured quantities. Swing phase simulations were used to characterize the effects of speed on the peak stretch, maximum force, and negative work of the biceps femoris long head (BF), the most often injured hamstring muscle. Perturbations of the double float simulations were used to assess the influence of individual muscles on BF stretch.

Peak hamstring musculotendon stretch occurred at 90% of the gait cycle (late swing) and was independent of speed. Peak hamstring force and negative musculotendon work increased significantly with speed (p<0.05). Muscles in the lumbo-pelvic region had greater influence on hamstring stretch than muscles acting about the knee and ankle. In particular, the hip flexors were found to induce substantial hamstring stretch in the opposite limb, with that influence increasing with running speed. We conclude that hamstring strain injury during sprinting may be related to the performance of large amounts of negative work over repeated strides and/or resulting from a perturbation in pelvic muscle coordination that induces excessive hamstring stretch in a single stride.     

Effects of long-term immobilization and recovery on human triceps surae and collagen turnover in the Achilles tendon in patients with healing ankle fracture.

The aim of the present study was to analyze how human tendon connective tissue responds to an approximately 7-wk period of immobilization and a remobilization period of a similar length, in patients with unilateral ankle fracture, which is currently unknown. Calf muscle cross-sectional area (CSA) decreased by 15% (5,316 to 4,517 mm2) and strength by 54% (239 to 110 N.m) in the immobilized leg after 7 wk. During the 7-wk remobilization, the CSA increased by 9% (to 4,943 mm2) and strength by 37% (to 176 Nm). Achilles tendon CSA did not change significantly during either immobilization or remobilization. Local collagen turnover was measured as the peritendinous concentrations of NH2-terminal propeptide of type I collagen (PINP) and COOH-terminal telopeptide region of type I collagen (ICTP), markers thought to be indexes of type I collagen synthesis and degradation, respectively. Both markers were increased (PINP: 257 vs. 56 ng/ml; ICTP: 9.8 vs. 2.1 microg/l) in the immobilized leg compared with the control leg after the 7 wk of immobilization, and levels decreased again in the immobilized leg during the recovery period (PINP: 103 vs. 44 ng/ml; ICTP: 4.2 vs. 1.9 microg/l). A significant reduction in calf muscle CSA and strength was found in relation to 7 wk of immobilization. Immobilization increased both collagen synthesis and degradation in tendon near tissue. However, it cannot be excluded that the facture of the ankle in close proximity could have affected these data. Remobilization increased muscle size and strength and tendon synthesis and degradation decreased to baseline levels. These dynamic changes in tendon connective tissue turnover were not associated with macroscopic changes in tendon size.     

Impact of ankle foot orthosis stiffness on Achilles tendon and gastrocnemius function during unimpaired gait

Ankle foot orthoses (AFOs) are designed to improve gait for individuals with neuromuscular conditions and have also been used to reduce energy costs of walking for unimpaired individuals. AFOs influence joint motion and metabolic cost, but how they impact muscle function remains unclear. This study investigated the impact of different stiffness AFOs on medial gastrocnemius muscle (MG) and Achilles tendon (AT) function during two walking speeds. We performed gait analyses for eight unimpaired individuals. Each individual walked at slow and very slow speeds with a 3D printed AFO with no resistance (free hinge condition) and four levels of ankle dorsiflexion stiffness: 0.25 Nm/°, 1 Nm/°, 2 Nm/°, and 3.7 Nm/°. Motion capture, ultrasound, and musculoskeletal modeling were used to quantify MG and AT lengths with each AFO condition. Increasing AFO stiffness increased peak AFO dorsiflexion moment with decreased peak knee extension and peak ankle dorsiflexion angles. Overall musculotendon length and peak AT length decreased, while peak MG length increased with increasing AFO stiffness. Peak MG activity, length, and velocity significantly decreased with slower walking speed. This study provides experimental evidence of the impact of AFO stiffness and walking speed on joint kinematics and musculotendon function. These methods can provide insight to improve AFO designs and optimize musculotendon function for rehabilitation, performance, or other goals.     

