Background: Associate Professor in Muscle Function and Movement (Leeds, 2011-), Lecturer (Leeds 2001-2011). I studied Biology at the University of Leeds and remained there to carry out a PhD on the effects of training and fatigue on the mechanical properties of skeletal muscle. I carried out postdoctoral research at Northeastern University in Boston and the University of Cambridge. I returned to the University of Leeds as a lecturer in 2011.
Contact: Garstang 5.61a, +44(0) 113 34 32897,
The physiology and biomechanics of animal movement
Overview. My research focuses on the physiology and biomechanics of animal movement. The overall goal of the lab is to understand how muscle mechanical function and energetics determine the overall metabolic energy expenditure and physical performance of locomotion. We use an integrative and comparative approach to reveal the underlying cellular, physiological and biomechanical determinants of physical performance in order to understand the evolution of musculoskeletal design.
Mechanical function of muscles during locomotion. A large part of our work has investigated muscle function during flight because during flight the bulk of the active muscles have a simple mechanical function – to generate power. We have studied the explosive take-off flights performed by some birds to investigate the physiological adaptations that allow muscles to generate maximal short-term power output. One of the species that we studied – the blue-breasted quail – develops the highest power of any skeletal muscle yet measured (per gram of muscle; Askew and Marsh, 2001). Bird flight also proved to be a useful system in which to investigate power modulation strategies. The need to modulate power is relevant to many activities including acceleration, incline locomotion, and speed related variation in power requirements. We have been able to determine the importance of power modulation through changes in muscle recruitment and length trajectory as well as at the level of the organism by the use of intermittent flight behaviour (Askew and Ellerby, 2007; Morris and Askew, 2010).
In vivo zebra finch pectoralis fascicle strain and EMG activity at 10 m s-1. Fascicle length was measured by sonomicrometry. Muscle activity was measured using a bipolar EMG electrode.
Energetics of Locomotion. My laboratory also investigates metabolic energy use during locomotion with the primary aim of quantifying energy use of the locomotory muscles in order to gain a better understanding of the determinants of whole organismal energy use. We use whole organism metabolic rate is determined by measuring O2 consumption and CO2 production during locomotion (Fig. 2; Morris and Askew, 2010). In leaf-cutter ants this technique has been used to investigate the behavioural adaptations that occur during the negotiation of hilly terrain (Holt and Askew, 2012). We also investigated the effects that wearing armour would have had on the locomotion of Medieval knights (Askew et al., 2011), finding that when wearing the armour the energy expenditure was doubled when they walked or ran. This is much more than wearing a backpack of equivalent mass due to the increased cost of swinging the loaded limbs and impaired breathing when wearing armour. These findings can help historians interpret the feats of battle. At the level of the muscle, we quantify energy use by measuring the total enthalpy output (the sum of the mechanical work and heat production) and also, indirectly, by measuring regional blood flow. Our research has demonstrated that a muscle doing stretch-shorten work uses no more energy than when force is produced isometrically, challenging the view that elastic tendons reduce locomotor costs by replacing muscle work (Holt et al., 2014). We have also developed approaches that allow energy use can also be quantified at the level of the cross-bridges (Askew et al., 2010).
Using respirometry to measure metabolic energy expenditure during locomotion - while walking in Medieval armour and during flight in a cockatiel
Integrating the Mechanics and Energetics of Locomotion. During all modes of locomotion, muscles convert chemical energy into mechanical work that is ultimately transferred to the environment to produce movement. To achieve a full understanding of the system, we need to be able to trace the transfer of energy between all levels of organisation from the contractile proteins to the momentum transferred to the animal's wake and relate this to the animal's locomotor performance, morphology and ecology. In collaboration with Dr Richard Bomphrey (Royal Veterinary College) we are using an integrative, multidisciplinary approach to determine, in insect flight, the transfer of energy from biochemical potential energy, through the muscles, to the surrounding air.
Funding. Research in my laboratory is funded by the BBSRC, EPSRC, and NERC.
Latest: PhD Studentship available on "Integrating the mechanics and energetics of running" in collaboration with Professor Stuart Egginton. See findaphd.com. Application deadline 19th January 2015.
Running energetics – in this paper we tested the idea that tendons save energy by reducing cyclic muscle work by comparing the cost of force generation during constant length muscle actions with active stretch-shorten cycles.
Holt, N.C, Roberts, T.J. and Askew, G.N. The energetic benefits of tendon springs in running: is the reduction of muscle work important? J. Exp. Biol. 217, 4365-4371. 2014. doi: 10.1242/ jeb.112813
Peacock’s train is not such a drag – here I measured the effect that the elaborate plumage of male peafowl has on their take-off performance.
Askew, G.N. The elaborate plumage in peacocks is not such a drag. J. Exp. Biol. 217, 3237-3241. 2014.
Paper highlighted in Inside JEB. J. Exp. Biol. 217, 3189; doi: 10.1242/ jeb.112342 and online (ScienceNow, Huffington Post, Times of India, Nature World News)
Photo by G. Askew
Energetics of locomotion in Medieval armour – here we measured how much effort is required to walk and run while wearing armour.
Askew, G.N., Formenti, F., Minetti, A.E. Limitations imposed by wearing armour on Medieval soldiers’ locomotor performance. Proc. R. Soc. Lond. B 279, 640-644. 2012.
Ant energetics – in this paper we measured the energetic cost of walking uphill and downhill in leaf cutter ants in order to understand the behavior of route selection.
Holt, N. and Askew, G.N. Locomotion on a slope: metabolic energy use, behavioural adaptations and the implications for route selection. J. Exp. Biol. 215, 2545-2550. 2012.
