How animals move: an integrative view.

Recent advances in integrative studies of locomotion have revealed several general principles. Energy storage and exchange mechanisms discovered in walking and running bipeds apply to multilegged locomotion and even to flying and swimming. Nonpropulsive lateral forces can be sizable, but they may benefit stability, maneuverability, or other criteria that become apparent in natural environments. Locomotor control systems combine rapid mechanical preflexes with multimodal sensory feedback and feedforward commands. Muscles have a surprising variety of functions in locomotion, serving as motors, brakes, springs, and struts. Integrative approaches reveal not only how each component within a locomotor system operates but how they function as a collective whole.

[1]  G. Cavagna,et al.  MECHANICAL WORK IN RUNNING. , 1964, Journal of applied physiology.

[2]  G. C. Joyce,et al.  The mechanical properties of cat soleus muscle during controlled lengthening and shortening movements , 1969, The Journal of physiology.

[3]  P. Rack,et al.  The short range stiffness of active mammalian muscle and its effect on mechanical properties , 1974, The Journal of physiology.

[4]  Douglas G. Stuart,et al.  Neural Control of Locomotion , 1976, Advances in Behavioral Biology.

[5]  G. Cavagna,et al.  The sources of external work in level walking and running. , 1976, The Journal of physiology.

[6]  J. Rayner A New Approach to Animal Flight Mechanics , 1979 .

[7]  Robert K. Colwell,et al.  Elevation and the Morphology, Flight Energetics, and Foraging Ecology of Tropical Hummingbirds , 1979, The American Naturalist.

[8]  N. V. KOKSHAYSKY,et al.  Tracing the wake of a flying bird , 1979, Nature.

[9]  Richard B. Stein,et al.  What muscle variable(s) does the nervous system control in limb movements? , 1982, Behavioral and Brain Sciences.

[10]  T. Daniel Mechanics and energetics of medusan jet propulsion , 1983 .

[11]  C. Ellington The Aerodynamics of Hovering Insect Flight. I. The Quasi-Steady Analysis , 1984 .

[12]  C. Ellington THE AERODYNAMICS OF HOVERING INSECT FLIGHT. V. A VORTEX THEORY , 1984 .

[13]  S. Vogel FLOW-ASSISTED SHELL REOPENING IN SWIMMING SCALLOPS , 1985 .

[14]  R. Josephson Mechanical Power output from Striated Muscle during Cyclic Contraction , 1985 .

[15]  W. Barnes,et al.  THE CUTICULAR STRESS DETECTOR (CSD2) OF THE CRAYFISH II. ACTIVITY DURING WALKING AND INFLUENCES ON LEG COORDINATION , 1986 .

[16]  G. Spedding,et al.  The generation of circulation and lift in a rigid two-dimensional fling , 1986, Journal of Fluid Mechanics.

[17]  Adrian L. R. Thomas,et al.  Vortex flow visualizations reveal change in upstroke function with flight speed in bats , 1986, Nature.

[18]  C. Capaday,et al.  Difference in the amplitude of the human soleus H reflex during walking and running. , 1987, The Journal of physiology.

[19]  S. Childress,et al.  Scanning currents in Stokes flow and the efficient feeding of small organisms , 1987, Journal of Fluid Mechanics.

[20]  R. Blickhan,et al.  Locomotion Energetics of the Ghost Crab: II. Mechanics of the Centre of Mass During Walking and Running , 1987 .

[21]  G. Spedding The Wake of a Kestrel (Falco Tinnunculus) in Flapping Flight , 1987 .

[22]  R. M. Alexander,et al.  Elastic mechanisms in animal movement , 1988 .

[23]  R. Blickhan The spring-mass model for running and hopping. , 1989, Journal of biomechanics.

[24]  R. M. Alexander,et al.  Optimization and gaits in the locomotion of vertebrates. , 1989, Physiological reviews.

[25]  Wolfgang Wieser,et al.  Energy transformations in cells and organisms : proceedings of the 10th Conference of the European Society for Comparative Physiology and Biochemistry, May 1989, Innsbruck, Austria , 1989 .

[26]  M. Koehl,et al.  THE INTERACTION OF BEHAVIORAL AND MORPHOLOGICAL CHANGE IN THE EVOLUTION OF A NOVEL LOCOMOTOR TYPE: “FLYING” FROGS , 1990, Evolution; international journal of organic evolution.

[27]  T. Daniel,et al.  Dynamic Mechanical Properties of Extensor Muscle Cells of the Shrimp Pandalus Danae: Cell Design for Escape Locomotion , 1990 .

[28]  Roland Hengstenberg,et al.  Gaze control in the blowfly Calliphora: a multisensory, two-stage integration process , 1991 .

[29]  R. Full,et al.  Mechanics of a rapid running insect: two-, four- and six-legged locomotion. , 1991, The Journal of experimental biology.

[30]  R. Blickhan,et al.  Leg design in hexapedal runners. , 1991, The Journal of experimental biology.