Contribution of muscle short-range stiffness to initial changes in joint kinetics and kinematics during perturbations to standing balance: A simulation study

Simulating realistic musculoskeletal dynamics is critical to understanding neural control of muscle activity evoked in sensorimotor feedback responses that have inherent neural transmission delays. Thus, the initial mechanical response of muscles to perturbations in the absence of any change in muscle activity determines which corrective neural responses are required to stabilize body posture. Muscle short-range stiffness, a history-dependent property of muscle that causes a rapid and transient rise in muscle force upon stretch, likely affects musculoskeletal dynamics in the initial mechanical response to perturbations. Here we identified the contributions of short-range stiffness to joint torques and angles in the initial mechanical response to support surface translations using dynamic simulation. We developed a dynamic model of muscle short-range stiffness to augment a Hill-type muscle model. Our simulations show that short-range stiffness can provide stability against external perturbations during the neuromechanical response delay. Assuming constant muscle activation during the initial mechanical response, including muscle short-range stiffness was necessary to account for the rapid rise in experimental sagittal plane knee and hip joint torques that occurs simultaneously with very small changes in joint angles and reduced root mean square errors between simulated and experimental torques by 56% and 47%, respectively. Moreover, forward simulations lacking short-range stiffness produced unreasonably large joint angle changes during the initial response. Using muscle models accounting for short-range stiffness along with other aspects of history-dependent muscle dynamics may be important to advance our ability to simulate inherently unstable human movements based on principles of neural control and biomechanics.     

More than energy cost: multiple benefits of the long Achilles tendon in human walking and running

Elastic strain energy that is stored and released from long, distal tendons such as the Achilles during locomotion allows for muscle power amplification as well as for reduction of the locomotor energy cost: as distal tendons perform mechanical work during recoil, plantar flexor muscle fibres can work over smaller length ranges, at slower shortening speeds, and at lower activation levels. Scant evidence exists that long distal tendons evolved in humans (or were retained from our more distant Hominoidea ancestors) primarily to allow high muscle–tendon power outputs, and indeed we remain relatively powerless compared to many other species. Instead, the majority of evidence suggests that such tendons evolved to reduce total locomotor energy cost. However, numerous additional, often unrecognised, advantages of long tendons may speculatively be of greater evolutionary advantage, including the reduced limb inertia afforded by shorter and lighter muscles (reducing proximal muscle force requirement), reduced energy dissipation during the foot–ground collisions, capacity to store and reuse the muscle work done to dampen the vibrations triggered by foot–ground collisions, reduced muscle heat production (and thus core temperature), and attenuation of work-induced muscle damage. Cumulatively, these effects should reduce both neuromotor fatigue and sense of locomotor effort, allowing humans to choose to move at faster speeds for longer. As these benefits are greater at faster locomotor speeds, they are consistent with the hypothesis that running gaits used by our ancestors may have exerted substantial evolutionary pressure on Achilles tendon length. The long Achilles tendon may therefore be a singular adaptation that provided numerous physiological, biomechanical, and psychological benefits and thus influenced behaviour across multiple tasks, both including and additional to locomotion. While energy cost may be a variable of interest in locomotor studies, future research should consider the broader range of factors influencing our movement capacity, including our decision to move over given distances at specific speeds, in order to understand more fully the effects of Achilles tendon function as well as changes in this function in response to physical activity, inactivity, disuse and disease, on movement performance.     

The effects of athletic training and muscle contractile character on the pressor response to isometric exercise of the human triceps surae

In this study, the influence of athletic training status and the contractile character of the active muscle on the magnitude of the pressor response (PR) to voluntary and electrically evoked isometric plantar flexion was investigated. Subjects were 10 sprint-trained athletes (sprint) (100-m, 200-m and 400-m) [mean (SD) age, 21 (2) years], 14 endurance trained athletes (distance) [22 (2) years] and 8 untrained men (control) [23 (3) years]. Twitch time to peak tension (TPT) in the sprint group [108 (7) ms] was significantly less (P<0.001) than that of the distance group [124 (10) ms]. During voluntary contraction, the mean change in systolic blood pressure (SBP), diastolic blood pressure (DBP) and heart rate (fc) was not significantly different between groups. During electrically evoked contractions, mean changes in SBP, DBP and fc were not significantly different between the sprint, distance and control groups. However, division of the sprint group into 400-m (sprint I) and 100/200-m athletes (sprint II) showed that an increase in DBP of 1.6 kPa (12 mm Hg) in sprint I was significantly less (P<0.05) than the 2.5 kPa (19 mm Hg) increase observed for both the distance and control groups. Prediction of the DBP response from our previously published relationship between TPT and DBP showed close agreement in all subject groups except sprint I; in these subjects the observed DBP response was only 55% of that predicted. Attenuation of the PR in the involuntary experiment suggests that some aspect of sprint training, but not endurance training, modifies the muscle afferent input to the PR in man.     

Molecular dynamics study on the conformational flexibility and energetics in aqueous solution of methylated beta-cyclodextrins

All possible methylated beta-cyclodextrins (CDs) with C7-symmetry have been studied by molecular dynamics simulations, in gas phase and in water solution. Energetic and structural information were obtained from the trajectory analysis. CD flexibility increases with degree of methylation, very likely due to the concomitant reduction of the intramolecular hydrogen bonds. Solvation-free energy was computed for each of the studied CDs using the MM/GBSA method. An analysis of radial distribution functions was used to determine distribution of solvent molecules around the O2, O3, and O6. The number of solvent molecules around these oxygens decreases with an increase in the degree of methylation. The DeltaS contribution from solvent thus becomes more positive when the degree of methylation increases and, consequently, the overall DeltaG in water diminishes.     

Power in sports: A literature review on the application,assumptions, and terminology of mechanical power in sport research

The quantification of mechanical power can provide valuable insight into athlete performance because it is the mechanical principle of the rate at which the athlete does work or transfers energy to complete a movement task. Estimates of power are usually limited by the capabilities of measurement systems, resulting in the use of simplified power models. This review provides a systematic overview of the studies on mechanical power in sports, discussing the application and estimation of mechanical power, the consequences of simplifications, and the terminology. The mechanical power balance consists of five parts, where joint power is equal to the sum of kinetic power, gravitational power, environmental power, and frictional power. Structuring literature based on these power components shows that simplifications in models are done on four levels, single vs multibody models, instantaneous power (IN) versus change in energy (EN), the dimensions of a model (1D, 2D, 3D), and neglecting parts of the mechanical power balance. Quantifying the consequences of simplification of power models has only been done for running, and shows differences ranging from 10% up to 250% compared to joint power models. Furthermore, inconsistency and imprecision were found in the determination of joint power, resulting from inverse dynamics methods, incorporation of translational joint powers, partitioning in negative and positive work, and power flow between segments. Most inconsistency in terminology was found in the definition and application of ‘external’ and ‘internal’ work and power. Sport research would benefit from structuring the research on mechanical power in sports and quantifying the result of simplifications in mechanical power estimations.     

The effect of Knee-Ankle-Foot orthosis stiffness on the parameters of walking

The purpose of this simulation study was to analyze the effect of variation in Knee-Ankle-Foot-Orthosis stiffness on the joint power and the energy cost of walking. The effect of contractile tissue was simulated using linear elastic spring and viscous dampers in knee and ankle joints. Then, joint angles, ground reaction force, were collected from Twenty chronic hemiparesis subjects (15 males and 5 females) and twenty control subjects (14 males and 6 females), and spring stiffness were considered as the inputs. In this new study, the generated muscle torques were optimized by changing the stiffness as the desired output in the mathematical model attained by the MATLAB SimMechanics toolbox. Finally, the simulated mathematical model was introduced as an appropriate substitute in obtaining the optimized stiffness with a more convenient and efficient designed orthosis.     

Acute effects of capacitive and resistive electric transfer (CRet) on the Achilles tendon

This study aimed to investigate the acute effects of capacitive and resistive electric transfer (CRet) on Achilles tendon elongation during muscle contraction, as well as the circulation in the peritendinous region. Sixteen healthy men participated in this study. All 16 participants underwent 2 interventions: (1) CRet trial and (2) CRet without power (sham trial). Tendon elongation was measured four times. Using near-infrared spectroscopy, the blood circulation (volume of total-hemoglobin (Hb), oxygenated hemoglobin (oxy-Hb), and deoxygenated hemoglobin (deoxy-Hb)) was measured for 5 min before the intervention and for 30 min after the intervention. The differences between the measurements obtained before and after intervention were compared between the two interventions. The changes in tendon elongation and deoxy-Hb were not significantly different between the interventions. Total- and oxy-Hb were significantly increased in the CRet trial compared with the sham trial. In addition, the increases in total-Hb and oxy-Hb lasted for 30 min after the CRet intervention (CRet vs. sham: oxy-Hb: F = 8.063, p = 0.001, total-Hb: F = 4.564, p = 0.011). In conclusion, CRet significantly improved blood circulation in the peritendinous region.     

The effect of beta-adrenergic blockade on leg blood flow with repeated maximal contractions of the triceps surae muscle group in man

The effect of beta-adrenergic blockade on torque output and leg blood flow was examined in seven healthy young men during repeated maximal isometric voluntary contractions of the triceps surae muscle group. Exercise was performed in either a bent- or straight-leg position during each of four drug treatments: placebo, propranolol, metoprolol, oxprenolol. Contractions were sustained for 5 s with 5 s relaxation for a total of 10 min followed by a 10-min recovery. Leg blood flow was measured during the 5 s relaxation separating contractions using strain gauge plethysmography. Torque output decreased during the 10-min contractions with no differences between the four drug treatments. Leg blood flow was lower with beta-blockade during the initial stages of exercise and recovery in the bent-leg position but no differences were observed after 3 min exercise or recovery. Leg blood flow in the straight-leg position was not different between any of the four drug treatments, but it was significantly less than in bent-leg exercise. The lower blood flows during the initial stages of exercise in the beta-blocked conditions probably reflect a slowing of the central cardiovascular response because of beta 1-receptor blockade of the heart rather than on the beta 2-receptors effects on peripheral vascular resistance. It is concluded that local vasodilator substances released from the working muscle may play a more important role than beta 2-receptor stimulation of smooth muscle in skeletal muscle resistance vessels in regulating local muscle blood flow during maximal exercise of the triceps surae muscle group.     

The energetics of reproduction and parental care in the terrestrial isopod Porcellio laevis

Parental care is a behavioral strategy that contributes to increased fitness of progeny. Among terrestrial arthropods, many isopods provide extensive parental care. Few studies have quantified the underlying cost of parental care in terms of energy. We used the terrestrial woodlouse Porcellio laevis (Latreille) as a study model to examine how energetic acquisition and expenditure in females is affected during the incubation period and how parental care affects energy balance in this species. We determined the basic reproductive biology (i.e. fecundity, reproductive output, egg volume, egg loss), energy expenditure (i.e. metabolic rate), and energy acquisition (i.e. food consumption, digestibility) of ovigerous females in different stages of embryonic development. Non-ovigerous females were used as the control group. Our results show that P. laevis displays variability in life-history traits compared with populations from other zones around the world. Ovigerous females exhibited a lower ingestion rate and lower digestibility than control females, thus indicating a lower capacity for energy acquisition. Furthermore, energy expenditure was higher in ovigerous females when compared to non-ovigerous females. In particular, females in early embryonic development stored 5.1-fold less daily energy than females without eggs.

The results presented here show that the parental care provided by female P. laevis is energetically costly. Overall, our work brings us much closer to understanding the proximate mechanisms of the costs of parental care in terrestrial isopods. Both proximal mechanisms and consequences of providing care on future reproduction, should be considered in explaining the evolution of parental care.     

The regulation of aggrecanase ADAMTS-4 expression in human Achilles tendon and tendon-derived cells

Several members of the ADAMTS (A Disintegrin And Metalloproteinase with ThromboSpondin motifs) family have been identified as aggrecanases, whose substrates include versican, the principal large proteoglycan in the tendon extracellular matrix. We have characterized the expression of ADAMTS-4 in human Achilles tendon and tendon-derived cells. ADAMTS-4 mRNA levels were higher in ruptured tendon compared with normal tendon or chronic painful tendinopathy. In tissue extracts probed by Western blotting, mature ADAMTS-4 (68 kDa) was detected only in ruptured tendons, while processed ADAMTS-4 (53 kDa) was detected also in chronic painful tendinopathy and in normal tendon. In cultured Achilles tendon cells, transforming growth factor-β (TGF-β) stimulated ADAMTS-4 mRNA expression (typically 20-fold after 24 h), while interleukin-1 induced a smaller, shorter-term stimulation which synergised markedly with that induced by TGF-β. Increased levels of immunoreactive proteins consistent with mature and processed forms of ADAMTS-4 were detected in TGF-β-stimulated cells. ADAMTS-4 mRNA was expressed at higher levels by tendon cells in collagen gels than in monolayer cultures. In contrast, the expression of ADAMTS-1 and -5 mRNA was lower in collagen gels compared with monolayers, and these mRNA showed smaller or opposite responses to growth factors and cytokines compared with that of ADAMTS-4 mRNA. We conclude that both ADAMTS-4 mRNA and ADAMTS-4 protein processing may be differentially regulated in normal and damaged tendons and that both the matrix environment and growth factors such as TGF-β are potentially important factors controlling ADAMTS aggrecanase activities in tendon pathology.     

The energetics and evolution of oxidoreductases in deep time

The core metabolic reactions of life drive electrons through a class of redox protein enzymes, the oxidoreductases. The energetics of electron flow is determined by the redox potentials of organic and inorganic cofactors as tuned by the protein environment. Understanding how protein structure affects oxidation–reduction energetics is crucial for studying metabolism, creating bioelectronic systems, and tracing the history of biological energy utilization on Earth. We constructed ProtReDox ( https://protein-redox-potential.web.app ), a manually curated database of experimentally determined redox potentials. With over 500 measurements, we can begin to identify how proteins modulate oxidation–reduction energetics across the tree of life. By mapping redox potentials onto networks of oxidoreductase fold evolution, we can infer the evolution of electron transfer energetics over deep time. ProtReDox is designed to include user-contributed submissions with the intention of making it a valuable resource for researchers in this field.     

The interaction of muscle moment arm,knee laxity,and torque in a multi-scale musculoskeletal model of the lower limb

IntroductionMusculoskeletal modeling allows insight into the interaction of muscle force and knee joint kinematics that cannot be measured in the laboratory. However, musculoskeletal models of the lower extremity commonly use simplified representations of the knee that may limit analyses of the interaction between muscle forces and joint kinematics. The goal of this research was to demonstrate how muscle forces alter knee kinematics and consequently muscle moment arms and joint torque in a musculoskeletal model of the lower limb that includes a deformable representation of the knee.MethodsTwo musculoskeletal models of the lower limb including specimen-specific articular geometries and ligament deformability at the knee were built in a finite element framework and calibrated to match mean isometric torque data collected from 12 healthy subjects. Muscle moment arms were compared between simulations of passive knee flexion and maximum isometric knee extension and flexion. In addition, isometric torque results were compared with predictions using simplified knee models in which the deformability of the knee was removed and the kinematics at the joint were prescribed for all degrees of freedom.ResultsPeak isometric torque estimated with a deformable knee representation occurred between 45° and 60° in extension, and 45° in flexion. The maximum isometric flexion torques generated by the models with deformable ligaments were 14.6% and 17.9% larger than those generated by the models with prescribed kinematics; by contrast, the maximum isometric extension torques generated by the models were similar. The change in hamstrings moment arms during isometric flexion was greater than that of the quadriceps during isometric extension (a mean RMS difference of 9.8 mm compared to 2.9 mm, respectively).DiscussionThe large changes in the moment arms of the hamstrings, when activated in a model with deformable ligaments, resulted in changes to flexion torque. When simulating human motion, the inclusion of a deformable joint in a multi-scale musculoskeletal finite element model of the lower limb may preserve the realistic interaction of muscle force with knee kinematics and torque.     

The influence of increased muscle spindle sensitivity on Achilles tendon jerk and H-reflex in relaxed human subjects

Whether the fusimotor system contributes to reflex gain changes during reinforcement maneuvers is re-examined in the light of new data. Recently, from direct recordings of spindle afferent activity originating from ankle flexor muscles, we showed that mental computation increased the muscle spindle mechanical sensitivity in completely relaxed human subjects without concomitant alpha-motoneuron activation, providing evidence for selective fusimotor drive activation. In the present study, the effects of mental computation were investigated on monosynaptic reflexes elicited in non-contracting soleus muscle either by direct nerve stimulation (Hoffmann reflex, H) or by tendon tap (Tendinous reflex, T). The aim was to relate the time course of the changes in reflex size to the increase in spindle sensitivity during mental task in order to explore whether fusimotor activation can influence the size of the monosynaptic reflex. The results show changes in reflex amplitude that parallel the increase in muscle spindle sensitivity. When T-reflex is consistently facilitated during mental effort, the H-reflex is either depressed or facilitated, depending on the subjects. These findings suggest that the increased activity in muscle spindle primary endings may account for mental computation-induced changes in both tendon jerk and H-reflex. The facilitation of T-reflex is attributed to the enhanced spindle mechanical sensitivity and the inhibition of H-reflex is attributed to post-activation depression following the increased Ia ongoing discharge. This study supports the view that the fusimotor sensitization of muscle spindles is responsible for changes in both the mechanically and electrically elicited reflexes. It is concluded that the fusimotor drive contributed to adjustment of the size of tendon jerk and H-reflex during mental effort. The possibility that a mental computation task may also operate by reducing the level of presynaptic inhibition is discussed on the basis of H-reflex facilitation.     

The effects of alterations in temperature and flow regime on organic carbon dynamics in Mediterranean river networks

It is only recently that freshwaters have been identified as important quantitative components of the carbon (C) cycle at global and regional scales. To date there are no studies that quantitatively predict the effects of alterations in temperature and flow regimes, individually, or in concert, on organic C dynamics in streams. To address this need, we applied a mechanistic model to simulate organic C dynamics in Mediterranean river networks under 27 different scenarios of altered temperature and flow regimes. We predict that the organic C dynamics in freshwaters in the Mediterranean, as well as in other semiarid regions, will be highly sensitive to global climate change owing to major increases in the degree of intermittency as well as in flood frequency and magnitude. Results indicate that flow regime alterations increase C export rates, whereas temperature alterations increase instream metabolism of organic C. However, flow regime alterations exhibit a much greater influence on C dynamics than do changes in the temperature regime. Reservoirs partly counteract the effects of flow extremes on C export rates, and their role in the C dynamics increases with increasing flow variability. The present study is one of the first studies to quantify the complex interactions between the flow and the temperature regime on C dynamics, emphasizing the key role of extreme events such as dry periods and floods, compared with overall trend effects. This information is pivotal in understanding the impact of future climate change on global C dynamics.