Comprehensive Physiology Wiley Online Library

Invertebrate Locomotor Systems

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Mechanisms of Locomotion
1.1 Swimming
1.2 Crawling
1.3 Walking, Running, and Rolling
1.4 Jumping
1.5 Flying
1.6 Comparison of Locomotor Dynamics
2 Production of Locomotion: Musculoskeletal Systems
2.1 Filament, Sarcomere, and Muscle Level
3 Energetics of Locomotion
3.1 Aerobic Metabolism
3.2 Anaerobic Metabolism
3.3 Endurance and Metabolism
3.4 Metabolic Cost of Transport
4 Conclusions
4.1 Trends
5 Future Research
5.1 Comparative Muscle Physiology
5.2 Comparative Bioenergetics and Exercise Physiology
5.3 Comparative Biomechanics
5.4 Collaboration
5.5 Direct‐Experiments Using Innovative Technology
5.6 Cybercreatures and Experiments: Computer Modeling
6 Appendix: List of Symbols
Figure 1. Figure 1.

Schematic diagram showing integration of comparative bioenergetics, exercise physiology, functional morphology, muscle physiology, and comparative biomechanics. A: Energy available from several sources is transduced to segments (appendages or body sections) through muscles, depending on geometry. The chapter is organized in sections from right to left. B: Magnification of musculoskeletal role in transducing energy. Musculoskeletal parameters determine musculoskeletal forces. Musculoskeletal forces and joint geometry determine moment. Moment gives rise to acceleration, velocity, and angle change. Velocity and angle change (change in muscle length) feedback to affect musculoskeletal force production. Moment and joint angular velocity determine amount of energy transferred. Direction of movement and moment determine whether energy enters, leaves, or is transferred from a segment. Segmental energy and morphology of all segments determine power output of whole body and represent locomotor mechanical energy in A.

Figure 2. Figure 2.

Legs of animals in general, and of invertebrates in particular, can provide biological inspiration for the design of new robot legs. A: Legs operating in a more upright posture in a vertical plane parallel to the body (horizontal first joint axis for body–leg attachment), as seen in many birds and mammals, are gravitationally loaded and muscles must bear part of the body's weight 24. Animals using a horizontal first axis, however, can take advantage of gravity in swinging their legs. B–D: Legs operating in a sprawled posture (vertical first joint axis for body–leg attachment, like some lizards, amphibians, and arthropods) can potentially decouple gravitational loading of muscles from moving forward. To move forward, vertical first axis legs must project out to the side, resulting in increased static stability. Robots inspired from the design of cockroaches 62,63, whose legs operate in a horizontal plane, demonstrate a linear step, large step lengths, and proficiency at climbing.

adapted from ref. 62
Figure 3. Figure 3.

Speed as a function of body mass. A: Speeds of fliers, jumpers, runners, jetters, swimmers, and crawlers. Shown are maximum speeds available from the literature. Some speeds are reported as truly maximum. The majority are near maximum aerobic speed (speed at which maximal oxygen consumption is attained). Many are preferred speeds. For jumpers, take‐off speeds shown. The shaded area bounds the speed of crawlers. Data include runners 22,41,69,83,134,136,181,199,201,202,207,208,212,259,261,263,264,265,266,278,290,343,344,345,346,347,383,392,413,458,523, crawlers 97,145,277,292,353, jumpers 11,48,53,179,180,282,352, swimmers 7,65,114,119,184,241,276,278,280,290,293,328,369,386,409,466,468,482,511, jetters 106,127,128,130,132,294,338,395,398,399,473,474,475, and fliers 45,152,155,156,173,273,367. (Ref. 158 published after submission of figures.) B: Regression lines of major taxonomic groups (Table 1). Data to generate regressions include insect runners 41,136,181,207,208,212,259,263,266,343,344,345,346,347,383,392,458; crustacean runners 69,83,199,202,261,264,265,278,290,523; crustacean swimmers 7,119,184,241,276,280,293,328,369,386,409,466,468,482; crustacean underwater runners 114,276,278,280,290; bee, fly, and mosquito fliers 156,173,273; and lepidopteran fliers 152,155. Note the change in speed axis from A.

Figure 4. Figure 4.

Cycle frequency as a function of body mass. A: Cycle frequency of fliers, runners, jetters, and swimmers. Frequencies shown are the maximum available from the literature. Some frequencies are reported as truly maximum. The majority are preferred speeds. Data include runners 22,69,134,207,208,212,261; swimmers 7,65,114,290,409; jetters 29,106,127,140,338,473,495; and fliers such as mosquitoes 84,273,366,507, aphids, white flies 84,524, dragonflies 84,366,367, bees, wasps 84,96,156,171,306,366,507, beetles 84,507, flies 84,273,318,366,507,524, flower flies 84,318,507, butterflies 84,152,507, saturnid moths 39,84,96,507, sphinx moths 94,96,366,507, and crane flies 84,171,366,507. B: Regression lines of major taxanomic groups (Table 2). Note the change in cycle frequency axis from A.

Figure 5. Figure 5.

Whole‐animal mass‐specific mechanical power output as a function of body mass. Shaded areas and connected points represent the range of possible values for fliers, assuming 0 (upper bound) and 100% (lower bound) elastic storage. Speeds selected are the maximum available from the literature. Some speeds are reported as truly maximum. Many are preferred speeds. Data include insects 207,208 and crabs 69 for terrestrial locomotion; crustaceans 385,467, insects (65, 3890), jellyfish 140, and molluscs 127,397,398 for aquatic locomotion; and insects 94,102,153,173,299,504 for aerial locomotion.

Figure 6. Figure 6.

Sarcomere morphometrics. Muscles were divided into fast and slow groups based on a variety of criteria, which included fiber type, enzyme level or activity, and contraction kinetics. In general, if a study designated a muscle as fast or slow, this characterization was used. All insect flight muscles were considered to be fast. All crawlers—larvae, worms, and nematodes—have slow muscles. Walking and swimming organisms have both fiber types. Bars represent ± 1 S.D. Numbers in parentheses are sample size. Lmyo, myosin filament length; Lcsarc, length of a sarcomere. A: Sarcomere length. Data include insects such as locusts 48,51,285,504, cockroaches 163,185,287,296,382,415,454,479, water bugs 5,25,163, water beetles 162, katydids 5,164, lepidopterans 89,422, dragonflies 444, larvae 86,88,243,401,422, nymphs 87, flies 405,434, and hemipterans 125; crustaceans such as lobsters 287,295,296,415, crayfish 1,415,461,528, crabs 188,283,476, and merostomatans such as horseshoe crabs 490; arachnids such as spiders 321,436 and scorpions 26,425; molluscs such as scallops 379 and cuttlefish 286; annelids such as worms 376,462 and polychaetes 365; chaetognaths such as arrow worms 161; and coelenterates such as jellyfish 107,439. B: Myosin filament length. Data include insects such as locusts 118,285, cockroaches 5,163,239,240,287,296,415,479, water bugs 5,25,163, water beetles 162, katydids 5,164, lepidopterians 5,513, dragon‐flies 444, larvae 86,88,243,401, and nymphs 87,90; crustaceans such as lobsters 295,296,297,415, crayfish 415,528, and crabs 188,283,476; arachnids such as spiders 321,436; molluscs such as scallops 379 and cuttlefish 286; annelids such as worms 376,462, and polychaetes 365; nematodes 426; and coelenterates such as jellyfish 107,439. C; Ratio of thin/thick filament number. Data include insects such as locusts 118,285, cockroaches 5,239,240,287,296,415,454,479, water bugs 5,25, katydids 5,164, lepidopterians 89,422,513, dragonflies 444, larvae 86,88,243,401,422, nymphs 87,90, and flies 405; crustaceans such as lobsters 287,296,415 and crabs 188,283; arachnids such as spiders 436 and scorpions 26,425; molluscs such as scallops 379 and cuttlefish 286; annelids 376,462; nematodes 426; and coelenterates such as jellyfish 107. D: Sarcomere diagram shows designated lengths.

Figure 7. Figure 7.

Active length‐tension or strain‐stress curves of invertebrate muscles. Stress is normalized to peak isometric tension. Strain is normalized as a fraction of the length that gives peak isometric stress. Shaded areas represent strains that correspond to stresses above 50% maximal isometric stress. Data include bee flight 242, locust flight 353, crayfish 528, frog 271, leech 378, and fly larvae 243 muscles.

Figure 8. Figure 8.

Mechanical power output of invertebrate muscle and its determinants as a function of cycle frequency. Data are from isolated muscle studies using the work‐loop method (Fig. 9) and come from unpublished data of Stevenson and Casey. A: Mass‐specific mechanical power output and work per cycle. Gilmour and Ellington 223 have demonstrated values as high as 110 W/kg from asynchronous, glycerinated muscle fibers of bumblebees. B: Muscle stress, strain, and strain rate. Data include insects 312,349,380,453, crustaceans 455, and molluscs 362.

Figure 9. Figure 9.

Work‐loop method to determine power output of an isolated muscle. A: Muscle is subjected to cyclic length changes (strain) while force per area (stress) is measured. The stimulation pattern is controlled. B: Area under the stress‐strain curve represents energy produced (output) during shortening or absorbed (input) during lengthening. The difference between the curves represents the net work per cycle. Counterclockwise loops result in energy production, whereas clockwise loops result in energy absorption by the muscle. C: Variation in work‐loop shape. Rectangular shape observed at low frequencies (<30 Hz). Triangular shape observed at intermediate frequencies (30–60 Hz). Ellipsoid shape observed at high frequencies (60–180 Hz).

after ref. 311
Figure 10. Figure 10.

Three‐dimensional musculoskeletal model of themeta‐thoracic leg of the cockroach Blaberus discoidalis. Left: Shaded polygons represent the exoskeleton, which was reconstructed from serial sections. Right: Heavy lines represent the lines of action of Hill‐type muscles. The model is articulated at the coxa/trochanter–femur and femurtibia joints so that muscle lengths, moment arms, forces, and joint moments can be estimated for a range of body positions 195. The computer model was created using SIMM.

MusculoGraphics, Evanston, IL
Figure 11. Figure 11.

Whole‐animal mass‐specific metabolic power input as a function of body mass. Oxygen consumption rates are the highest available for the species. In some cases, they are known to be maximal rates. A: Oxygen consumption as a function of body mass for runners, crawlers, swimmers, jetters, and fliers. Shaded areas represent jetters; top right area includes molluscs such as squid; bottom area includes jellyfish. Insect runners can warm their bodies. The regression line shown is for the Q10 corrected data set not shown (Q10 = 2). Data include runners (22,41,42,43,44,190,199,200,201,202,210,212,258,259,261,262,263,264,265,266,301,343,345,346,347,368,374,427,433,514,522; W. J. Van Aardt, unpublished data), runners with elevated body temperature 37,209, crawlers 97,277,292, swimmers 119,184,276,278,279,293,328,369,371,428,466, jetters 29,130,338,395,399,474,495,511, and fliers 36,39,42,91,92,93,96,100,102,104,108,109,110,173,250,251,273,318,333,384,393,427,447,502,507,521. B: Oxygen consumption rates by taxonomic groups (Table 3). Data include insect runners 41,42,43,44,210,212,259,263,266,301,343,345,346,347,368,427; crustacean runners (190,199,200,202,261,262,264,265,514,522; W. J. Van Aardt, unpublished data); swimmers 119,184,276,278,279,293,328,369,371,428,466,474; moth fliers 39,91,92,93,96,100; and bee, fly, and mosquito fliers 36,42,96,102,104,108,109,110,173,250,251,273,318,333,384,393,427,447,502,507,521.

Figure 12. Figure 12.

Aerobic factorial scopes for invertebrate activity. Oxygen consumption rates are the highest available for the species. In some cases, they are known to be maximal rates. Bars represent ± 1 S.D. Numbers in parentheses are sample size. Data include aquatic groups such as coelenterates 338, gastropods 277,292, crustaceans 119,184,276,278,279,293,328,369,371,428,466, and cephalopods 29,395,399,495,511; terrestrial groups such as gastropods 277,292, myriapods 201, crustaceans (190,199,200,202,261,262,264,265,514,522; W. J. Van Aardt, unpublished data), arachnids 22,258,347,374,433, and insects 41,42,43,44,210,212,259,263,266,301,343,345,346,347,368,427; insects with elevated body temperature 37,209; and aerial groups such as insects 36,39,42,91,92,93,96,100,102,104,108,10,173,250,251,273,318,333,384,393,427,447,502,507,521.

Figure 13. Figure 13.

Mass‐specific metabolic cost of transport as a function of body mass. Values represent minimum cost of locomotion available. Shaded area represents hydrostatic crawlers and burrowers. Data include runners 41,43,44,55,159,191,199,200,201,209,212,259,260,263,264,265,301,343,345,346,347, crawlers 97,143,267,277,292, burrowers 54,291,469, swimmers 119,184,241,276,278,280,293,328,369,468,474,511, jetters 130,338,395,399,495, and fliers 173,273,478,483,503,521.

Figure 14. Figure 14.

Hypothesized relationships of invertebrate locomotion based on literature review. A: Mass‐specific mechanical power output (see Fig. 5), the product of cycle frequency and mass‐specific mechanical work output per cycle. B: Cycle frequency (see Fig. 4). C: Mass‐specific metabolic power input (see Fig. 11), the product of cycle frequency and mass‐specific metabolic cost input per cycle. D: Mass‐specific mechanical work output per cycle, the product of cycle distance and mass‐specific mechanical work output per cycle, the product of cycle distance and mass‐specific resistive force or mass‐specific mechanical energy of transport. E: Cycle distance. F: Mass‐specific metabolic cost input per cycle (see Fig. 12), the product of cycle distance and mass‐specific metabolic cost of transport. G: Mass‐specific resistive force or mass‐specific mechanical energy of transport. H: Speed (see Fig. 3), the product of cycle frequency and cycle distance. I: Mass‐specific metabolic cost of transport (see Fig. 13).

Figure 15. Figure 15.

Mass‐specific metabolic cost input per cycle as a function of body mass. Regression lines represent a range of body masses for a given species. Data include runners 22,190,209,210,212,261, a crawler 97, and fliers 42,251,273,447,502,507 such as moths 96 and bees 96,102,521.

Figure 16. Figure 16.

Trends for variation in power output. Two hypothetical animals of the same mass are compared. The morphology of a hypothetical appendage is shown to illustrate mechanical advantage, muscle strain, and the angle swept by the appendage. Left column represents trends for an animal that generates lower power output than the animal represented at right. Equations show how variables (muscle moment, joint angle, and muscle power output) are used to derive mass‐specific muscle power output (PM*). See text for explanation.

Figure 17. Figure 17.

Cyber‐invertebrates. Dynamic models of legged animals designed by M. Raibert, MIT, and Boston Dynamics in collaboration with R. J. Full at U.C. Berkeley. Simulations consist of equations of motion, ground contact models, a numerical integrator, and a three‐dimensional graphics program. All models obey the laws of Newtonian physics and are not simple kinematic representations. The Hexahopper (center) is an abstracted insect with telescoping legs attached to a hemispherical body. The model exhibits dynamic locomotion without an aerial phase as it bounces along. Three legs act as one spring‐mass system. The Hexabug (right) uses articulated legs attached to a long body. The legs have the length dimensions of cockroach legs. The Playback cockroach (left) uses the actual morphology of cockroach legs along with three‐dimensional kinematics during running. The kinematic motion data can be played through the control system and the resulting dynamics analyzed. The controller uses the motion data to drive servos at each joint to apply moments which attempt to maintain target positions.



Figure 1.

Schematic diagram showing integration of comparative bioenergetics, exercise physiology, functional morphology, muscle physiology, and comparative biomechanics. A: Energy available from several sources is transduced to segments (appendages or body sections) through muscles, depending on geometry. The chapter is organized in sections from right to left. B: Magnification of musculoskeletal role in transducing energy. Musculoskeletal parameters determine musculoskeletal forces. Musculoskeletal forces and joint geometry determine moment. Moment gives rise to acceleration, velocity, and angle change. Velocity and angle change (change in muscle length) feedback to affect musculoskeletal force production. Moment and joint angular velocity determine amount of energy transferred. Direction of movement and moment determine whether energy enters, leaves, or is transferred from a segment. Segmental energy and morphology of all segments determine power output of whole body and represent locomotor mechanical energy in A.



Figure 2.

Legs of animals in general, and of invertebrates in particular, can provide biological inspiration for the design of new robot legs. A: Legs operating in a more upright posture in a vertical plane parallel to the body (horizontal first joint axis for body–leg attachment), as seen in many birds and mammals, are gravitationally loaded and muscles must bear part of the body's weight 24. Animals using a horizontal first axis, however, can take advantage of gravity in swinging their legs. B–D: Legs operating in a sprawled posture (vertical first joint axis for body–leg attachment, like some lizards, amphibians, and arthropods) can potentially decouple gravitational loading of muscles from moving forward. To move forward, vertical first axis legs must project out to the side, resulting in increased static stability. Robots inspired from the design of cockroaches 62,63, whose legs operate in a horizontal plane, demonstrate a linear step, large step lengths, and proficiency at climbing.

adapted from ref. 62


Figure 3.

Speed as a function of body mass. A: Speeds of fliers, jumpers, runners, jetters, swimmers, and crawlers. Shown are maximum speeds available from the literature. Some speeds are reported as truly maximum. The majority are near maximum aerobic speed (speed at which maximal oxygen consumption is attained). Many are preferred speeds. For jumpers, take‐off speeds shown. The shaded area bounds the speed of crawlers. Data include runners 22,41,69,83,134,136,181,199,201,202,207,208,212,259,261,263,264,265,266,278,290,343,344,345,346,347,383,392,413,458,523, crawlers 97,145,277,292,353, jumpers 11,48,53,179,180,282,352, swimmers 7,65,114,119,184,241,276,278,280,290,293,328,369,386,409,466,468,482,511, jetters 106,127,128,130,132,294,338,395,398,399,473,474,475, and fliers 45,152,155,156,173,273,367. (Ref. 158 published after submission of figures.) B: Regression lines of major taxonomic groups (Table 1). Data to generate regressions include insect runners 41,136,181,207,208,212,259,263,266,343,344,345,346,347,383,392,458; crustacean runners 69,83,199,202,261,264,265,278,290,523; crustacean swimmers 7,119,184,241,276,280,293,328,369,386,409,466,468,482; crustacean underwater runners 114,276,278,280,290; bee, fly, and mosquito fliers 156,173,273; and lepidopteran fliers 152,155. Note the change in speed axis from A.



Figure 4.

Cycle frequency as a function of body mass. A: Cycle frequency of fliers, runners, jetters, and swimmers. Frequencies shown are the maximum available from the literature. Some frequencies are reported as truly maximum. The majority are preferred speeds. Data include runners 22,69,134,207,208,212,261; swimmers 7,65,114,290,409; jetters 29,106,127,140,338,473,495; and fliers such as mosquitoes 84,273,366,507, aphids, white flies 84,524, dragonflies 84,366,367, bees, wasps 84,96,156,171,306,366,507, beetles 84,507, flies 84,273,318,366,507,524, flower flies 84,318,507, butterflies 84,152,507, saturnid moths 39,84,96,507, sphinx moths 94,96,366,507, and crane flies 84,171,366,507. B: Regression lines of major taxanomic groups (Table 2). Note the change in cycle frequency axis from A.



Figure 5.

Whole‐animal mass‐specific mechanical power output as a function of body mass. Shaded areas and connected points represent the range of possible values for fliers, assuming 0 (upper bound) and 100% (lower bound) elastic storage. Speeds selected are the maximum available from the literature. Some speeds are reported as truly maximum. Many are preferred speeds. Data include insects 207,208 and crabs 69 for terrestrial locomotion; crustaceans 385,467, insects (65, 3890), jellyfish 140, and molluscs 127,397,398 for aquatic locomotion; and insects 94,102,153,173,299,504 for aerial locomotion.



Figure 6.

Sarcomere morphometrics. Muscles were divided into fast and slow groups based on a variety of criteria, which included fiber type, enzyme level or activity, and contraction kinetics. In general, if a study designated a muscle as fast or slow, this characterization was used. All insect flight muscles were considered to be fast. All crawlers—larvae, worms, and nematodes—have slow muscles. Walking and swimming organisms have both fiber types. Bars represent ± 1 S.D. Numbers in parentheses are sample size. Lmyo, myosin filament length; Lcsarc, length of a sarcomere. A: Sarcomere length. Data include insects such as locusts 48,51,285,504, cockroaches 163,185,287,296,382,415,454,479, water bugs 5,25,163, water beetles 162, katydids 5,164, lepidopterans 89,422, dragonflies 444, larvae 86,88,243,401,422, nymphs 87, flies 405,434, and hemipterans 125; crustaceans such as lobsters 287,295,296,415, crayfish 1,415,461,528, crabs 188,283,476, and merostomatans such as horseshoe crabs 490; arachnids such as spiders 321,436 and scorpions 26,425; molluscs such as scallops 379 and cuttlefish 286; annelids such as worms 376,462 and polychaetes 365; chaetognaths such as arrow worms 161; and coelenterates such as jellyfish 107,439. B: Myosin filament length. Data include insects such as locusts 118,285, cockroaches 5,163,239,240,287,296,415,479, water bugs 5,25,163, water beetles 162, katydids 5,164, lepidopterians 5,513, dragon‐flies 444, larvae 86,88,243,401, and nymphs 87,90; crustaceans such as lobsters 295,296,297,415, crayfish 415,528, and crabs 188,283,476; arachnids such as spiders 321,436; molluscs such as scallops 379 and cuttlefish 286; annelids such as worms 376,462, and polychaetes 365; nematodes 426; and coelenterates such as jellyfish 107,439. C; Ratio of thin/thick filament number. Data include insects such as locusts 118,285, cockroaches 5,239,240,287,296,415,454,479, water bugs 5,25, katydids 5,164, lepidopterians 89,422,513, dragonflies 444, larvae 86,88,243,401,422, nymphs 87,90, and flies 405; crustaceans such as lobsters 287,296,415 and crabs 188,283; arachnids such as spiders 436 and scorpions 26,425; molluscs such as scallops 379 and cuttlefish 286; annelids 376,462; nematodes 426; and coelenterates such as jellyfish 107. D: Sarcomere diagram shows designated lengths.



Figure 7.

Active length‐tension or strain‐stress curves of invertebrate muscles. Stress is normalized to peak isometric tension. Strain is normalized as a fraction of the length that gives peak isometric stress. Shaded areas represent strains that correspond to stresses above 50% maximal isometric stress. Data include bee flight 242, locust flight 353, crayfish 528, frog 271, leech 378, and fly larvae 243 muscles.



Figure 8.

Mechanical power output of invertebrate muscle and its determinants as a function of cycle frequency. Data are from isolated muscle studies using the work‐loop method (Fig. 9) and come from unpublished data of Stevenson and Casey. A: Mass‐specific mechanical power output and work per cycle. Gilmour and Ellington 223 have demonstrated values as high as 110 W/kg from asynchronous, glycerinated muscle fibers of bumblebees. B: Muscle stress, strain, and strain rate. Data include insects 312,349,380,453, crustaceans 455, and molluscs 362.



Figure 9.

Work‐loop method to determine power output of an isolated muscle. A: Muscle is subjected to cyclic length changes (strain) while force per area (stress) is measured. The stimulation pattern is controlled. B: Area under the stress‐strain curve represents energy produced (output) during shortening or absorbed (input) during lengthening. The difference between the curves represents the net work per cycle. Counterclockwise loops result in energy production, whereas clockwise loops result in energy absorption by the muscle. C: Variation in work‐loop shape. Rectangular shape observed at low frequencies (<30 Hz). Triangular shape observed at intermediate frequencies (30–60 Hz). Ellipsoid shape observed at high frequencies (60–180 Hz).

after ref. 311


Figure 10.

Three‐dimensional musculoskeletal model of themeta‐thoracic leg of the cockroach Blaberus discoidalis. Left: Shaded polygons represent the exoskeleton, which was reconstructed from serial sections. Right: Heavy lines represent the lines of action of Hill‐type muscles. The model is articulated at the coxa/trochanter–femur and femurtibia joints so that muscle lengths, moment arms, forces, and joint moments can be estimated for a range of body positions 195. The computer model was created using SIMM.

MusculoGraphics, Evanston, IL


Figure 11.

Whole‐animal mass‐specific metabolic power input as a function of body mass. Oxygen consumption rates are the highest available for the species. In some cases, they are known to be maximal rates. A: Oxygen consumption as a function of body mass for runners, crawlers, swimmers, jetters, and fliers. Shaded areas represent jetters; top right area includes molluscs such as squid; bottom area includes jellyfish. Insect runners can warm their bodies. The regression line shown is for the Q10 corrected data set not shown (Q10 = 2). Data include runners (22,41,42,43,44,190,199,200,201,202,210,212,258,259,261,262,263,264,265,266,301,343,345,346,347,368,374,427,433,514,522; W. J. Van Aardt, unpublished data), runners with elevated body temperature 37,209, crawlers 97,277,292, swimmers 119,184,276,278,279,293,328,369,371,428,466, jetters 29,130,338,395,399,474,495,511, and fliers 36,39,42,91,92,93,96,100,102,104,108,109,110,173,250,251,273,318,333,384,393,427,447,502,507,521. B: Oxygen consumption rates by taxonomic groups (Table 3). Data include insect runners 41,42,43,44,210,212,259,263,266,301,343,345,346,347,368,427; crustacean runners (190,199,200,202,261,262,264,265,514,522; W. J. Van Aardt, unpublished data); swimmers 119,184,276,278,279,293,328,369,371,428,466,474; moth fliers 39,91,92,93,96,100; and bee, fly, and mosquito fliers 36,42,96,102,104,108,109,110,173,250,251,273,318,333,384,393,427,447,502,507,521.



Figure 12.

Aerobic factorial scopes for invertebrate activity. Oxygen consumption rates are the highest available for the species. In some cases, they are known to be maximal rates. Bars represent ± 1 S.D. Numbers in parentheses are sample size. Data include aquatic groups such as coelenterates 338, gastropods 277,292, crustaceans 119,184,276,278,279,293,328,369,371,428,466, and cephalopods 29,395,399,495,511; terrestrial groups such as gastropods 277,292, myriapods 201, crustaceans (190,199,200,202,261,262,264,265,514,522; W. J. Van Aardt, unpublished data), arachnids 22,258,347,374,433, and insects 41,42,43,44,210,212,259,263,266,301,343,345,346,347,368,427; insects with elevated body temperature 37,209; and aerial groups such as insects 36,39,42,91,92,93,96,100,102,104,108,10,173,250,251,273,318,333,384,393,427,447,502,507,521.



Figure 13.

Mass‐specific metabolic cost of transport as a function of body mass. Values represent minimum cost of locomotion available. Shaded area represents hydrostatic crawlers and burrowers. Data include runners 41,43,44,55,159,191,199,200,201,209,212,259,260,263,264,265,301,343,345,346,347, crawlers 97,143,267,277,292, burrowers 54,291,469, swimmers 119,184,241,276,278,280,293,328,369,468,474,511, jetters 130,338,395,399,495, and fliers 173,273,478,483,503,521.



Figure 14.

Hypothesized relationships of invertebrate locomotion based on literature review. A: Mass‐specific mechanical power output (see Fig. 5), the product of cycle frequency and mass‐specific mechanical work output per cycle. B: Cycle frequency (see Fig. 4). C: Mass‐specific metabolic power input (see Fig. 11), the product of cycle frequency and mass‐specific metabolic cost input per cycle. D: Mass‐specific mechanical work output per cycle, the product of cycle distance and mass‐specific mechanical work output per cycle, the product of cycle distance and mass‐specific resistive force or mass‐specific mechanical energy of transport. E: Cycle distance. F: Mass‐specific metabolic cost input per cycle (see Fig. 12), the product of cycle distance and mass‐specific metabolic cost of transport. G: Mass‐specific resistive force or mass‐specific mechanical energy of transport. H: Speed (see Fig. 3), the product of cycle frequency and cycle distance. I: Mass‐specific metabolic cost of transport (see Fig. 13).



Figure 15.

Mass‐specific metabolic cost input per cycle as a function of body mass. Regression lines represent a range of body masses for a given species. Data include runners 22,190,209,210,212,261, a crawler 97, and fliers 42,251,273,447,502,507 such as moths 96 and bees 96,102,521.



Figure 16.

Trends for variation in power output. Two hypothetical animals of the same mass are compared. The morphology of a hypothetical appendage is shown to illustrate mechanical advantage, muscle strain, and the angle swept by the appendage. Left column represents trends for an animal that generates lower power output than the animal represented at right. Equations show how variables (muscle moment, joint angle, and muscle power output) are used to derive mass‐specific muscle power output (PM*). See text for explanation.



Figure 17.

Cyber‐invertebrates. Dynamic models of legged animals designed by M. Raibert, MIT, and Boston Dynamics in collaboration with R. J. Full at U.C. Berkeley. Simulations consist of equations of motion, ground contact models, a numerical integrator, and a three‐dimensional graphics program. All models obey the laws of Newtonian physics and are not simple kinematic representations. The Hexahopper (center) is an abstracted insect with telescoping legs attached to a hemispherical body. The model exhibits dynamic locomotion without an aerial phase as it bounces along. Three legs act as one spring‐mass system. The Hexabug (right) uses articulated legs attached to a long body. The legs have the length dimensions of cockroach legs. The Playback cockroach (left) uses the actual morphology of cockroach legs along with three‐dimensional kinematics during running. The kinematic motion data can be played through the control system and the resulting dynamics analyzed. The controller uses the motion data to drive servos at each joint to apply moments which attempt to maintain target positions.

References
 1. Abbott, B. C., and I. Parnas. Electrical and mechanical responses in deep abdominal extensor muscles of crayfish and lobster. J. Gen. Physiol. 48: 919–931, 1965.
 2. Adamczewska, A. M., and S. Morris. Exercise in the terrestrial Christmas Island red crab Gecarcoidea natalis. J. Exp. Biol. 188: 257–274, 1994.
 3. Ahn, A. N., and R. J. Full. Multiple muscle kinematic simulation of running roaches. Physiologist 37: A86, 1994.
 4. Aidley, D. J. Excitation‐contraction coupling and mechanical properties. In: Insect Muscle, edited by P. N. R. Usherwood. London: Academic, 1975, p. 337–356.
 5. Aidley, D. J. Muscular contraction. In: Comprehensive Insect Physiology, Biochemistry, and Pharmacology. Nervous System: Structure and Motor Function, edited by G. A. Kerkut and L. I. Gilbert, New York: Pergamon, 1985, p. 407–437.
 6. Alexander, D. E. Kinematics of swimming in two species of Idotea (Isopoda: Valvifera). J. Exp. Biol. 138: 37–49, 1988.
 7. Alexander, D. E., and T. Chen. Comparison of swimming speed and hydrodynamic drag in two species of Idotea (Isopoda). J. Crust. Biol. 10: 406–412, 1990.
 8. Alexander, R. M. Rubber‐like properties of the inner hinge‐ligament of Pectinidae. J. Exp. Biol. 44: 119–130, 1966.
 9. Alexander, R. M. Terrestrial locomotion. In: Mechanics and Energetics of Animal Locomotion, edited by R. M. Alexander and T. Goldspink. New York: Wiley, 1977, p. 168–203.
 10. Alexander, R. M. Locomotion of Animals. Tertiary Level Biology edited by R. M. Alexander, Glasgow: Blackie, 1982.
 11. Alexander, R. M. (ed). Animal Mechanics (2nd ed.), Biology Series Oxford: Blackwell, 1983.
 12. Alexander, R. M. The maximum forces exerted by animals. J. Exp. Biol. 115: 231–238, 1985.
 13. Alexander, R. M. Bending of cylindrical animals with helical fibres in their skin or cuticle. J. Theor. Biol. 124: 97–110, 1987.
 14. Alexander, R. M. Elastic Mechanisms in Animal Movement Cambridge: Cambridge Univ. Press, 1988.
 15. Alexander, R. M. Three uses for springs in legged locomotion. Int. J. Rob. Res. 9: 53–61, 1990.
 16. Alexander, R. M. Exploring Biomechanics New York: Scientific American Library, 1992.
 17. Alexander, R. M. (Ed). Mechanics of Animal Locomotion, Advances in Comparative and Environmental Physiology Berlin: Springer‐Verlag, 1992, vol. 11.
 18. Alexander, R. M., and H. C. Bennet‐Clark. Storage of elastic strain energy in muscle and other tissues. Nature 265: 114–117, 1977.
 19. Alexander, R. M., and G. Goldspink (Eds). Mechanics and Energetics of Animal Locomotion London: Chapman and Hall, 1977.
 20. Anderson, B. D., and R. J. Full. Locomotor mechanics of the centipede Scolopendra heros. Am. Zool. 34: 45A, 1994.
 21. Anderson, B. D., J. W. Shultz, and B. C. Jayne. Axial kinematics and muscle activity during terrestrial locomotion of the centipede Scolpendra heros. J. Exp. Biol. 198: 1185–1195, 1995.
 22. Anderson, J. F., and K. N. Prestwich. The physiology of exercise at and above maximal aerobic capacity in a theraphosid (tarantula) spider, Brachypelma simihi (F.O. Pickard‐Cambridge). J. Comp. Physiol. [B] 155: 529–539, 1985.
 23. Angle, C. Genghis: A Six‐Legged Autonomous Walking Robot Cambridge, MA: MIT, 1989.
 24. Angle, C. Design of an artificial creature Cambridge, MA: MIT, 1991.
 25. Ashhurst, D. E. The fibrillar flight muscles of giant water‐bugs: an electron‐microscope study. J. Cell Sci. 2: 435–444, 1967.
 26. Auber‐Thomay, M. Remarques sur l'ultrastructure des myofibrilles chez des scorpions. J. Microsc. (Paris) 2: 233–236, 1963.
 27. Azuma, A., and T. Watanabe. Flight performance of a dragonfly. J. Exp. Biol. 137: 221–252, 1988.
 28. Baldwin, J., and W. R. England. The properties and functions of alanopine dehydrogenase and octopine dehydrogenase from the pedal retractor muscle of Strombidae (class Gastropoda). Pacific Sci. 36: 381–394, 1982.
 29. Baldwin, J., and A. K. Lee. Contributions of aerobic and anaerobic energy production during swimming in the bivalve mollusc Limaria fragilis (family Limidae). J. Comp. Physiol. [B] 129: 361–364, 1979.
 30. Baldwin, J., and G. M. Morris. Re‐examination of the contributions of aerobic and anaerobic energy production during swimming in the bivalve mollusc Limaria fragilis (family Limidae). Aust. J. Marine Freshw. Res. 34: 909–914, 1983.
 31. Barlow, D., and M. A. Sleigh. Water propulsion speeds and power output by comb plates of the ctenophore Pleurobrachia pileus under different conditions. J. Exp. Biol. 183: 149–163, 1993.
 32. Barlow, D., M. A. Sleigh, and R. J. White. Water flows around the comb plates of the ctenophore Pleurobrachia plotted by computer: a model system for studying propulsion by antiplectic metachronism. J. Exp. Biol. 177: 113–128, 1993.
 33. Barnes, W.J.P. Leg co‐ordination during walking in the crab, Uca pugnax. J. Comp. Physiol. 96: 237–256, 1975.
 34. Barnes, W.J.P. Nervous control of locomotion in Crustacea. In: Simple Nervous Systems, edited by P. N. R. Usherwood and D. R. Newth. London: Arnold, 1975, p. 415–441.
 35. Barr, D., and B. P. Smith. Stable swimming by diagonal phase synchrony in arthropods. Can. J. Zool. 58: 782–795, 1980.
 36. Bartholomew, G. A., and M. C. Barnhart. Tracheal gases, respiratory gas exchange, body temperature and flight in some tropical cicadas. J. Exp. Biol. 111: 131–144, 1984.
 37. Bartholomew, G. A., and T. M. Casey. Body temperature and oxygen consumption during rest and activity in relation to body size in some tropical beetles. J. Therm. Biol. 2: 173–176, 1977.
 38. Bartholomew, G. A., and T. M. Casey. Endothermy during terrestrial activity in large beetles. Science 195: 882–883, 1977.
 39. Bartholomew, G. A., and T. M. Casey. Oxygen consumption of moths during rest, pre‐flight warm‐up, and flight in relation to body size and wing morphology. J. Exp. Biol. 76: 11–25, 1978.
 40. Bartholomew, G. A., and B. Heinrich. Endothermy in African dung beetles during flight, ball making, and ball rolling. J. Exp. Biol. 73: 65–83, 1978.
 41. Bartholomew, G. A., and J.R.B. Lighton. Ventilation and oxygen consumption during rest and locomotion in a tropical cockroach, Blaberus giganticus. J. Exp. Biol. 118: 449–454, 1985.
 42. Bartholomew, G. A., and J.R.B. Lighton. Endothermy and energy metabolism of a giant tropical fly, Pantophthalmus tabaninus Thunberg. J. Comp. Physiol. [B] 156: 461–467, 1986.
 43. Bartholomew, G. A., J.R.B. Lighton, and D. H. Feener. Energetics of trail running, load carriage, and emigration in the column‐raiding army ant Eciton hamatum. Physiol. Zool. 61: 57–68, 1988.
 44. Bartholomew, G. A., J.R.B. Lighton, and G. N. Louw. Energetics of locomotion and patterns of respiration in tenebrionid beetles from the Namib Desert. J. Comp. Physiol. [B] 155: 155–162, 1985.
 45. Bauer, C. K., and M. Gewecke. Flight behaviour of the water beetle Dytiscus marginalis L. (Coleopera, Dytiscidae). In: Insect Locomotion, edited by M. Gewecke and G. Wendler. Berlin: Verlag Paul Parey, 1985, p. 205–214.
 46. Beer, R. D., N. J. Chiel, R. D. Quinn, and P. Larsson. A distributed neural network architecture for hexapod robot locomotion. Neural Comput. 4: 356–365, 1992.
 47. Beis, I., and E. A. Newsholme. The contents of adenine nucleotides, phosphagens and some glycolytic intermediates in resting muscles from vertebrates and invertebrates. Biochem. J. 152: 23–32, 1975.
 48. Bennet‐Clark, H. C. The energetics of the jump of the locust, Schistocerca gregaria. J. Exp. Biol. 63: 53–83, 1975.
 49. Bennet‐Clark, H. C. Energy storage in jumping animals. In: Perspectives in Experimental Biology, edited by S. Davies. Oxford: Pergamon, 1976, p. 467–479.
 50. Bennet‐Clark, H. C. Aerodynamics of insect jumping. In: Aspects of Animal Movement, edited by H. Y. Elder and E. R. Trueman, Cambridge: Cambridge Univ. Press, 1980, p. 151–167.
 51. Bennet‐Clark, H. C. Jumping in Orthoptera. In: Biology of Grasshoppers, edited by R. F. Capman and A. Joern. New York: Wiley, 1990, p. 173–203.
 52. Bennet‐Clark, H. C., and G. M. Alder. The effect of air resistance on the jumping performance of insects. J. Exp. Biol. 82: 105–121, 1979.
 53. Bennet‐Clark, H. C., and E.C.A. Lucey. The jump of the flea; a study of the energetics and a model of the mechanism. J. Exp. Biol. 47: 59–76, 1967.
 54. Berrigan, D., and J.R.B. Lighton. Bioenergetic and kinematic consequences of limblessness in larval Diptera. J. Exp. Biol. 179: 245–259, 1993.
 55. Berrigan, D., and J.R.B. Lighton. Energetics of pedestrian locomotion in adult male blowflies, Protophormia terraenovae (Diptera: Calliphoridae). Physiol. Zool. 67: 1140–1153, 1994.
 56. Bessonov, A. P., and N. V. Umnov. Features of kinematics of turn of walking vehicles. Program, 3rd CISM‐IFToMM Symposium on theory and practice of robots and manipulators, 1978.
 57. Biewener, A. A. Mammalian terrestrial locomotion and size. Bioscience 39: 776–783, 1989.
 58. Biewener, A. A. Scaling body support in mammals: limb posture and muscle mechanics. Science 245: 45–48, 1989.
 59. Biewener, A. A. (Ed). Biomechanics: Structures and Systems, A Practical Approach Practical Approach Series Oxford: IRL Oxford Univ. Press, 1992.
 60. Biewener, A. A., and R. J. Full. Force platform and kinematic analysis. In: Biomechanics: Structures and Systems, A Practical Practical Approach Series, edited by A. A. Biewener, Oxford: IRL Oxford Univ. Press, 1992, p. 45–73.
 61. Bill, R. G., and W. F. Herrnkind. Drag reduction by formation movement in spiny lobsters. Science 193: 1146–1148, 1976.
 62. Binnard, M. B. Leg Design for a Small Walking Robot Cambridge, MA: MIT, 1992.
 63. Binnard, M. B. Design of a Small Pneumatic Walking Robot Cambridge, MA: MIT, 1995.
 64. Blake, R. W. Crab carapace hydrodynamics. J. Zool. 207: 407–423, 1985.
 65. Blake, R. W. Hydrodynamics of swimming in the water boatman, Cenocorixa bifida. Can. J. Zool. 64: 1606–1613, 1986.
 66. Blake, R. W. (Ed). Efficiency and Economy in Animal Physiology, Cambridge: Cambridge Univ. Press, 1991.
 67. Blickhan, R. The spring‐mass model for running and hopping. J. Biomech. 22: 1217–1227, 1989.
 68. Blickhan, R., and F. G. Barth. Strains in the exoskeleton of spiders. J. Comp. Physiol. [A] 157: 115–147, 1985.
 69. Blickhan, R., and R. J. Full. Locomotion energetics of the ghost crab II. Mechanics of the center of mass during walking and running. J. Exp. Biol. 130: 155–174, 1987.
 70. Blickhan, R., and R. J. Full. Mechanical work in terrestrial locomotion. In: Biomechanics: Structures and Systems. A Practical Approach. Practical Approach Series edited by A. A. Biewener, Oxford: IRL Oxford Univ. Press, 1992, p. 75–96.
 71. Blickhan, R., and R. J. Full. Similarity in multilegged locomotion: bouncing like a monopode. J Comp. Physiol. [A] 173: 509–517, 1993.
 72. Blickhan, R., R. J. Full, and L. H. Ting. Exoskeletal strain: evidence for a trot‐gallop transition in rapid running ghost crabs. J. Exp. Biol. 179: 301–321, 1993.
 73. Bone, Q. Jet propulsion in salps (Tunicata, Thaliacea). J. Zool. 201: 481–506, 1983.
 74. Booth, C. E., and B. R. McMahon. Lactate dynamics during locomotor activity in the blue crab, Callinectes sapidus. J. Exp. Biol. 118: 461–465, 1985.
 75. Booth, C. E., and B. R. McMahon. Aerobic capacity of the blue crab, Callinectes sapidus. 65: 1074–1091, 1992.
 76. Booth, C. E., B. R. McMahon, P. L. de Fur, and P.R.H. Wilkes. Acid‐base regulation during exercise and recovery in the blue crab, Callinectes sapidus. Respir. Physiol. 58: 359–376, 1984.
 77. Booth, C. E., B. R. McMahon, and A. W. Pinder. Oxygen uptake and the potentiating effects of increased hemolymph lactate on oxygen transport during exercise in the blue crab, Callinectes sapidus. J. Comp. Physiol. 148: 111–121, 1982.
 78. Bowdan, E. Walking and rowing in the water strider, Gerris remigis. II. Muscle activity associated with slow and rapid mesothoracic leg movement. J. Comp. Physiol. [A] 123: 51–57, 1978.
 79. Bowerman, R. F. The control of arthropod walking. Comp. Biochem. Physiol. A 56: 231–247, 1977.
 80. Bowerman, R. F. Arachnid locomotion. In: Locomotion and Energetics in Arthropods, edited by C. F. Herreid II and C. R. Fourtner. New York: Plenum, 1981, p. 73–102.
 81. Brodsky, A. K. Vortex formation in the tethered flight of the peacock butterfly Inachis io L. (Lepidoptera, Nymphalidae) and some aspects of insect flight evolution. J. Exp. Biol. 161: 77–95, 1991.
 82. Brown, A. C. Oxygen diffusion into the foot of the whelk Bullia digitalis (Dillwyn) and its possible signifigance in respiration. J. Exp. Mar. Biol. Ecol. 79: 1–7, 1984.
 83. Burrows, M., and G. Hoyle. The mechanism of rapid running in the ghost crab, Ocypode ceratophthalma. J. Exp. Biol. 58: 327–349, 1973.
 84. Byrne, D. N., S. L. Buchmann, and H. G. Spangler. Relationship between wing loading, wingbeat frequency and body mass in homopterous insects. J. Exp. Biol. 135: 9–23, 1988.
 85. Caldwell, R. L. A unique form of locomotion in a stomatopod—backward somersaulting. Nature 282: 71–73, 1979.
 86. Carnevali, D. C., and R. Valvassori. Intersegmental abdominal muscles of Calopterix splendens and Anax imperator. J. Submicrosc. Cytol. 5: 227–236, 1973.
 87. Carnevali, M.D.C. Ultrastructural pattern differences in some muscle fibres of nymphal Ephemera danica (Ephemeroptera). J. Submicrosc. Cytol. 7: 219–230, 1975.
 88. Carnevali, M.D.C. Z‐line and supercontraction in the hydraulic muscular systems of insect larvae. J. Exp. Zool. 203: 15–30, 1978.
 89. Carnevali, M.D.C., and J. F. Reger. Slow‐acting flight muscles of saturniid moths. J. Ultrastruct. Res. 79: 241–249, 1982.
 90. Carnevali, M.D.C., and A. Saita. Atypical myofilament arrangement in some abdominal muscles of mayfly nymphs. Program, II International conference on Ephemeroptera, Warsawa‐Krakowl, 1979.
 91. Casey, T. M. Flight energetics in sphinx moths: heat production and heat loss in Hyles lineata during free flight. J. Exp. Biol. 64: 545–560, 1976.
 92. Casey, T. M. Flight energetics of sphinx moths: power input during hovering flight. J. Exp. Biol. 64: 529–543, 1976.
 93. Casey, T. M. Flight energetics and heat change of gypsy moths in relation to air temperature. J. Exp. Biol. 88: 133–145, 1980.
 94. Casey, T. M. A comparison of mechanical and energetic estimates of flight cost for hovering sphinx moths. J. Exp. Biol. 91: 117–129, 1981.
 95. Casey, T. M. Insect flight energetics. In: Locomotion and Energetics in Arthropods, edited by C. F. Herreid II and C. R. Fourtner. New York: Plenum, 1981, p. 419–452.
 96. Casey, T. M. Oxygen consumption during flight. In: Insect Plight, edited by G. Goldsworthy and C. Wheeler. Boca Raton, FL: CRC, 1989, p. 257–272.
 97. Casey, T. M. Energetics of caterpillar locomotion: biomechanical constraints of a hydraulic skeleton. Science 252: 112–114, 1991.
 98. Casey, T. M. Energetics of locomotion. In: Mechanics of Animal Locomotion, edited by R. M. Alexander, Berlin: Springer‐Verlag, 1992, p. 251–275.
 99. Casey, T. M., and C. P. Ellington. Energetics of insect flight. In: Energy Transformations in Cells and Animals. Proc. 10th Conf. Eur. Soc. Comp. Physiol. Biochem, edited by W. Wieser and E. Gneiger. Innsbruck: Georg Thieme‐Verlag, 1989, p. 200–210.
 100. Casey, T. M., J. R. Hegel, and C. S. Buser. Physiology and energetics of pre‐flight warm‐up in the eastern tent caterpillar moth Malacosoma americanum. J. Exp. Biol. 94: 119–135, 1981.
 101. Casey, T. M., and B. A. Joos. Morphometrics, conductance, thoracic temperature, and flight energetics of noctuid and geometrid moths. Physiol. Zool. 56: 160–173, 1983.
 102. Casey, T. M., and M. L. May. Flight energetics of euglossine bees in relation to morphology and wing stroke frequency. J. exp. Biol. 116: 271–289, 1985.
 103. Cavagna, G. A., N. C. Heglund, and C. R. Taylor. Mechanical work in terrestrial locomotion: two basic mechanisms for minimizing energy expenditure. Am. J. Physiol. 233 (Regulatory Integrative Comp. Physiol. 4): R243–R261, 1977.
 104. Chadwick, L. E., and D. Gilmour. Respiration during flight in Drosophila repleta Wollaston: the oxygen consumption considered in relation to the wing‐rate. Physiol. Zool. 13: 398–410, 1940.
 105. Chamberlain, J. A., Jr. Motor performance and jet propulsion in Nautilus: implications for cephalopod paleobiology and evolution. Bull. Am. Malacol. Union 1980: 37–42, 1980.
 106. Chamberlain, J. A., Jr. Locomotion in Nautilus. In: Nautilus: The Biology and Paleobiology of a Living Fossil, edited by W. B. Saunders and N. H. Landman. New York: Plenum, 1987, p. 489–525.
 107. Chapman, D. M., C.F.A. Pantin, and E. A. Robson. Muscle in coelenterates. Rev. Can. Biol. 21: 267–278, 1962.
 108. Chappell, M. A. Temperature regulation of carpenter bees (Xylocopa californica) foraging in the Colorado Desert of southern California. Physiol. Zool. 55: 267–280, 1982.
 109. Chappell, M. A. Temperature regulation and energetics of the solitary bee Centris pallida during foraging and intermale mate competition. Physiol. Zool. 57: 215–225, 1984.
 110. Chappell, M. A., and K. R. Morgan. Temperature regulation, endothermy, resting metabolism, and flight energetics of tachinid flies (Nowickia sp.). Physiol. Zool. 60: 550–559, 1987.
 111. Cheer, A. Y. L., and M.A.R. Koehl. Paddles and rakes: fluid flow through bristled appendages of small organisms. J. Theor. Biol. 129: 17–39, 1987.
 112. Chih, C. P., and W. R. Ellington. Energy metabolism during contractile activity and environmental hypoxia in the phasic adductor muscle of the bay scallop Argopecten irradians concentricus. Physiol. Zool. 56: 623–631, 1983.
 113. Clarac, F. Decapod crustacean leg coordination during walking. In: Locomotion and Energetics in Arthropods, edited by C. F. Herreid II and C. R. Fourtner. New York: Plenum, 1981, p. 31–71.
 114. Clarac, F., and W.J.P. Barnes. Peripheral influences on the coordination of the legs during walking in decapod crustaceans. Semin. Soc. Exp. Biol. 24: 249–269, 1985.
 115. Clarac, F., and H. Cruse. Comparison of forces developed by the leg of the rock lobster when walking free or on a treadmill. Biol. Cybern. 43: 109–114, 1982.
 116. Clark, R. B., and D. J. Tritton. Swimming mechanisms in nereidiform polychaetes. J. Zool. 161: 257–271, 1970.
 117. Cloupeau, M., J. F. Devillers, and D. Devezeaux. Direct measurements of instantaneous lift in desert locust; comparison with Jensen's experiments on detached wings. J. Exp. Biol. 80: 1–15, 1979.
 118. Cochrane, D. G., H. Y. Elder, and P.N.R. Usherwood. Physiology and ultrastructure of phasic and tonic skeletal muscle fibres in the locust, Schistocerca gregaria. J. Cell Sci. 10: 419–441, 1972.
 119. Cowles, D. L., and J. J. Childress. Swimming speed and oxygen consumption in the bathypelagic mysid Gnathophausia ingens. Biol. Bull. 175: 111–121, 1988.
 120. Cruse, H. The function of the legs in the free walking stick insect, Carausius morosus. J. Comp. Physiol. [A] 112: 235–262, 1976.
 121. Cruse, H. A new model describing the coordination pattern of the legs of a walking stick insect. Biol. Cybern. 32: 107–113, 1979.
 122. Cruse, H. What mechanisms coordinate leg movement in walking arthropods?. Trends Nat. Sci. 13: 15–21, 1990.
 123. Cruse, H., et al. A modular artificial neural net for controlling a six‐legged walking system. Biol. Cybern. 72: 421–430, 1995.
 124. Cruse, H., and S. Epstein. Peripheral influences on the movement of the legs in a walking insect Carausius morosus. J. Exp. Biol. 101: 161–170, 1982.
 125. Cullen, M. J. The distribution of asynchronous muscle in insects with particular reference to the Hemiptera: an electron microscope study. J. Entomol. [A] 49: 17–41, 1974.
 126. Curtin, N. A., and R. C. Woledge. Power output and force‐velocity relationship of live fibres from white myotomal muscle of the dogfish, Scyliorhinus canicula. J. exp. Biol. 140: 187–197, 1988.
 127. Dadswell, M. J., and D. Weihs. Size‐related hydrodynamic characteristics of the giant scallop, Placopecten magellanicus (Bivalvia, Pectinidae). Can. J. Zool. 68: 778–785, 1990.
 128. Daniel, T. L. Mechanics and energetics of medusan jet propulsion. Can. J. Zool. 61: 1406–1420, 1983.
 129. Daniel, T. L. Unsteady aspects of aquatic locomotion. Am. Zool. 24: 121–134, 1984.
 130. Daniel, T. L. Cost of locomotion: unsteady medusan swimming. J. Exp. Biol. 119: 149–164, 1985.
 131. Daniel, T. L., C. Jordan, and D. Grunbaum. Hydromechanics of swimming. In: Advances in Comparative and Environmental Physiology edited by Alexander, R. M. Berlin: Springer‐Verlag, 1992, p. 17–49.
 132. Daniel, T. L., and E. Meyhofer. Size limits in escape locomotion of carridean shrimp. J. Exp. Biol. 143: 245–265, 1989.
 133. Dean, J. A model of leg coordination in the stick insect, Carausius morosus. II. Description of the kinematic model and simulation of normal step patterns. Biol. Cybern. 64: 403–411, 1991.
 134. Delcomyn, F. The locomotion of the cockroach Periplaneta americana. J. Exp. Biol. 54: 443–452, 1971.
 135. Delcomyn, F. Insect locomotion on land. In: Locomotion and Energetics of Arthropods, edited by C. F. Herreid II and C. R. Fourtner. New York: Plenum, 1981, p. 103–125.
 136. Delcomyn, F. Walking and running. In: Comprehensive Insect Physiology, Biochemistry, and Pharmacology, Nervous System: Structure and Motor Function, edited by G. A. Kerkut and L. I. Gilbert, New York: Pergamon, 1985, p. 439–466.
 137. De Mont, M. E. Tuned oscillations in the swimming scallop Pecten maximus. Can. J. Zool. 68: 786–791, 1990.
 138. De Mont, M. E. Locomotion in soft bodied animals. In: Advances in Comparative and Environmental Physiology, edited by Alexander, R. M. Berlin: Springer‐Verlag, 1992, p. 167–190.
 139. De Mont, M. E., and J. M. Gosline. Mechanics of jet propulsion in the hydromedusan jellyfish, Polyorchis penicillatus. I. Mechanical properties of the locomotor structure. J. Exp. Biol. 134: 313–332, 1988.
 140. De Mont, M. E., and J. M. Gosline. Mechanics of jet propulsion in the hydromedusan jellyfish, Polyorchis penicillatus. II. Energetics of the jet cycle. J. Exp. Biol. 134: 333–345, 1988.
 141. De Mont, M. E., and J. M. Gosline. Mechanics of jet propulsion in the hydromedusan jellyfish, Polyorchis penicillatus III. A natural resonating bell: the presence and importance of a resonant phenomenon in the locomotor structure. J. exp. Biol. 134: 347–361, 1988.
 142. De Mont, M. E., and E. I. Hokkanen. Hydrodynamics of animal movement. In: Biomechanics: Structures and Systems, A Practical Approach. Practical Approach Series, edited by A. A. Biewener, Oxford: IRL Oxford Univ. Press, 1992, p. 263–284.
 143. Denny, M. W. Locomotion: the cost of gastropod crawling. Science 208: 1288–1290, 1980.
 144. Denny, M. W. Mechanical properties of pedal mucus and their consequences for gastropod structure and performance. Am. Zool. 24: 23–36, 1984.
 145. Denny, M. W., and J. M. Gosline. The physical properties of the pedal mucus of the terrestrial slug, Ariolimax columbianus. J. Exp. Biol. 88: 375–393, 1980.
 146. Dickinson, M. H. The effect of wing rotation on unsteady aerodynamic performance at low Reynolds numbers. J. Exp. Biol. 192: 179–206, 1994.
 147. Dickinson, M. H., and K. G. Gotz. Unsteady aerodynamic performance of model wings at low Reynolds numbers. J. Exp. Biol. 174: 45–64, 1993.
 148. Dickinson, M. H., and J. Lighton. Muscle efficiency and elastic storage in the flight motor of Drosophila. Science 268: 87–90, 1995.
 149. Dobrolyubov, A. I. The mechanism of locomotion of some terrestrial animals by travelling waves of deformation. J. Theor. Biol. 119: 457–466, 1986.
 150. Drewes, C. D., and C. R. Fourtner. Helical swimming in a freshwater oligochaete. Biol. Bull. 185: 1–9, 1993.
 151. Du Bois, M. B., and R. Jander. Leg coordination and swimming in an ant, Campnotus americanus. Physiol. Entomol. 10: 267–270, 1985.
 152. Dudley, R. Biomechanics of flight in neotropical butterflies: morphometrics and kinematics. J. Exp. Biol. 150: 37–53, 1990.
 153. Dudley, R. Biomechanics of flight in neotropical butterflies: aerodynamics and mechanical power requirements. J. Exp. Biol. 159: 335–357, 1991.
 154. Dudley, R. Aerodynamics of flight. In: Biomechanics: Structures and Systems. A Practical Approach. Practical Approach Series, edited by A. A. Biewener, Oxford: IRL Oxford Univ. Press, 1992, p. 97–121.
 155. Dudley, R., and P. J. De Vries. Flight physiology of migrating Urania fulgens (Uraniidae) moths: kinematics and aerodynamics of natural free flight. J. Comp. Physiol. [A] 167: 145–154, 1990.
 156. Dudley, R., and C. P. Ellington. Mechanics of forward flight in bumblebees. I. Kinematics and morphology. J. Exp. Biol. 148: 19–52, 1990.
 157. Dudley, R., and C. P. Ellington. Mechanics of forward flight in bumblebees. II. Quasi‐steady lift and power requirements. J. Exp. Biol. 148: 53–88, 1990.
 158. Dudley, R., and R. B. Srygley. Flight physiology of neotropical butterflies: allometry of airspeeds during natural free flight. J. Exp. Biol. 191: 125–139, 1994.
 159. Duncan, F. D., and R. M. Crewe. A comparison of the energetics of foraging of three species of Leptogenys (Hymenoptera, Formicidae). Physiol. Entomol. 18: 372–378, 1993.
 160. Duncan, F. D., and J.R.B. Lighton. The burden within: the energy cost of load carriage in the honeypot ant, Myrmecocystus. Physiol. Zool. 67: 190–203, 1994.
 161. Duvert, M., and C. Salat. The primary body‐wall musculature in the arrow‐worm Sagitta setosa (Chaetognatha): an ultra‐structural study. Tissue Cell 12: 723–738, 1980.
 162. Edwards, G. A., P. De Souza Santos, H. Lopes De Souza Santos, and P. Sawaya I. Electron microscope studies of insect muscle. I. Flight and coxal muscle of Hydrophilus piceus. Ann. Entomol. S. Am. 47: 343–354, 1954.
 163. Edwards, G. A., P. De Souza Santos, H. Lopes De Souza Santos, and P. Sawaya. II. Electron microscope studies of insect muscle. II. Flight and leg muscles of Belostoma and Periplaneta. Ann. Entomol. S. Am. 47: 459–467, 1954.
 164. Elder, H. Y. High frequency muscles used in sound production by a katydid. II. Ultrastructure of the singing muscles. Biol. Bull. 141: 434–448, 1971.
 165. Elder, H. Y. Peristaltic mechanisms. In: Aspects of Animal Movement, edited by H. Y. Elder and E. R. Trueman, Cambridge: Cambridge Univ. Press, 1980, p. 71–92.
 166. Elder, H. Y., and E. R. Trueman (Eds). Aspects of Animal Movement. Society for Experimental Biology Seminar Series Cambridge: Cambridge Univ. Press, 1980.
 167. Ellington, C. P. The aerodynamics of hovering insect flight. I. The quasi‐steady analysis. Phil. Trans. R. Soc. Lond. [B] 305: 1–15, 1984.
 168. Ellington, C. P. The aerodynamics of hovering insect flight. II. Morphological parameters. Phil. Trans. R. Soc. Lond. [B] 305: 17–40, 1984.
 169. Ellington, C. P. The aerodynamics of hovering insect flight. IV. Aerodynamic mechanisms. Phil. Trans. R. Soc. Lond. [B] 305: 79–113, 1984.
 170. Ellington, C. P. The aerodynamics of hovering insect flight. V. A vortex theory. Phil. Trans. R. Soc. Lond. [B] 305: 115–144, 1984.
 171. Ellington, C. P. The aerodynamics of hovering insect flight. VI. Lift and power requirements. Phil. Trans. R. Soc. Lond. [B] 305: 145–181, 1984.
 172. Ellington, C. P. Limitations on animal flight performance. J. Exp. Biol. 160: 71–91, 1991.
 173. Ellington, C. P., K. E. Machin, and T. M. Casey. Oxygen consumption of bumblebees in forward flight. Nature 347: 472–473, 1990.
 174. Ellington, W. R. The recovery from anaerobic metabolism in invertebrates. J. Exp. Zool. 228: 431–444, 1983.
 175. Ellis, C. H. The mechanism of extension in the legs of spiders. Biol. Bull. 86: 41–50, 1944.
 176. England, W. R., and J. Baldwin. Anaerobic energy metabolism in the tail musculature of the Australian yabby Cherax destructor (Crustacea, Decapoda, Parastacidae): role of phosphagens and anaerobic glycolysis during escape behavior. Physiol. Zool. 56: 614–622, 1983.
 177. Ennos, A. R. The importance of torsion in the design of insect wings. J. Exp. Biol. 140: 137–160, 1988.
 178. Ennos, A. R., and R. J. Wootton. Functional wing morphology and aerodynamics of Panorpa germanica (Insecta: Mecoptera). J. Exp. Biol. 143: 267–284, 1989.
 179. Evans, M.E.G. The jump of the click beetle (Coleoptera: Elateridae)—a preliminary study. J. Zool. 167: 319–336, 1972.
 180. Evans, M.E.G. The jump of the click bettle (Coleoptera: Elateridae)—energetics and mechanics. J. Zool. 169: 181–194, 1973.
 181. Evans, M.E.G. Locomotion in the Coleoptera Adephaga, especially Carabidae. J. Zool. 181: 189–226, 1977.
 182. Evans, M.E.G., and T. G. Forsythe. A comparison of adaptations to running, pushing and burrowing in some adult Coleoptera: especially Carabidae. J. Zool. 202: 513–534, 1984.
 183. Fauci, L. J., and C. S. Peskin. A computational model of aquatic animal locomotion. J. Comput. Physics 77: 85–108, 1988.
 184. Foulds, J. B., and J. C. Roth. Oxygen consumption during simulated vertical migration in Mysis relicta (Crustacea, Mysidacea). Can. J. Zool. 54: 377–385, 1976.
 185. Fourtner, C. R. The ultrastructure of the metathoracic femoral extensors of the cockroach, Periplaneta americana. J. Morphol. 156: 127–140, 1978.
 186. Fourtner, C. R. Role of muscle in insect posture and locomotion. In: Locomotion and Energetics in Arthropods, edited by C. F. Herreid II and C. R. Fourtner. New York: Plenum, 1981, p. 195–213.
 187. Franklin, R., R. Jander, and K. Ele. The coordination mechanics and evolution of swimming by a grasshopper, Melanoplus differentialis (Orthoptera: Acrididae). J. Kansas Entomol. Soc. 50: 189–199, 1977.
 188. Franzini‐Armstrong, C. Natural variability in the length of thin and thick filaments in single fibres from a crab, Portunus depurator. J. Cell Sci. 6: 559–592, 1970.
 189. Full, R. J. Locomotion without lungs: energetics and performance of a lungless salamander, Plethodon jordani. Am. J. Physiol. 251 (Regulatory Integrative Comp. Physiol. 22): R775–R780, 1986.
 190. Full, R. J. Locomotion energetics of the ghost crab: I. Metabolic cost and endurance. J. Exp. Biol. 130: 137–153, 1987.
 191. Full, R. J. Mechanics and energetics of terrestrial locomotion: from bipeds to polypeds. In: Energy Transformation in Cells and Organisms. Proc. 10th Conf. Eur. Soc. Comp. Physiol. Biochem. Stuttgart: Georg Thieme‐Verlag, Insbruck. p. 175–182, 1989.
 192. Full, R. J. The concepts of efficiency and economy in land locomotion. In: Efficiency and Economy in Animal Physiology, edited by R. W. Blake, Cambridge: Cambridge Univ. Press, 1991, p. 97–131.
 193. Full, R. J. Integration of individual leg dynamics with whole body movement in arthropod locomotion. In: Biological Neural Networks in Invertebrate Neuroethology and Robotics, edited by R. Beer, R. Ritzmann, and T. McKenna. New York: Academic, 1992, p. 3–20.
 194. Full, R. J. The importance of mechanical systems in understanding arthropod neural control of locomotion. Proc. Annu. Yale Workshop Adaptive Learning Syst. 8: 21–26, 1994.
 195. Full, R. J., and A. Ahn. Static forces and moments generated in the insect leg: comparison of a three‐dimensional musculoskeletal computer model with experimental measurements. J. Exp. Biol. 198: 1285–1298, 1995.
 196. Full, R. J., B. D. Anderson, C. M. Finnerty, and M. E. Feder. Exercising with and without lungs: I. The effects of metabolic cost, maximal oxygen transport and body size on terrestrial locomotion in salamander species. J. Exp. Biol. 138: 471–485, 1988.
 197. Full, R. J., R. Blickhan, and L. H. Ting. Leg design in hexpedal runners. J. Exp. Biol. 158: 369–390, 1991.
 198. Full, R. J., K. Earls, M. A. Wong, and R. L. Caldwell. Locomotion like a wheel?. Nature 365: 495, 1993.
 199. Full, R. J., and C. F. Herreid II. The aerobic response to exercise of the fastest land crab. Am. J. Physiol. 244 (Regulatory Integrative Comp. Physiol. 15): R530–R536, 1983.
 200. Full, R. J., and C. F. Herreid II. Fiddler crab exercise: energetic cost of running sideways. J. Exp. Biol. 109: 141–161, 1984.
 201. Full, R. J., and C. F. Herreid II. Energetics of multilegged locomotion. Proc. Int. Union Physiol. Sci. 16: 403, 1986.
 202. Full, R. J., C. F. Herreid II, and J. A. Assad. Energetics of the exercising wharf crab, Sesarma cinereum. Physiol. Zool. 58: 605–615, 1985.
 203. Full, R. J., and M.A.R. Koehl. Drag and lift in running insects. J. Exp. Biol. 176: 89–103, 1993.
 204. Full, R. J., R. Kram, and B. Wong. Instantaneous power at the leg joints of running roaches. Am. Zool. 33: 140A, 1993.
 205. Full, R. J., and C. Min. Do insects have a maximal oxygen consumption?. Physiologist 33: A89, 1990.
 206. Full, R. J., and K. Prestwich. Anaerobic metabolism of bouncing gaits in ghost crabs. Am. Zool. 26: 88A, 1986.
 207. Full, R. J., and M. S. Tu. The mechanics of six‐legged runners. J. Exp. Biol. 148: 129–146, 1990.
 208. Full, R. J., and M. S. Tu. Mechanics of a rapid running insect: two‐, four‐ and six‐legged locomotion. J. Exp. Biol. 156: 215–231, 1991.
 209. Full, R. J., and A. Tullis. Capacity for sustained terrestial locomotion in an insect: energetics, thermal dependence and kinematics. J. Comp. Physiol. [B] 160: 573–581, 1990.
 210. Full, R. J., and A. Tullis. Energetics of ascent: insects on inclines. J. Exp. Biol. 149: 307–317, 1990.
 211. Full, R. J., and R. B. Weinstein. Integrating the physiology, mechanics and behavior of rapid running ghost crabs: slow and steady doesn't always win the race. Am. Zool. 32: 382–395, 1992.
 212. Full, R. J., D. A. Zuccarello, and A. Tullis. Effect of variation in form on the cost of terrestrial locomotion. J. Exp. Biol. 150: 233–246, 1990.
 213. Gabriel, J. M. The effect of animal design on jumping performance. J. Zool. 204: 533–539, 1984.
 214. Gabriel, J. M. The development of the locust jumping mechanism I. Allometric growth and its effect on jumping performance. J. Exp. Biol. 118: 313–326, 1985.
 215. Gabrielli, G., and T. Karman. What price speed?. Mech. Eng. 72: 775, 1950.
 216. Gade, G. Effects of oxygen deprivation during anoxia and muscular work on the energy metabolism of the crayfish, Orconectes limosus. Comp. Biochem. Physiol. A 77: 495–502, 1984.
 217. Gans, C. Arthropod locomotion as seen through a vertebrate eye. In: Locomotion and Energetics in Arthropods, edited by C. F. Herreid II and C. R. Fourtner. New York: Plenum, 1981, p. 527–539.
 218. Gans, C., and J. A. Burr. Unique locomotory mechanism of Mermis nigrescens, a large nematode that crawls over soil and climbs through vegetation. J. Morphol. 222: 133–148, 1994.
 219. Gans, C., and F. de Vree. Functional bases of fiber length and angulation in muscle. J. Morphol. 192: 63–85, 1987.
 220. Gatten, R. E., K. Miller, and R. J. Full. Energetics at rest and during locomotion. In: Environmental Physiology of the Amphibians, edited by M. E. Feder and W. W. Burggren, Chicago: Univ. of Chicago Press, 1992, p. 314–377.
 221. Gewecke, M., and G. Wendler (Eds). Insect Locomotion Berlin: Verlag‐Paul Parey, 1985.
 222. Glimour, K. M., and C. P. Ellington. In vivo muscle lengths in bumblebees and the in vitro effects on work and power. J. Exp. Biol. 183: 101–113, 1993.
 223. Gilmour, K. M., and C. P. Ellington. Power output of glycerinated bumblebee flight muscle. J. Exp. Biol. 183: 77–100, 1993.
 224. Gosline, J. M. The elastic properties of rubber‐like and highly extensible tissues. In: The Mechanical Properties of Biological Materials, edited by J. F. V. Vincent and J. D. Currey. London: Soc. Exp. Biol., 1980, p. 331–357.
 225. Gosline, J. M., and M. E. De Mont. Jet‐propelled swimming in squid. Sci. Am. 252: 96–103, 1985.
 226. Gosline, J. M., and R. E. Shadwick. The role of elastic energy storage mechanisms in swimming: an analysis of mantle elasticity in escape jetting in the squid, Loligo opalescens. Can. J. Zool. 61: 1421–1431, 1983.
 227. Gosline, J. M., J. D. Steeves, A. D. Harmon, and De Mont, M. E. Patterns of circular and radial mantle muscle activity in respiration and jetting of the squid Loligo opalescens. J. Exp. Biol. 104: 97–109, 1983.
 228. Graham, D. Unusual step patterns in the free walking grasshopper Neoconocephalus robustus I. General features of the step patterns. J. Exp. Biol. 73: 147–157, 1978.
 229. Graham, D. Insects are both impeded and propelled by their legs during walking. J. Exp. Biol. 104: 129–137, 1983.
 230. Graham, D. Pattern and control of walking in insects. Adv. Insect Physiol. 18: 31–140, 1985.
 231. Granzier, H.L.M., and K. Wang. Interplay between passive tension and strong and weak binding cross‐bridges in insect indirect flight muscle. J. Gen. Physiol. 101: 235–270, 1993.
 232. Gray, J. Animal locomotion. In: London: Weidenfeld & Nicolson, 1968, p. 479.
 233. Gray, J., and G. J. Hancock. The propulsion of sea‐urchin spermatozoa. J. Exp. Biol. 32: 802–814, 1955.
 234. Gray, J., and H. W. Lissmann. The locomotion of nematodes. J. Exp. Biol. 41: 135–154, 1964.
 235. Greenewalt, C. H. Dimensional relationships for flying animals. Smithson. Misc. Coll. 144: 1–46, 1962.
 236. Grieshaber, M. Breakdown and formation of high‐energy phosphates and octopine in the adductor muscle of the scallop, Chlamys opercularis (L.), during escape swimming and recovery. J. Comp. Physiol. [B] 126: 269–276, 1978.
 237. Grodnitsky, D. L., and P. P. Morozov. Vortex formation during tethered flight of functionally and morphologically two‐winged insects, including evolutionary considerations on insect flight. J. Exp. Biol. 182: 11–40, 1993.
 238. Gunzel, D., and W. Rathmayer. Non‐uniformity of sarcomere length can explain the “catch‐like” effect of arthropod muscle. J. Muscle Res. Cell Biol. 15: 535–546, 1994.
 239. Hagopian, M. The myofilament arrangement in the femoral muscle of the cockroach, Leucophaea maderae Fabricius. J. Cell Biol. 28: 545–562, 1966.
 240. Hagopian, M., and D. Spiro. The filament lattice of cockroach thoracic muscle. J. Cell Biol. 36: 433–442, 1968.
 241. Halcrow, K., and C. M. Boyd. The oxygen consumption and swimming activity of the amphipod Gammarus oceanicus at different temperatures. Comp. Biochem. Physiol. 23: 233–242, 1967.
 242. Hanson, J., and J. Lowy. Contractile apparatus in invertebrate animals. In: The structure and function of muscle, edited by G. H. Bourne, New York: Academic, 1960, p. 265–335.
 243. Hardie, J. The tension/length relationship of an insect (Calliphora erythrocephala) supercontracting muscle. Experientia 32: 714–716, 1976.
 244. Hargreaves, B. R. Energetics of crustacean swimming. In: Locomotion and Energetics in Arthropods, edited by C. F. Herreid II and C. R. Fourtner. New York: Plenum, 1981, p. 453–490.
 245. Harris, J., and H. Ghiradella. The forces exerted on the substrate by walking and stationary crickets. J. Exp. Biol. 85: 263–279, 1980.
 246. Harrison, J. F., J. E. Phillips, and T. T. Gleeson. Activity physiology of the two‐striped grasshopper, Melanoplus bivittatus: gas exchange, hemolymph acid‐base status, lactate production, and the effect of temperature. Physiol. Zool. 64: 451–472, 1991.
 247. Heffernan, J. M., and S. A. Wainwright. Locomotion of the holothurian Euapta lappa and redefinition of peristalsis. Biol. Bull. 147: 95–104, 1974.
 248. Heglund, N. C., G. A. Cavagna, and C. R. Taylor. Energetics and mechanics of terrestrial locomotion. III. Energy changes of the centre of mass as a function of speed and body size in birds and mammals. J. Exp. Biol. 97: 41–56, 1982.
 249. Heglund, N. C., and C. R. Taylor. Speed, stride frequency and energy cost per stride: how do they change with body size and gait?. J. Exp. Biol. 138: 301–318, 1988.
 250. Heinrich, B. Thermoregulation in bumblebees. II. Energetics of warm‐up and free flight. J. Comp. Physiol. 96: 155–166, 1975.
 251. Heinrich, B. Heat exchange in relation to blood flow between thorax and abdomen in bumblebees. J. Exp. Biol. 64: 561–585, 1976.
 252. Heinrich, B. Keeping a cool head: honeybee thermoregulation. Science 205: 1269–1271, 1979.
 253. Heinrich, B. Mechanisms of body‐temperature regulation in honeybees, Apis mellifera II. Regulation of thoracic temperature at high air temperatures. J. Exp. Biol. 85: 73–87, 1980.
 254. Heinrich, B. (Ed). Insect Thermoregulation New York: Wiley, 1981.
 255. Heinrich, B. Temperature regulation during locomotion in insects. In: Locomotion and Energetics in Arthropods, edited by C. F. Herreid II and C. R. Fourtner. New York: Plenum, 1981, p. 391–417.
 256. Henry, R. P., C. E. Booth, F. N. Lollier, and P. J. Walsh. Post‐exercise lactate production and metabolism in three species of aquatic and terrestrial decapod crustaceans. J. Exp. Biol. 186: 215–234, 1994.
 257. Herreid II, C. F. Energetics of pedestrian arthropods. In: Locomorion and Energetics in Arthropods, edited by C. F. Herreid II and C. R. Fourtner. New York: Plenum, 1981, p. 491–526.
 258. Herreid II, C. F., and C. R. Fourtner (Eds). Locomotion and Energetics in Arthropods New York: Plenum, 1981.
 259. Herreid II, C. F., and R. J. Full. Cockroaches on a treadmill: aerobic running. J. Insect Physiol. 30: 395–403, 1984.
 260. Herreid II, C. F., and R. J. Full. Energetics of hermit crabs during locomotion: the cost of carrying a shell. J. Exp. Biol. 120: 297–308, 1986.
 261. Herreid II, C. F., and R. J. Full. Locomotion of hermit crabs (Coenobita compressus) on beach and treadmill. J. Exp. Biol. 120: 283–296, 1986.
 262. Herreid II, C. F., and R. J. Full. Energetics and locomotion. In: Biology of the Land Crab, edited by W. Burggren and B. R. McMahon, New York: Cambridge Univ. Press, 1988, p. 333–377.
 263. Herreid II, C. F., R. J. Full, and D. A. Prawel. Energetics of cockroach locomotion. J. Exp. Biol. 94: 189–202, 1981.
 264. Herreid II, C. F., L. W. Lee, and G. M. Shah. Respiration and heart rate in exercising land crabs. Respir. Physiol. 36: 109–120, 1979.
 265. Herreid II, C. F., P. J. O'Mahoney, and R. J. Full. Locomotion in land crabs: respiratory and cardiac response of Gecarcinus lateralis. Comp. Biochem. Physiol. A 74: 117–124, 1983.
 266. Herreid II, C. F., D. A. Prawel, and R. J. Full. Energetics of running cockroaches. Science 212: 331–333, 1981.
 267. Herreid II, C. F., D. A. Sperrazza, R. Weinstein, and R. J. Full. Energetics of locomotion of crawling caterpillars. Physiologist 28: 277, 1985.
 268. Hertel, H. (Ed). Structure, Form, Movements New York: Reinhold, 1966.
 269. Hill, A. V. The heat of shortening and the dynamic constants of muscle. Proc. R. Soc. Lond. [B] 126: 136–195, 1938.
 270. Hill, A. V. The dimensions of animals and their muscular dynamics. Sci. Prog. 38: 209–230, 1950.
 271. Hill, A. V. The mechanics of active muscle. Proc. R. Soc. Lond. [B] 141: 104–117, 1953.
 272. Hitzemann, K. Untersuchungen uber den Energie‐Stoffwechsel in der Sprungmusculatur von Locusta migratoria (L.). Munster: Univ. of Munster, 1979. Dissertation.
 273. Hocking, B. The intrinsic range and speed of flight of insects. Trans. R. Entomol. Soc. Lond. 104: 223–345, 1953.
 274. Holberton, D. V. Locomotion of protozoa and single cells. In: Mechanics and Energetics of Animal Locomotion, edited by R. M. Alexander and G. Goldspink. London: Chapman and Hall, 1977, p. 279–332.
 275. Holwill, M. E. J. Low Reynolds number undulatory propulsion in organisms of different sizes. In: Scale Effects in Animal Locomotion, edited by T. J. Pedley, London: Academic, 1977, p. 233–283.
 276. Houlihan, D. F., C. K. Govind, and A. El Haj. Energetics of swimming in Callinectes sapidus and walking in Homarus americanus. Comp. Biochem. Physiol. A 82: 267–279, 1985.
 277. Houlihan, D. F., and A. J. Innes. Oxygen consumption, crawling speeds, and cost of transport in four Mediterranean intertidal gastropods. J. Comp. Physiol. [B] 147: 113–121, 1982.
 278. Houlihan, D. F., and A. J. Innes. The cost of walking in crabs: aerial and aquatic oxygen consumption during activity of two species of intertidal crab. Comp. Biochem. Physiol. A 77: 325–334, 1984.
 279. Houlihan, D. F., and E. Mathers. Effects of captivity and exercise on the energetics of locomotion and muscle of Carcinus maenus (L.). J. Exp. Mar. Biol. Ecol. 92: 125–142, 1985.
 280. Houlihan, D. F., E. Mathers, and A. J. El Haj. Walking performance and aerobic and anaerobic metabolism of Carcinus maenas (L.) in sea water at 15°C. J. Exp. Mar. Biol. Ecol. 74: 211–230, 1984.
 281. Hoy, M. G., F. E. Zajac, and M. E. Gordon. A musculoskeletal model of the human lower extremity: the effect of muscle, tendon, and moment arm on the moment‐angle relationship of musculotendon actuators at the hip, knee, and ankle. J. Biomech. 23: 157–169, 1990.
 282. Hoyle, G. Neuromuscular mechanisms of a locust skeletal muscle. Proc. R. Soc. Lond. [B] 143: 343–367, 1955.
 283. Hoyle, G. Specificity of muscle. In: Invertebrate Nervous Systems, edited by C.A.G. Wiersma. Chicago: Univ. of Chicago Press, 1967, p. 151–167.
 284. Hoyle, G. Arthropod walking. In: Neural Control of Locomotion, edited by R. M. Herman, S. Grillner, P.S.G. Stein, and D. G. Stuart. New York: Plenum, 1976, p. 137–179.
 285. Hoyle, G. Distributions of nerve and muscle fibre types in locust jumping muscle. J. Exp. Biol. 73: 205–233, 1978.
 286. Hoyle, G. Muscles and their Neural Control New York: Wiley, 1983.
 287. Hoyle, G., and P. A. McNeill. Correlated physiological and ultrastructural studies on specialised muscles. Ib. Ultrastructure of white and pink fibers of the levator of the eyestalk of Podopbthalmus vigil (Weber). J. Exp. Zool. 167: 487–522, 1968.
 288. Hughes, G. M. The co‐ordination of insect movements. I. The walking movements of insects. J. Exp. Biol. 29: 267–283, 1951.
 289. Hughes, G. M., and P. J. Mill. Locomotion:terrestrial. In: The Physiology of Insecta, edited by M. Rockstein. New York: Academic, 1974, p. 335–379.
 290. Hui, C. A. Walking of the shore crab Pachygrapsus crassipes in its two natural environments. J. Exp. Biol. 165: 213–227, 1992.
 291. Hunter, R. D., and H. Y. Elder. Burrowing dynamics and energy cost of transport in the soft‐bodied marine invertebrates Polyphysia crassa and Priapulus caudatus. J. Zool. 218: 209–222, 1989.
 292. Innes, A. J., and D. F. Houlihan. Aerobic capacity and cost of locomotion of a cool temperate gastropod: a comparison with some Mediterranean species. Comp. Biochem. Physiol. A 80: 487–493, 1985.
 293. Ivlev, V. S. Energy consumption during the motion of shrimps. Zool. Zh, 42: 1465–1471, 1963.
 294. Jacklyn, P. M., and D. A. Ritz. Hydrodynamics of swimming in scyllarid lobsters. J. Exp. Mar Biol. Ecol. 101: 85–99, 1986.
 295. Jahromi, S. S., and H. L. Atwood. Correlation of structure, speed of contraction, and total tension in fast and slow abdominal muscle fibers of the lobster (Homarus americanus). J. Exp. Zool. 171: 25–38, 1969.
 296. Jahromi, S. S., and H. L. Atwood. Structural features of muscle fibres in the cockroach leg. J. Insect. Physiol. 15: 2255–2262, 1969.
 297. Jahromi, S. S., and H. L. Atwood. Structural and contractile properties of lobster leg‐muscle fibers. J. Exp. Zool. 176: 475–486, 1971.
 298. Jamon, M., and F. Clarac. Locomotion patterns on freely moving crayfish (Procambarus clarkii). J. Exp. Biol. 198: 683–700, 1995.
 299. Jensen, M. Biology and physics of locust flight III. The aerodynamics of locust flight. Phil. Trans. R. Soc. Lond. [B] 239: 511–552, 1956.
 300. Jensen, M., and T. Weis‐Fogh. Biology and physics of locust flight V. Strength and elasticity of locust cuticle. Phil. Trans. R. Soc. Long. [B] 245: 137–169, 1962.
 301. Jensen, T. F., and I. Holm‐Jensen. Energetic cost of running in workers of three ant species Formica fusca L., Formica rufa L., and Camponotus herculaneaus L. (Hymenoptera, Formicidae). J. Comp. Physiol. [B] 137: 151–156, 1980.
 302. John‐Alder, H. B., and A. F. Bennett. Thermal dependence of endurance and locomotory energetics in a lizard. Am. J. Physiol. 241 (Regulatory Integrative Comp. Physiol. 12): R342–R349, 1981.
 303. Johnson, W., P. D. Soden, and E. R. Trueman. A study in jet propulsion: an analysis of the motion of the squid, Loligo vulgaris. J. Exp. Biol. 56: 155–165, 1972.
 304. Johnston, I. A., J. D. Fleming, and T. Crockford. Thermal acclimation and muscle contractile properties in cyprinid fish. Am. J. Physiol. 259 (Regulatory Integrative Comp. Physiol. 30): R231–R236, 1990.
 305. Jones, H. D. Locomotion. In: Pulmonates, edited by V. Fretter and J. Peake. London: Academic, 1975.
 306. Joos, B., P. A. Young, and T. M. Casey. Wingstroke frequency of foraging and hovering bumblebees in relation to morphology and temperature. Physiol. Entomol. 16: 191–200, 1991.
 307. Jordan, C. E. A model of rapid‐start swimming at intermediate Reynolds number: undulatory locomotion in the chaetognath Sagitta elegans. J. Exp. Biol. 163: 119–137, 1992.
 308. Josephson, R. K. Extensive and intensive factors determining the performance of striated muscle. J. Exp. Zool. 194: 135–154, 1975.
 309. Josephson, R. K. Temperature and the mechanical performance of insect muscle. In: Insect Thermoregulation, edited by B. Heinrich. New York: Wiley, 1981, p. 9–44.
 310. Josephson, R. K. Contraction dynamics of flight and stridulatory muscles of tettigoniid insects. J. Exp. Biol. 108: 77–96, 1984.
 311. Josephson, R. K. Mechanical power output from striated muscle during cyclic contraction. J. Exp. Biol. 114: 493–512, 1985.
 312. Josephson, R. K. The mechanical power output of a tettigoniid wing muscle during singing and flight. J. Exp. Biol. 117: 357–368, 1985.
 313. Josephson, R. K. A synchronous insect muscle with an operating frequency greater than 500 Hz. J. Exp. Biol. 118: 185–208, 1985.
 314. Josephson, R. K. Power output from skeletal muscle during linear and sinusoidal shortening. J. Exp. Biol. 147: 533–537, 1989.
 315. Josephson, R. K. Contraction dynamics and power output of skeletal muscle. Annu. Rev. Physiol. 55: 527–546, 1993.
 316. Josephson, R. K., and R. D. Stevenson. The efficiency of a flight muscle from the locust Schistocerca americana. J. Physiol. (Lond.) 413–429, 1991.
 317. Josephson, R. K., and D. R. Stokes. Strain, muscle length and work output in a crab muscle. J. Exp. Biol. 145: 45–61, 1989.
 318. Kammer, A. E., and B. Heinrich. Insect flight metabolism. Adv. Insect Physiol. 14: 133–228, 1978.
 319. Katz, S. L., and J. M. Gosline. Ontogenetic scaling and mechanical behaviour of the tibiae of the African desert locust (Schistocerca gregaria). J. Exp. Biol. 168: 125–150, 1992.
 320. Katz, S. L., and J. M. Gosline. Ontogenetic scaling of jump performance in African desert locust (Schistocerca gregaria). J. Exp. Biol. 177: 81–111, 1993.
 321. Kawaguti, S., and Y. Kamichima. Electron microscopy on the long‐sarcomere muscle of the spider leg. Biol. J. Okayama Univ. 15: 73–86, 1969.
 322. Keller, J. B., and M. S. Falkovitz. Crawling of worms. J. Theor. Biol. 104: 417–442, 1983.
 323. Kelly, R. E., and R. V. Rice. Abductin: a rubber‐like protein from the internal triangular hinge ligament of pecten. Science 155: 208–210, 1967.
 324. Kier, W. M. The musculature of squid arms and tentacles: ultrastructural evidence for functional differences. J. Morphol. 185: 223–239, 1985.
 325. Kier, W. M. The fin musculature of cuttlefish and squid (Mollusca, Cephalopoda): morphology and mechanics. J. Zool. 217: 23–38, 1989.
 326. Kier, W. M., and K. K. Smith. Tongues, tentacles, and trunks: the biomechanics of movement in muscular hydrostats. Zool. J. Linn. Soc. 83: 307–324, 1985.
 327. Klarner, D. B., D. Klarner, and W. J. P. Barnes. The cuticular stress detector (CSD2) of the crayfish. II. Activity during walking and influences on leg coordination. J. Exp. Biol. 122: 161–175, 1986.
 328. Klyashtorin, L. B., and V. I. Kuz'micheva. Level of energy expenditure of planktonic crustaceans on active movements. Oceanology 15: 592–595, 1975.
 329. Koehl, M.A.R., and J. R. Strickler. Copepod feeding currents: food capture at low Reynolds number. Limnol. Oceanogr. 26: 1062–1073, 1981.
 330. Kohlhage, K. The economy of paddle‐swimming: the role of added waters and viscosity in the locomotion of Daphnia magna. Zool. Better. 35: 47–54, 1994.
 331. Kozacik, J. J. Stepping patterns in the cockroach, Periplaneta americana. J. Exp. Biol. 90: 357–360, 1981.
 332. Krebs, H. A. The August Krogh principle: “For many problems there is an animal on which it can be most conveniently studied.” J. Exp. Zool. 194: 221–226, 1975.
 333. Krogh, A., and T. Weis‐Fogh. The respiratory exchange of the desert locust (Shistocerca gregaria) before, during and after flight. J. Exp. Biol. 28: 344–357, 1951.
 334. Kutsch, W., and P. Stevenson. Time‐correlated flights of juvenile and mature locusts: a comparison between free and tethered animals. J. Insect Physiol. 27: 455–459, 1981.
 335. La Barbera, M. Why the wheels won't go. Am. Nat. 121: 395–408, 1983.
 336. Lanzavecchia, G., M. de Eguileor, and R. Valvassori. Muscles. In: The Ultrastructure of Polychaeta, edited by W. Westheide and C. O. Hermans, Stuttgart: Gustav Fischer‐Verlag, 1988, p. 71–88.
 337. Lanzavecchia, G., M. De Eguileor, R. Valvassori, and P. Lanzavecchia, Jr. Analysis and reconstruction of unusual obliquely striated fibres in lumbriculids (Annelida, Oligochaeta). J. Muscle Res. Cell Motil. 8: 209–219, 1987.
 338. Larson, R. J. Costs of transport for the scyphomedusa Stomolophus meleagris L. Agassiz. Can. J. Zool. 65: 2690–2695, 1987.
 339. Lieber, R. L., M. E. Leonard, C. L. Brown, and C. L. Trestik. Frog semitendinosis tendon load‐strain and stress‐strain properties during passive loading. Am. J. Physiol. 261 (Cell Physiol. 30): C86–C92, 1991.
 340. Lighthill, J., and R. Blake. Biofluid dynamics of balistiform and gymnotiform locomotion. Part 1. Biological background, and analysis by elongated‐body theory. J. Fluid Mech. 212: 183–207, 1990.
 341. Lighthill, M. J. Large‐amplitude elongated‐body theory of fish locomotion. Proc. R. Soc. Lond. [B] 179: 125–138, 1971.
 342. Lighton, J. B., J. A. Weier, and D.H.J. Feener. The energetics of locomotion and load carriage in the desert harvester ant Pogonomyrmex rugosus. J. Exp. Biol. 183: 49–61, 1993.
 343. Lighton, J.R.B. Minimum cost of transport and ventilatory patterns in three African beetles. Physiol. Zool. 58: 390–399, 1985.
 344. Lighton, J. R. B., and G. A. Bartholomew. Standard energy metabolism of a desert harvester ant, Pogonomyrmex regosus: effects of temperature, body mass, group size, and humidity. Proc. Natl. Acad. Sci. U.S.A. 85: 4665–4769, 1988.
 345. Lighton, J.R.B., G. A. Bartholomew, and D. H. Feener. Energetics of locomotion and load carriage and a model of the energy cost of foraging in the leaf‐cutting ant Atta colombica Guer. Physiol. Zool. 60: 524–537, 1987.
 346. Lighton, J.R.B., and D. H. Feener. A comparison of energetics and ventilation of desert ants during voluntary and forced locomotion. Nature 342: 174–175, 1989.
 347. Lighton, J.R.B., and R. G. Gillespie. The energetics of mimicry: the cost of pedestrian transport in a formicine ant and its mimic, a clubionid spider. Physiol. Entomol. 14: 173–177, 1989.
 348. Long, G. L. The stereospecific distribution and evolutionary significance of invertebrate lactate dehydrogenases. Comp. Biochem. Physiol. [B] 55: 77–83, 1976.
 349. Machin, K. E., and J.W.S. Pringle. The physiology of insect fibrillar muscle. II. Mechanical properties of a beetle flight muscle. Proc. R. Soc. Lond. [B] 151: 204–255, 1959.
 350. Mackie, G. O., and Q. Bone. Locomotion and propagated skin impulses in salps (Tunicata: Thaliacea). Biol. Bull. 153: 180–197, 1977.
 351. Madin, L. P. Aspects of jet propulsion in salps. Can. J. Zool. 68: 765–777, 1990.
 352. Maitland, D. P. Locomotion by jumping in the Mediterranean fruit‐fly larva Ceratitis capitata. Nature 355: 159–161, 1992.
 353. Malamud, J. G. The tension in a locust flight muscle at varied muscle lengths. J. Exp. Biol. 144: 479–494, 1989.
 354. Malamud, J. G., and R. K. Josephson. Force‐velocity relationships of a locust flight muscle at different times during a twitch contraction. J. Exp. Biol. 159: 65–87, 1991.
 355. Manton, S. M. The evolution of arthropodan locomotory mechanisms. Part I. The locomotion of Peripatus. J. Linn. Soc. Zool. 41: 529–570, 1950.
 356. Manton, S. M. Locomotory habits and the evolution of the larger arthropodan groups. Symp. Soc. Exp. Biol. 7: 339–353, 1953.
 357. Manton, S. M. (Ed). The Arthropoda: Habits, Functional Morphology, and Evolution Oxford: Clarendon, 1977.
 358. Manton, S. M., and J. P. Harding. The evolution of arthropodan locomotory mechanisms. Part 10. Locomotory habits, morphology and evolution of the hexapod classes. J. Linn. Soc. Zool. 51: 203–400, 1972.
 359. Marden, J. H. Maximum lift production during takeoff in flying animals. J. Exp. Biol. 130: 235–258, 1987.
 360. Marsh, R. L. Deactivation rate and shortening velocity as determinants of contractile frequency. Am. J. Physiol. 259 (Regulatory Integrative Comp. Physiol. 30): R223–R230, 1990.
 361. Marsh, R. L., and J. M. Olson. Power output of scallop adductor muscle during contractions replicating the in vivo mechanical cycle. J. Exp. Biol. 193: 139–156, 1994.
 362. Marsh, R. L., J. M. Olson, and S. K. Guzik. Mechanical performance of scallop adductor muscle during swimming. Nature 357: 411–413, 1992.
 363. Martinez, M., R. J. Full, and M. A. R. Koehl. Kinematics of intertidal crabs: locomoting through water and air. Physiol. Zool. 68: 58, 1995.
 364. Matsumoto, G. I. Swimming movements of ctenophores, and the mechanics of propulsion by ctene rows. Hydrobiology 216/217: 319–325, 1991.
 365. Matsuno, A., and Y. Kawamura. Cell types differing in thick myofilament diameter in obliquely striated muscle of a polychaete annelid, Neanthes. Tissue Cell 23: 481–487, 1991.
 366. May, M. L. Wingstroke frequency of dragonflies (Odonata: Anisoptera) in relation of temperature and body size. J. Comp. Physiol. [B] 144: 229–240, 1981.
 367. May, M. L. Dragonfly flight: power requirements at high speed and acceleration. J. Exp. Biol. 158: 325–342, 1991.
 368. May, M. L., D. L. Pearson, and T. M. Casey. Oxygen consumption of active and inactive adult tiger beetles. Physiol. Entomol. 11: 171–179, 1986.
 369. McFarland, W. N., and P. E. Pickens. The effects of season, temperature, and salinity on standard and active oxygen consumption of the grass shrimp, Palaemonetes vulgaris (Say). Can. J. Zool. 43: 571–585, 1965.
 370. McGhee, R. B. Some finite state aspects of legged locomotion. Math. Biosci. 2: 67–84, 1968.
 371. McMahon, B. R., D. G. McDonald, and C. M. Wood. Ventilation, oxygen uptake and haemolymph oxygen transport, following enforced exhausting activity in the dungeness crab Cancer magister. J. Exp. Biol. 80: 271–285, 1979.
 372. McMahon, T. A., and G. C. Cheng. The mechanics of running: how does stiffness couple with speed?. J. Biomech. 23: 65–78, 1990.
 373. McMahon, T. A., G. Valiant, and E. C. Frederick. Groucho running. J. Appl. Physiol.: Respir. Environ. Exerc. Physiol. 62: 2326–2337, 1987.
 374. McQueen, D. J. Active respiration rates for the burrowing wolf spider Geolycosa domifex (Hancock). Can. J. Zool. 58: 1066–1074, 1980.
 375. Meyhofer, E., and T. Daniel. Dynamic mechanical properties of extensor muscle cells of the shrimp Pandalus danae: cell design for escape locomotion. J. Exp. Biol. 151: 435–452, 1990.
 376. Mill, P. J., and M. F. Knapp. The fine structure of obliquely striated body wall muscles in the earthworm, Lumbricus ter‐restris Linn. J. Cell Sci. 7: 233–261, 1970.
 377. Mill, P. J., and R. S. Pickard. Jet‐propulsion in anisopteran dragonfly larvae. J. Comp. Physiol. 97: 329–338, 1975.
 378. Miller, J. B. The length‐tension relationship of the dorsal longitudinal muscle of a leech. J. Exp. Biol. 62: 43–53, 1975.
 379. Millman, B. M., and P. M. Bennett. Structure of the cross‐striated adductor muscle of the scallop. J. Mol. Biol. 103: 439–467, 1976.
 380. Mizisin, A. P., and R. K. Josephson. Mechanical power output of locust flight muscle. J. Comp. Physiol. [A] 160: 413–419, 1987.
 381. Moore, J. D., and E. R. Trueman. Swimming of the scallop, Chlamys opercularis (L.). J. Exp. Mar. Biol. Ecol. 6: 179–185, 1971.
 382. Morgan, C. R., and D. R. Stokes. Ultrastructural heterogeneity of the mesocoxal muscles of Periplaneta americana. Cell Tissue Res. 201: 305–314, 1979.
 383. Morgan, K. R. Temperature regulation, energy metabolism and mate‐searching in rain beetles (Pleocotna spp.), winter‐active, endothermic scarabs (Coleoptera). J. Exp. Biol. 128: 107–122, 1987.
 384. Morgan, K. R., T. E. Shelly, and L. S. Kimsey. Body temperature regulation, energy metabolism, and foraging in light‐seeking and shade‐seeking robber flies. J. Comp. Physiol. [B] 155: 561–570, 1985.
 385. Morris, M. J., G. Gust, and J. J. Torres. Propulsion efficiency and cost of transport for copepods: a hydromechanical model of crustacean swimming. Mar. Biol. 86: 283–295, 1985.
 386. Morris, M. J., K. Kohlhage, and G. Gust. Mechanics and energetics of swimming in the small copepod Acanthocyclops robustus (Cyclopoida). Mar. Biol. 107: 83–91, 1990.
 387. Nachtigall, W. Mechanics of swimming in water‐beetles. In: Aspects of Animal Movement, edited by H. Y. Elder and E. R. Trueman, Cambridge: Cambridge Univ. Press, 1980, p. 107–124.
 388. Nachtigall, W. The biophysics of locomotion on land. In: Biophysics, edited by Hoppe W., Lohmann, W., Markel, H., and Zielger, H. Berlin: Springer‐Verlag, 1983, p. 580–587.
 389. Nachtigall, W. Swimming in aquatic insects. In: Comprehensive Insect Physiology, Biochemistry and Pharmacology. Nervous System: Structure and Motor Function, edited by G. A. Kerkut and L. I. Gilbert, New York: Pergamon, 1985, p. 467–490.
 390. Nachtigall, W. Mechanics and aerodynamics of flight. In: Insect Flight, edited by G. Goldsworthy and C. Wheeler. Boca Raton, FL: CRC, 1989, p. 1–29.
 391. Nachtigall, W., and U. Hanauer‐Thieser. Drag and lift coefficients of the bee's body; implications for flight dynamics. J. Comp. Physiol. [B] 162: 267–277, 1992.
 392. Nicolson, S. W., G. A. Bartholomew, and M. K. Seely. Ecological correlates of locomotion speed, morphometrics and body temperature in three Namib Desert tenebrionid beetles. S. Afr. J. Zool. 19: 131–134, 1984.
 393. Nicolson, S. W., and G. N. Louw. Simultaneous measurement of evaporative water loss, oxygen consumption, and thoracic temperature during flight in a carpenter bee. J. Exp. Zool. 222: 287–296, 1982.
 394. Niebur, E., and P. Erdos. Theory of the locomotion of nematodes—dynamics of undulatory progression on a surface. Biophys J. 60: 1132–1146, 1991.
 395. O'Dor, R. K. Respiratory metabolism and swimming performance of the squid, Loligo opalescens, Can. J. Fish. Aquat. Sci. 39: 580–587, 1982.
 396. O'Dor, R. K. The forces acting on swimming squid. J. Exp. Biol. 137: 421–442, 1988.
 397. O'Dor, R. K. Limitations on locomotor performance in squid. J. Appl. Physiol.: Respir. Environ. Exerc. Physiol. 64: 128–134, 1988.
 398. O'Dor, R. K., and D. M. Webber. Invertebrate athletes: tradeoffs between transport efficiency and power density in cephalopod evolution. J. Exp. Biol. 160: 93–112, 1991.
 399. O'Dor, R. K., J. Wells, and M. J. Wells. Speed, jet pressure and oxygen consumption relationships in free‐swimming Nautilus. J. Exp. Biol. 154: 383–396, 1990.
 400. Onnen, T., and E. Zebe. Energy metabolism in the tail muscles of the shrimp Crangon crangon during work and subsequent recovery. Comp. Biochem. Physiol. A 74: 833–838, 1983.
 401. Osborne, M. P. Supercontraction in the muscles of the blowfly larva: an ultrastructural study. J. Insect Physiol. 13: 1471–1482, 1967.
 402. Otten, E. Optimal design of vertebrate and insect sarcomeres. J. Morphol. 191: 49–62, 1987.
 403. Pabst, D. A., and M. D. Healy. Worms and whale tails: flattening the circular cylinder. Am. Zool. 29: 182A, 1989.
 404. Parry, D. A., and R.H.J. Brown. The hydraulic mechanism of the spider leg. J. Exp. Biol. 36: 423–433, 1959.
 405. Pasquali‐Ronchetti, I. The ultrastructural organization of femoral muscles in Musca domestica (Diptera). Tissue Cell 2: 339–354, 1970.
 406. Pedley, T. J. (Ed). Scale Effects in Animal Locomotion London: Academic, 1977.
 407. Pennycuick, C. J. Adapting skeletal muscle to be efficient. In: Efficiency and Economy in Animal Physiology, edited by R. W. Blake, Cambridge: Cambridge Univ. Press, 1991, p. 33–42.
 408. Pennycuick, C. J., and M. A. Rezende. The specific power output of aerobic muscle, related to the power density of mitochondria. J. Exp. Biol. 108: 377–392, 1984.
 409. Plotnik, R. E. Lift based mechanisms for swimming in eurypterids and portunid crabs. Trans. R. Soc. Edinburgh 76: 325–337, 1985.
 410. Pond, C. M. The role of the “walking legs” in aquatic and terrestrial locomotion of the crayfish Austropotamobius pallipes (Lereboullet). J. Exp. Biol. 62: 447–454, 1975.
 411. Prestwich, K. N. Anaerobic metabolism in spiders. Physiol. Zool. 56: 112–121, 1983.
 412. Prestwich, K. N. The roles of aerobic and anaerobic metabolism in active spiders. Physiol. Zool. 56: 122–132, 1983.
 413. Prestwich, K. N. The constraints on maximal activity in spiders: I. Evidence against the fluid insufficiency hypothesis. J. Comp. Physiol. [B] 158: 437–447, 1988.
 414. Prestwich, K. N. The constraints on maximal activity in spiders: II. Limitations imposed by phosphagen depletion and anaerobic metabolism. J. Comp. Physiol. [B] 158: 449–456, 1988.
 415. Pringle, J.W.S. Arthropod muscle. In: The Structure and Function of Muscle, edited by G. H. Bourne, New York: Academic, 1972, p. 491–541.
 416. Prosser, C. L. (Ed). Comparative Animal Physiology (3rd ed.). Philadelphia: Saunders, 1973.
 417. Queathem, E. The ontogeny of grasshopper jumping performance. J. Insect Physiol. 37: 129–138, 1991.
 418. Queathem, L., and R. J. Full. Variation in jump force production within an instar of the grasshopper Schistocerca americana. J. Zool. 235: 605–620, 1995.
 419. Quinn, R. D., and K. S. Espenschied. Control of a hexapod robot using a biologically inspired neural network. In: Biological Neural Networks in Invertebrate Neuroethology and Robotics, edited by R. Beer, R. Ritzmann, and T. McKenna. New York: Academic, 1992, p. 365–381.
 420. Raibert, M. H. Trotting, pacing and bounding by a quadruped robot. J. Biomech. 23 (suppl. 1): 79–98, 1990.
 421. Raibert, M. H., M. Chepponis, and H. B. Brown. Running on four legs as though they were one. IEEE J. Rob. Auto. RA‐2: 70, 1986.
 422. Rheuben, M. B., and A. E. Kammer. Comparison of slow larval and fast adult muscle innervated by the same motor neurone. J. Exp. Biol. 84: 103–118, 1980.
 423. Rome, L. C., R. P. Funke, R. M. Alexander, O. Lutz, H. Aldridge, and F. Scott. Why animals have different muscle fibre types. Nature 335: 824–827, 1988.
 424. Rome, L. C., P. T. Loughna, and G. Goldspink. Muscle fiber activity in carp as a function of swimming speed and muscle terperature. Am. J. Physiol. Regulatory Intergrative Comp. Physiol. 247: R272–R279, 1983.
 425. Root, T. M., and R. F. Bowerman. The scorpion walking leg motor system: muscle fine structure. Comp. Biochem. Physiol. A 69: 73–78, 1981.
 426. Rosenbluth, J. Ultrastructural organization of obliquely striated muscle fibers in Ascaris lumbricoides. Comp. Biochem. Physiol. A 25: 495–515, 1965.
 427. Rothe, U., and W. Nachtigall. Flight of the honey bee. V. Respiration quotients and metabolic rates during sitting, walking and flying. J. Comp. Physiol. [B] 158: 739–749, 1989.
 428. Rutledge, P. S., and A. W. Pritchard. Scope for activity in the crayfish Pacifastacus leniusculus. Am. J. Physiol. 240: 87–92, 1981.
 429. Satterlie, R. A., M. La Barbera, and A. N. Spencer. Swimming in the pteropod mollusc, Clione limacina. I. Behaviour and morphology. J. Exp. Biol. 116: 189–204, 1985.
 430. Schmidt‐Nielsen, K. Locomotion: energetic cost of swimming, flying and running. Science 177: 222–228, 1972.
 431. Schmidt‐Nielsen, K. Animal Physiology: Adaptation and Environment (4th ed.). Cambridge: Cambridge Univ. Press, 1990.
 432. Schottler, U. On the anaerobic metabolism of three species of Nereis (Annelida). Mar. Ecol. Prog. Ser. 1: 249–254, 1979.
 433. Seymour, R. S., and A. Vinegar. Thermal relations, water loss and oxygen consumption of a North American tarantula. Comp. Biochem. Physiol. A 44: 83–96, 1973.
 434. Shafiq, S. A. Electron microscopic studies on the indirect flight muscles of Drosophila melanogaster. II. Differentiation of myofibrils. J. Cell Biol. 17: 363–373, 1963.
 435. Shapely, H. Thermokinetics of Liometopum apiculatum Mayr. Proc. Natl. Acad. Sci. U.S.A. 6: 204–211, 1920.
 436. Sherman, R. G., and A. R. Luff. Structural features of the tarsal claw muscles of the spider Eurypelma marxi Simon. Can. J. Zool. 49: 1549–1556, 1971.
 437. Shultz, J. W. Evolution of locomotion in Arachnida: the hydraulic pressure pump of the giant whipscorpion, Mastigoproctus giganteus (Uropygi). J. Morphol. 210: 13–31, 1991.
 438. Siegmund, B., and M. K. Grieshaber. Opine metabolism of Arenicola marina L. Program, First Int. Congr. C. P. B., 1984.
 439. Singla, C. L. Locomotion and neuromuscular system of Aglantha digitate. Cell Tiss. Res. 188: 317–327, 1978.
 440. Sleigh, M. A., and J. R. Blake. Methods of ciliary propulsion and their size limitations. In: Scale Effects in Animal Locomotion, edited by T. J. Pedley, London: Academic, 1977, p. 243–256.
 441. Sleinis, S., and G. E. Silvey. Locomotion in a forward walking crab. J. Comp. Physiol. [A] 136: 301–312, 1980.
 442. Smatresk, N. J., A. J. Preslar, and J. N. Cameron. Post‐exercise acid‐base disturbance in Gecarcinus lateralis, a terrestrial crab. J. Exp. Zool. 210: 205–210, 1979.
 443. Smith, B. P., and D. Barr. Swimming by the water mite Limnochares americana Lundblad (Acari, Parasitengona, Limnocharidae). Can. J. Zool. 55: 2050–2059, 1977.
 444. Smith, D. S. The structure of intersegmental muscle fibers in an insect, Periplaneta americana. J. Cell Biol. 29: 449–459, 1966.
 445. Song, S., and K. Waldron (Eds). Machines that Walk: the Adaptive Suspension Vehicle Cambridge, MA: MIT, 1989.
 446. Song, S. M. Machines that Walk: the Adaptive Suspension Vehicle Columbus: Ohio State Univ. 1984.
 447. Sotavalta, O. The essential factor regulating the wing‐stroke frequency of insects in wing mutilation and loading experiments and in experiments at subatmospheric pressure. Ann. Zool. Soc. Zool.‐Bot. Fenn. Vanamo 15: 1–67, 1952.
 448. Sotavalta, O. Recordings of high wing‐stroke and thoracic vibration frequency in some midges. Biol. Bull. 104: 439–444, 1953.
 449. Spedding, G. R. The aerodynamics of flight. In: Mechanics of Animal Locomotion, edited by R. M. Alexander, Berlin: Springer‐Verlag, 1992, p. 52–111.
 450. Squire, J. M. Molecular Mechanisms in Muscular Contraction. Topics in Molecular and Structural Biology, edited by S. Neidle and W. Fuller. Boca Raton, FL: CRC, 1990.
 451. Stern‐Tomlinson, W., M. P. Nusbaum, L. E. Perez, and W. B. Kristan, Jr. A kinematic study of crawling behavior in the leech, Hirudo medicinalis. J. Comp. Physiol. [A] 158: 593–603, 1986.
 452. Stevens, E. D. Relation between work and power calculated from the force‐velocity curves to that done during oscillatory work. J. Muscle Res. Cell Motil. 14: 518–526, 1993.
 453. Stevenson, R. D., and R. K. Josephson. Effects of operating frequency and temperature on mechanical power output from moth flight muscle. J. Exp. Biol. 149: 61–78, 1990.
 454. Stokes, D. R. Insect muscles innervated by single motoneurons: structural and biochemical features. Am. Zool. 27: 1001–1010, 1987.
 455. Stokes, D. R., and R. K. Josephson. The mechanical power output of a crab respiratory muscle. J. Exp. Biol. 140: 287–299, 1988.
 456. Storey, K. B., and J. M. Storey. Energy metabolism in the mantle muscle of the squid, Loligo pealeii. J. Comp. Physiol. [B] 123: 169–175, 1978.
 457. Storey, K. B., and J. M. Storey. Octopine metabolism in the cuttlefish, Sepia officinalis: octopine production by muscle and its role as an aerobic substrate for non‐muscular tissues. J. Comp. Physiol. [B] 131: 311–319, 1979.
 458. Straub, R., and M. Heisenberg. Coordination of legs during straight walking and turning in Drosophila melagaster. J. Comp. Physiol. [A] 167: 403–412, 1990.
 459. Sunada, S., K. Kawacchi, I. Watanabe, and A. Azuma. Fundamental analysis of three‐dimentional “near flying.” J. Exp. Biol. 183: 217–248, 1993.
 460. Sunada, S., K. Kawacchi, I. Watanabe, and A. Azuma. Performance of a butterfly in take‐off flight. J. Exp. Biol. 183: 249–277, 1993.
 461. Tameyasu, T., and H. Sugi. The origin of the series elastic component in single crayfish muscle fibers. Experientia 35: 210–211, 1979.
 462. Tashiro, N. Mechanical properties of the longitudinal and circular muscle in the earthworm. J. Exp. Biol. 54: 101–110, 1971.
 463. Taylor, C. R., K. Schmidt‐Nielsen, and J. L. Raab. Scaling of energetic cost to body size in mammals. Am. J. Physiol. 210: 1104–1107, 1970.
 464. Thebault, M. T., J. P. Raffin, and J. Y. Le Gall. 31P NMR studies of the metabolic changes in the prawns Palameon serratus and P. elegans during exercise. Mar. Ecol. Progr. 111: 73–78, 1994.
 465. Ting, L. H., R. Blickhan, and R. J. Full. Dynamic and static stability in hexapedal runners. J. Exp. Biol. 197: 251–269, 1994.
 466. Torres, J. J. The Functional Relation of Metabolism to Environment in Selected Mesopelagic and Vertically Migrating Species. Santa Barbara: Univ. of California, 1980. Dissertation.
 467. Torres, J. J. Relationship of oxygen consumption to swimming speed in Euphausia pacifica. II. Drag, efficiency and a comparison with other swimming organisms. Mar. Biol. 78: 231–237, 1984.
 468. Torres, J. J., and J. J. Childress. Relationship of oxygen consumption to swimming speed in Euphausia pacifica I. Effects of temperature and pressure. Mar. Biol. 74: 79–86, 1983.
 469. Trevor, J. H. The dynamics and mechanical energy expenditure of the polychaetes Nephtys cirrosa, Nereis divesicolor and Arenicola marina during burrowing. Estuarine Coastal Mar. Sci. 6: 605–619, 1978.
 470. Trombitas, K., and G. H. Pollack. Elastic properties of connecting filaments along the sarcomere. In: Mechanism of Myofilament Sliding in Muscle Contraction, 1993, vol. 332, p. 71–79.
 471. Trueman, E. R. Observation on certain mechanical properties of the ligament of Pecten. J. Exp. Biol. 30: 453–467, 1950.
 472. Trueman, E. R. (Ed). Locomotion of Soft‐Bodied Animals New York: Elsevier, 1975.
 473. Trueman, E. R. Swimming by jet propulsion. In: Aspects of Animal Movement, edited by H. Y. Elder and E. R. Trueman, Cambridge: Cambridge Univ. Press, 1980, p. 93–105.
 474. Trueman, E. R., Q. Bone, and J.‐C. Braconnot. Oxygen consumption in swimming salps (Tunicata: Thaliacea). J. Exp. Biol. 110: 323–327, 1984.
 475. Trueman, E. R., and H. D. Jones. Crawling and burrowing. In: Mechanics and Energetics of Animal Locomotion, edited by R. M. Alexander and G. Goldspink. London: Chapman and Hall, 1977, p. 204–221.
 476. Tse, F. W., C. K. Govind, and H. L. Atwood. Diverse fiber composition of swimming muscles in the blue crab, Callinectes sapidus. Can. J. Zool. 61: 52–59, 1983.
 477. Tu, M. S., and M. H. Dickinson. Modulation of negative work output from a steering muscle of the blowfly Calliphora vicina. J. Exp. Biol. 192: 207–224, 1994.
 478. Tucker, V. Energetic cost of locomotion in animals. Comp. Biochem. Physiol. 34: 841–846, 1970.
 479. Usherwood, P.N.R. The nature of “slow” and “fast” contractions in the coxal muscles of the cockroach. J. Insect Physiol. 8: 31–52, 1962.
 480. van Leeuwen, J. L. Optimum power output and structural design of sarcomeres. J. Theor. Biol. 149: 229–256, 1991.
 481. van Leeuwen, J. L. Muscle function in locomotion. In: Mechanics of Animal Locomotion, edited by R. M. Alexander, Berlin: Springer‐Verlag, 1992, p. 191–249.
 482. Videler, J. J., and B. A. Nolet. Costs of swimming measured at optimum speed: scale effects, differences between swimming styles, taxonomic groups and submerged and surface swimming. Comp. Biochem. Physiol. A 97: 91–99, 1990.
 483. Vogel, S. Flight in Drosophila. I. Flight performance of tethered flies. J. Exp. Biol. 44: 567–578, 1966.
 484. Vogel, S. Flight in Drosophila III. Aerodynamic characteristics of fly wings and wing models. J. Exp. Biol. 46: 431–443, 1967.
 485. Vogel, S. Life in Moving Fluids. The Physical Biology of Flow Princeton: Princeton Univ. Press, 1981.
 486. Vogel, S. Flow‐assisted shell reopening in swimming scallops. Biol. Bull. 169: 624–630, 1985.
 487. Vogel, S. Flow‐assisted mantle cavity refilling in jetting squid. Biol. Bull. 172: 61–68, 1987.
 488. Wadepuhl, M. Computer simulation of the hydrostatic skeleton. The physical equivalent, mathematics and application to worm‐like forms. J. Theor. Biol. 136: 379–402, 1989.
 489. Wainwright, S. A. Axis and Circumference Cambridge, MA: Harvard Univ. Press, 1988.
 490. Walcott, B., and M. Burrows. The ultrastructure and physiology of the abdominal air‐guide retractor muscles in the giant water bug, Lethocerus. J. Insect Physiol. 15: 1855–1872, 1969.
 491. Waldron, K. J. Force and motion management in legged locomotion. IEEE J. Rob. Auto. RA‐2: 214–220, 1986.
 492. Ward, D. V., and S. A. Wainwright. Locomotory aspects of squid mantle structure. J. Zool. 167: 437–449, 1972.
 493. Yamakawa, M. Molloy, J. Falkenthal, S. and D. Maughan. Myosin light chain‐2 mutation affects flight, wing beat frequency, and indirect flight muscle contraction kinetics in Drosophila. J. Cell Biol. 119: 1523–1539, 1992.
 494. Webb, P. W. Mechanics of excape responses in crayfish (Orconectes virilis). J. Exp. Biol. 79: 245–263, 1979.
 495. Webber, D. M., and R. K. O'Dor. Monitoring the metabolic rate and activity of free‐swimming squid with telemetered jet pressure. J. Exp. Biol. 126: 205–224, 1986.
 496. Wegener, G., N. M. Bolas, and A. A. G. Thomas. Locust flight metabolism studied in vivo by 31P NMR spectroscopy. Comp. Biochem. Physiol. [B] 161: 247–256, 1991.
 497. Weihs, D. Periodic jet propulsion of aquatic creatures. Forsch. Zool. 24: 171–175, 1977.
 498. Weinstein, R. B., and R. J. Full. Intermittent locomotion increases distance capacity at low temperature in the ghost crab. Am. Zool. 31: 141A, 1991.
 499. Weinstein, R. B., and R. J. Full. Intermittent exercise alters endurance in an eight‐legged ectotherm. Am. J. Physiol. 262 (Regulatory Integrative Comp. Physiol): R852–R859, 1992.
 500. Weinstein, R., and R. J. Full. Thermal dependence of locomotor energetics of the ghost crab, Ocypode quadrata. Physiol. Zool. 67: 855–872, 1994.
 501. Weinstein, R., R. J. Full, and A. N. Ahn. Dehydration decreases locomotor performance of the ghost crab. Ocypode quadrata. Physiol. Zool. 67: 873–891, 1994.
 502. Weis‐Fogh, T. Fat combustion and metabolic rate of flying locusts (Schistocerca gregaria Forskal). Phil. Trans. R. Soc. Lond. [B]. 237: 1–36, 1952.
 503. Weis‐Fogh, T. Weight economy of flying insects. Trans. Ninth Int. Congr. Entomol. 1: 341–347, 1952.
 504. Weis‐Fogh, T. Tetanic force and shortening in locust flight muscle. J. Exp. Biol. 33: 668–684, 1956.
 505. Weis‐Fogh, T. A rubber‐like protein in insect cuticle. J. Exp. Biol. 37: 889–907, 1960.
 506. Weis‐Fogh, T. Energetics of hovering flight in hummingbirds and in Drosophila. J. Exp. Biol. 56: 79–104, 1972.
 507. Weis‐Fogh, T. Quick estimates of flight fitness in hovering animals, including novel mechanisms for lift production. J. Exp. Biol. 59: 169–230, 1973.
 508. Weis‐Fogh, T., and R. M. Alexander. The sustained power output from striated muscle, In: Scale Effects in Animal Locomotion, edited by T. J. Pedley, London: Academic, 1977, p. 511–525.
 509. Weis‐Fogh, T., and M. Jensen. Biology and physics of locust flight. I. Basic principles in insect flight. A critical review. Phil. Trans. R. Soc. Lond. [B] 239: 415–458, 1956.
 510. Wells, M. The dilemma of the jet set. New Sci. 17: 44–47, 1990.
 511. Wells, M. J., O'Dor, R. K. Mangold, K. and Jowells Oxygen consumption in movement by Octopus. Mar. Behav. Physiol. 9: 289–303, 1983.
 512. Wendler, G. The coordination of walking movements in arthropods. Symp. Soc. Exp. Biol. 20: 229–249, 1966.
 513. Wensler, R. J. The ultrastructure of the indirect flight muscles of the monarch butterfly, Danaus plexippus (L.) with implications for fuel utilization. Acta Zool (Stockh.) 58: 157–167, 1977.
 514. Wheatly, M. G., B. R. McMahon, W. W. Burggsen, and A. W. Pinder. A rotating respirometer to monitor voluntary activity and associated exchange of respiratory gases in the land hermit crab (Coenobita compressus). J. Exp. Biol. 119: 85–101, 1985.
 515. White, D.C.S. The elasticity of relaxed insect fibrillar flight muscle. J. Physiol. (Lond.) 343: 31–57, 1983.
 516. Williams, T. A. Locomotion in developing Artemia larvae: mechanical analysis of antennal propulsors based on large‐scale physical models. Biol. Bull. 187: 156–163, 1994.
 517. Williams, T. A. A model of rowing propulsion and the ontogeny of locomotion in Artemia larvae. Biol. Bull. 187: 164–173, 1994.
 518. Wilson, D. M. Insect walking. Annu. Rev. Entomol. 11: 103–122, 1966.
 519. Wilson, D. M. Stepping patterns in tarantula spiders. J. exp. Biol. 47: 133–151, 1967.
 520. Winter, D. A. Biomechanics and Motor Control of Human Movement (2nd ed.). New York: Wiley, 1990.
 521. Wolf, T. J., P. Schmid‐Hempel, C. P. Ellington, and R. D. Stevenson. Physiological correlates of foraging efforts in honeybees: oxygen consumption and nectar load. Funct. Ecol. 3: 417–424, 1989.
 522. Wood, C. M., and D. J. Randall. Haemolymph gas transport, acid‐base regulation, and anaerobic metabolism during exercise in the land crab (Cardisoma camifex). J. Exp. Zool. 218: 23–35, 1981.
 523. Wood, C. M., and D. J. Randall. Oxygen and carbon dioxide exchange during exercise in the land crab. (Cardisoma carnifex). J. Exp. Zool. 218: 7–22, 1981.
 524. Wootton, R. J., and D.J.S. Newman. Whitefly have the highest contraction frequencies yet recorded in non‐fibrillar flight muscles. Nature 280: 402–403, 1979.
 525. Wu, T. Y. Introduction to the scaling of aquatic animal locomotion. In: Scale Effects in Animal Locomotion, edited by T. J. Pedley, London: Academic, 1977, p. 203–232.
 526. Yates, G. T. Hydromechanics of body and caudal fin propulsion. In: Fish Biomechanics, edited by P. W. Webb and D. Weihs. New York: Praeger, 1983, p. 177–213.
 527. Yates, G. T. How microorganisms move through water. Am. Sci. 74: 358–365, 1986.
 528. Zachar, J., and D. Zacharova. The length‐tension diagram of single muscle fibres of the crayfish. Experientia 15: 451–452, 1966.
 529. Zajac, F. E. Muscle and tendon: properties, models, scaling, and application to biomechanics and motor control. Crit. Rev. Biomed. Eng. 17: 359–411, 1989.
 530. Zanker, J. M. The wing beat of Drosophila melanogaster I. Kinematics. Phil. Trans. R. Soc. Lond. [B] 327: 1–18, 1990.
 531. Zanker, J. M., and K. Gotz. The wing beat of Drosophila melanogaster. II. Dynamics. Phil. Trans. R. Soc. Lond. [B] 327: 19–44, 1990.
 532. Zebe, E. Anaerobic metabolism in Upogebia pugettensis and Callianassa californiensis (Crustacea, Thalassinidea). Comp. Biochem. Physiol. [B] 72: 613–617, 1982.
 533. Ziegler, C. Titin‐related proteins in invertebrate muscles. Comp. Biochem. Physiol. [A] 109: 823–833, 1994.
 534. Zollikofer, C. P. Stepping patterns in ants I. Influence of speed and curvature. J. Exp. Biol. 192: 95–106, 1994.
 535. Zollikofer, C. P. Stepping patterns in ants II. Influence of body morphology. J. