In vivo strain in the humerus of pigeons (Columba livia) during flight

Longitudinal and principal strain recordings were made in vivo at three sites (dorsal, anterior, and ventral) on the humeral midshaft of pigeons executing five modes of free flight: Take‐off, level flight, landing, vertical ascent, and near‐vertical descent. Strains were also recorded while the birds flew carrying weights that were 33%, 50%, or 100% of their body weight. The relative distribution of strain measured at the three surface midshaft sites and across the bone's cortex was found to be similar for all flight modes. Principal strains recorded in the dorsal and ventral humerus indicated considerable torsion produced by aerodynamic loading of the wing surface posterior to the bone. Measured torsional shear strains (maximum: 2,700–4,150 μ ε during level flight) were 1.5 times greater than longitudinal strains. In addition to torsion, the humerus is also subjected to significant dorsoventral bending owing to lift forces acting on the wing during the downstroke. Analysis of the cross‐sectional distribution of longitudinal strains at the humeral midshaft cortex shows that the orientation of bending shifts in a regular manner during the downstroke, indicating that the wing generates progressively more thurst (vs. lift) later in the downstroke. This shift is less during take‐off and vertical ascent when greater lift is required. Peak principal and longitudinal strains increased by an average of only 50% from landing to vertical ascending flight and take‐off (e.g., dorsal humerus: −1,503 to −2,329 μ ε) and did not exceed −2,600 μ epsiv; at any site, even when the birds flew carrying twice their body weight. Strains recorded when birds flew at two times their body weight (100% BW load) were similar in magnitude to those recorded during vertical ascent and take‐off and likely represent those developed during maximal performance. Strains developed within the midshaft were maximal in the anterodorsal and posteroventral cortices, not at the dorsal, ventral, and anterior sites at which strain was recorded. Consequently, maximum strains experienced by the bone are probably 20–25% greater than those recorded (ca. 3,200 μ ε), indicating a safety factor of about 3.5 for compressive strain failure. The much higher shear strains, however, indicate a lower safety factor (1.9), in which the bone's torsional strength is its most critical design feature. Finally, the magnitude and distribution of strains developed in the humerus of pigeons are generally similar to those recorded in the humerus of large fruit‐eating bats during flight. © 1995 Wiley‐Liss, Inc.

[1]  G. E. Goslow,et al.  A Cineradiographic Analysis of Bird Flight: The Wishbone in Starlings Is a Spring , 1988, Science.

[2]  L E Lanyon,et al.  Dynamic strain similarity in vertebrates; an alternative to allometric limb bone scaling. , 1984, Journal of theoretical biology.

[3]  L E Lanyon,et al.  The influence of mechanical function on the development and remodeling of the tibia. An experimental study in sheep. , 1979, The Journal of bone and joint surgery. American volume.

[4]  David R. Carrier,et al.  Wing bone stresses in free flying bats and the evolution of skeletal design for flight , 1992, Nature.

[5]  Colin J Pennycuick,et al.  Bird flight performance: a practical calculation manual , 1992 .

[6]  U. Norberg,et al.  Evolutionary convergence in foraging niche and flight morphology in insectivorous aerial-hawking birds and bats , 1986 .

[7]  Knut Schmidt-Nielsen,et al.  Power Input During Flight of the Fish Crow, Corvus Ossifragus , 1973 .

[8]  Jeremy M. V. Rayner,et al.  Bounding and undulating flight in birds , 1985 .

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

[10]  V. Tucker Respiratory Exchange and Evaporative Water Loss in the Flying Budgerigar , 1968 .

[11]  Andrew A. Biewener,et al.  Mechanics of locomotion and jumping in the horse (Equus): in vivo stress in the tibia and metatarsus , 1988 .

[12]  V. Tucker Metabolism during flight in the laughing gull, Larus atricilla. , 1972, The American journal of physiology.

[13]  L. Lanyon,et al.  Limb mechanics as a function of speed and gait: a study of functional strains in the radius and tibia of horse and dog. , 1982, The Journal of experimental biology.

[14]  C. Pennycuick Power requirements for horizontal flight in the pigeon Columba livia , 1968 .

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

[16]  John D. Currey,et al.  The Mechanical Adaptations of Bones , 1984 .

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

[18]  R. H. Brown,et al.  THE FLIGHT OF BIRDS , 1963 .

[19]  C. Pennycuick,et al.  STRUCTURAL LIMITATIONS ON THE POWER OUTPUT OF THE PIGEON'S FLIGHT MUSCLES , 1966 .

[20]  Jmv Rayner,et al.  Momentum and energy in the wake of a pigeon (Columba livia) in slow flight , 1984 .

[21]  R. Blickhan,et al.  Preferred speeds in terrestrial vertebrates: are they equivalent? , 1988, The Journal of experimental biology.

[22]  C. Pennycuick,et al.  The strength of the pigeon's wing bones in relation to their function. , 1967, The Journal of experimental biology.

[23]  D R Carter,et al.  Anisotropic analysis of strain rosette information from cortical bone. , 1978, Journal of biomechanics.

[24]  J. Bertram,et al.  Telemetered in vivo strain analysis of locomotor mechanics of brachiating gibbons , 1989, Nature.

[25]  R. H. Brown The flight of birds; the flapping cycle of the pigeon. , 1948, The Journal of experimental biology.

[26]  J. Larochelle,et al.  The metabolic cost of flight in unrestrained birds. , 1978, The Journal of experimental biology.

[27]  W. F. Riley,et al.  Experimental stress analysis , 1978 .

[28]  M. Labarbera,et al.  Contribution of internal bony trabeculae to the mechanical properties of the humerus of the pigeon (Columba livia) , 1993 .

[29]  C. R. Taylor,et al.  Bone strain: a determinant of gait and speed? , 1986, The Journal of experimental biology.

[30]  A A Biewener,et al.  Musculoskeletal design in relation to body size. , 1991, Journal of biomechanics.

[31]  A. Biewener Biomechanics of mammalian terrestrial locomotion. , 1990, Science.