[31]  C T Farley,et al.  A mechanical trigger for the trot-gallop transition in horses. , 1991, Science.

[32]  A. Brodsky Vortex Formation in the Tethered Flight of the Peacock Butterfly Inachis io L. (Lepidoptera, Nymphalidae) and some Aspects of Insect Flight Evolution , 1991 .

[33]  R. Griffiths Shortening of muscle fibres during stretch of the active cat medial gastrocnemius muscle: the role of tendon compliance. , 1991, The Journal of physiology.

[34]  V. Edgerton,et al.  Short communication. Sarcomere length changes during fish swimming. , 1992, The Journal of experimental biology.

[35]  K. Dial Activity patterns of the wing muscles of the pigeon (Columba livia) during different modes of flight , 1992 .

[36]  C. I. Smith,et al.  MYOTOMAL MUSCLE FUNCTION AT DIFFERENT LOCATIONS IN THE BODY OF A SWIMMING FISH , 1993 .

[37]  Randall D. Beer,et al.  Biological Neural Networks in Invertebrate Neuroethology and Robotics. Editors: Randall D. Beer, Roy E. Ritzmann, Thomas McKenna (Academic Press, Inc., Harcourt Brace Jovanovich, 1993) , 1996, SGAR.

[38]  A. Biewener,et al.  PECTORALIS MUSCLE FORCE AND POWER OUTPUT DURING DIFFERENT MODES OF FLIGHT IN PIGEONS (COLUMBA LIVIA) , 1993 .

[39]  C. T. Farley,et al.  Running springs: speed and animal size. , 1993, The Journal of experimental biology.

[40]  S. Gueron,et al.  Simulations of three-dimensional ciliary beats and cilia interactions. , 1993, Biophysical journal.

[41]  R. Marsh,et al.  Power output of scallop adductor muscle during contractions replicating the in vivo mechanical cycle. , 1994, The Journal of experimental biology.

[42]  R. Blickhan,et al.  Dynamic and static stability in hexapedal runners. , 1994, The Journal of experimental biology.

[43]  M. S. Tu,et al.  MODULATION OF NEGATIVE WORK OUTPUT FROM A STEERING MUSCLE OF THE BLOWFLY CALLIPHORA VICINA , 1994, The Journal of experimental biology.

[44]  P. Webb,et al.  Hydrostatic stability of fish with swim bladders: not all fish are unstable , 1994 .

[45]  L. Rome,et al.  Built for jumping: the design of the frog muscular system. , 1994, Science.

[46]  Full,et al.  Maximum single leg force production: cockroaches righting on photoelastic gelatin , 1995, The Journal of experimental biology.

[47]  K. Pearson Proprioceptive regulation of locomotion , 1995, Current Opinion in Neurobiology.

[48]  J. Costello,et al.  Flow and feeding by swimming scyphomedusae , 1995 .

[49]  C. Gilbert,et al.  Oculomotor control in calliphorid flies: Head movements during activation and inhibition of neck motor neurons corroborate neuroanatomical predictions , 1995, The Journal of comparative neurology.

[50]  R. Dudley,et al.  Limits to vertebrate locomotor energetics suggested by hummingbirds hovering in heliox , 1995, Nature.

[51]  F. Maytag Evolution , 1996, Arch. Mus. Informatics.

[52]  M. Martinez ISSUES FOR AQUATIC PEDESTRIAN LOCOMOTION , 1996 .

[53]  E. Marder,et al.  Principles of rhythmic motor pattern generation. , 1996, Physiological reviews.

[54]  Liu,et al.  A computational fluid dynamics study of tadpole swimming , 1996, The Journal of experimental biology.

[55]  J. Y. Cheng,et al.  Jet-propelled swimming in scallops: swimming mechanics and ontogenic scaling , 1996 .

[56]  L. Fauci A COMPUTATIONAL MODEL OF THE FLUID DYNAMICS OF UNDULATORY AND FLAGELLAR SWIMMING , 1996 .

[57]  Adrian L. R. Thomas,et al.  Leading-edge vortices in insect flight , 1996, Nature.

[58]  T J Roberts,et al.  Muscular Force in Running Turkeys: The Economy of Minimizing Work , 1997, Science.

[59]  M. Dickinson,et al.  Phosphorylation-dependent power output of transgenic flies: an integrated study. , 1997, Biophysical journal.

[60]  C. T. Farley,et al.  Mechanics of locomotion in lizards. , 1997, The Journal of experimental biology.

[61]  Randall D. Beer,et al.  The brain has a body: adaptive behavior emerges from interactions of nervous system, body and environment , 1997, Trends in Neurosciences.

[62]  Adrian L. R. Thomas,et al.  FLOW VISUALIZATION AND UNSTEADY AERODYNAMICS IN THE FLIGHT OF THE HAWKMOTH, MANDUCA SEXTA , 1997 .

[63]  M. S. Tu,et al.  The Function of Dipteran Flight Muscle , 1997 .

[64]  J. Marden,et al.  From Molecules to Mating Success: Integrative Biology of Muscle Maturation in a Dragonfly , 1998 .

[65]  A. Biewener,et al.  In vivo pectoralis muscle force-length behavior during level flight in pigeons (Columba livia) , 1998, The Journal of experimental biology.

[66]  W P Chan,et al.  Visual input to the efferent control system of a fly's "gyroscope". , 1998, Science.

[67]  Daniel P. Ferris,et al.  Running in the real world: adjusting leg stiffness for different surfaces , 1998, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[68]  Daniel P. Ferris,et al.  10 Biomechanics of Walking and Running: Center of Mass Movements to Muscle Action , 1998, Exercise and sport sciences reviews.

[69]  P. Willems,et al.  Mechanics and energetics of human locomotion on sand. , 1998, The Journal of experimental biology.

[70]  L. Rome Some advances in integrative muscle physiology. , 1998, Comparative biochemistry and physiology. Part B, Biochemistry & molecular biology.

[71]  Full,et al.  Underwater punting by an intertidal crab: a novel gait revealed by the kinematics of pedestrian locomotion in air versus water , 1998, The Journal of experimental biology.

[72]  Full,et al.  Energy absorption during running by leg muscles in a cockroach , 1998, The Journal of experimental biology.

[73]  K. Pearson,et al.  Enhancement and Resetting of Locomotor Activity by Muscle Afferentsa , 1998, Annals of the New York Academy of Sciences.

[74]  A. Biewener,et al.  In vivo muscle force-length behavior during steady-speed hopping in tammar wallabies. , 1998, The Journal of experimental biology.

[75]  R. Shadwick,et al.  Muscle Dynamics in Fish During Steady Swimming , 1998 .

[76]  Tobalske,et al.  Kinematics of flap-bounding flight in the zebra finch over a wide range of speeds , 1999, The Journal of experimental biology.

[77]  J. Bradbury,et al.  Directional acoustic radiation in the strut display of male sage grouse Centrocercus urophasianus. , 1999, The Journal of experimental biology.

[78]  Full,et al.  Many-legged maneuverability: dynamics of turning in hexapods , 1999, The Journal of experimental biology.

[79]  T. Nichols,et al.  The role of musculoskeletal mechanics in motor coordination. , 1999, Progress in brain research.

[80]  R J Full,et al.  Templates and anchors: neuromechanical hypotheses of legged locomotion on land. , 1999, The Journal of experimental biology.

[81]  R. Josephson Dissecting muscle power output. , 1999, The Journal of experimental biology.

[82]  D. Ellerby,et al.  Fish swimming: patterns in muscle function. , 1999, The Journal of experimental biology.

[83]  E. Zehr,et al.  What functions do reflexes serve during human locomotion? , 1999, Progress in Neurobiology.

[84]  R. Blickhan,et al.  Stabilizing function of skeletal muscles: an analytical investigation. , 1999, Journal of theoretical biology.

[85]  A. Biewener,et al.  Dynamics of muscle function during locomotion: accommodating variable conditions. , 1999, The Journal of experimental biology.

[86]  J. Videler,et al.  Aquatic vertebrate locomotion: wakes from body waves. , 1999, The Journal of experimental biology.

[87]  J. Vigoreaux,et al.  An Integrated View of Insect Flight Muscle: Genes, Motor Molecules, and Motion. , 1999, News in physiological sciences : an international journal of physiology produced jointly by the International Union of Physiological Sciences and the American Physiological Society.

[88]  G. E. Loeb,et al.  A hierarchical foundation for models of sensorimotor control , 1999, Experimental Brain Research.

[89]  T. Sinkjaer,et al.  The stretch reflex and H-reflex of the human soleus muscle during walking. , 1999, Motor control.

[90]  Lauder,et al.  Locomotor forces on a swimming fish: three-dimensional vortex wake dynamics quantified using digital particle image velocimetry. , 1999, The Journal of experimental biology.

[91]  R. Full,et al.  The role of the mechanical system in control: a hypothesis of self-stabilization in hexapedal runners , 1999 .

[92]  R. Stein,et al.  Modulation of stretch reflexes during imposed walking movements of the human ankle. , 1999, Journal of neurophysiology.

[93]  M. Dickinson,et al.  Wing rotation and the aerodynamic basis of insect flight. , 1999, Science.

[94]  Lauder,et al.  Locomotion in sturgeon: function of the pectoral fins. , 1999, The Journal of experimental biology.

[95]  M. Binder,et al.  Summation of effective synaptic currents and firing rate modulation in cat spinal motoneurons. , 2000, Journal of neurophysiology.

[96]  J. Duysens,et al.  Load-regulating mechanisms in gait and posture: comparative aspects. , 2000, Physiological reviews.

[97]  Markus Raffel,et al.  Particle Image Velocimetry: A Practical Guide , 2002 .