Photo by G. Askew
Power modulation in bird flight –this is a series of three papers on cockatiel flight in which we quantify muscle mechanical performance during flight using two methods and compared this to the metabolic cost of flight.
Morris, C.R. and Askew, G.N. Power modulation strategies and the mechanical power requirements of flight in the cockatiel (Nymphicus hollandicus). J. Exp. Biol., 213, 2770-2780. 2010.
Morris, C.R., Nelson, F.E. and Askew, G.N. The metabolic power requirements of flight and estimations of flight muscle efficiency in the cockatiel (Nymphicus hollandicus). J. Exp. Biol., 213, 2788-2796. 2010.
Morris, C.R. and Askew, G.N. Comparison between mechanical power requirements of flight estimated using an aerodynamic model and in vitro muscle performance in the cockatiel (Nymphicus hollandicus). J. Exp. Biol. 213, 2781-2787. 2010.
Papers highlighted in Inside JEB. Journal of Experimental Biology 213, I (2010). doi:10.1242/jeb.04911114
BLGY2330 - Terrestrial Ecology and Behaviour Field Course
BMSC1110/SPSC1220 - Foundation modules
SPSC2302 - Exercise Physiology in Sport, Health and Disease
SPSC2304 - Mechanics of Sport and Exercise 2
SPSC2304/2213 - Mechanics of Sport and Exercise
SPSC3061 - Research Project in Sport and Exercise Science II
Member of Graduate School Committee
Centre membership: The Earth and Biosphere Institute
Group Leader Dr Graham Askew (Associate Professsor in Muscle Function & Movement)
The physiology and biomechanics of animal movement
Dr Peter Tickle (Research Fellow)
Morris CR, Askew GN The mechanical power output of the pectoralis muscle of cockatiel (Nymphicus hollandicus): the in vivo muscle length trajectory and activity patterns and their implications for power modulation J EXP BIOL 213 2770-2780, 2010
Morris CR, Nelson FE, Askew GN The metabolic power requirements of flight and estimations of flight muscle efficiency in the cockatiel (Nymphicus hollandicus) J EXP BIOL 213 2788-2796, 2010
Morris CR, Askew GN Comparison between mechanical power requirements of flight estimated using an aerodynamic model and in vitro muscle performance in the cockatiel (Nymphicus hollandicus) J EXP BIOL 213 2781-2787, 2010
Askew GN, Tregear RT, Ellington CP The scaling of myofibrillar actomyosin ATPase activity in apid bee flight muscle in relation to hovering flight energetics J EXP BIOL 213 1195-1206, 2010
Miller G, Neilan M, Chia R, Gheryani N, Holt N, Charbit A, Wells S, Tucci V, Lalanne Z, Denny P, Fisher EMC, Cheeseman M, Askew GN, Dear TN ENU Mutagenesis Reveals a Novel Phenotype of Reduced Limb Strength in Mice Lacking Fibrillin 2 PLOS ONE 5 -, 2010
Holt NC, Roberts TJ, Askew GN The energetic benefits of tendon springs in running: Is the reduction of muscle work important? Journal of Experimental Biology 217 4365-4371, 2014
Holt NC, Askew GN Locomotion on a slope in leaf-cutter ants: metabolic energy use, behavioural adaptations and the implications for route selection on hilly terrain. J Exp Biol 215 2545-2550, 2012
Ellerby DJ, Askew GN Modulation of flight muscle power output in budgerigars Melopsittacus undulatus and zebra finches Taeniopygia guttata: in vitro muscle performance J EXP BIOL 210 3780-3788, 2007
Ellerby DJ, Askew GN Modulation of pectoralis muscle function in budgerigars Melopsitaccus undulatus and zebra finches Taeniopygia guttata in response to changing flight speed J EXP BIOL 210 3789-3797, 2007
Askew GN, Ellerby DJ The mechanical power requirements of avian flight BIOLOGY LETT 3 445-448, 2007
West TG, Donohoe PH, Staples JF, Askew GN Tribute to R.G. Boutilier: The role for skeletal muscle in the hypoxia-induced hypometabolic responses of submerged frogs J EXP BIOL 209 1159-1168, 2006
James RS, Wilson RS, Askew GN Effects of caffeine on mouse skeletal muscle power output during recovery from fatigue J APPL PHYSIOL 96 545-552, 2004
Askew GN, Cox VM, Altringham JD, Goldspink DF Mechanical properties of the latissimus dorsi muscle after cyclic training Journal of Applied Physiology 93 649-659, 2002
Askew GN, Marsh RL Muscle designed for maximum short-term power output: quail flight muscle. J Exp Biol 205 2153-2160, 2002
Askew GN, Marsh RL, Ellington CP The mechanical power output of the flight muscles of blue-breasted quail (Coturnix chinensis) during take-off. J Exp Biol 204 3601-3619, 2001
Askew GN, Marsh RL The mechanical power output of the pectoralis muscle of blue-breasted quail (Coturnix chinensis): the in vivo length cycle and its implications for muscle performance. J Exp Biol 204 3587-3600, 2001
Askew GN, Marsh RL Optimal relative shortening velocity (V/Vmax) of skeletal muscle during cyclical contractions: length-force effects and velocity dependent activation and deactivation Journal of Experimental Biology 201 1527-1540, 1998
Askew GN, Young IS, Altringham JD Fatigue of mouse soleus muscle using the work loop technique Journal of Experimental Biology 200 2907-2912, 1997
Askew GN, Marsh RL The effects of length trajectory on the mechanical power output of mouse skeletal muscles Journal of Experimental Biology 200 3119-3131, 1997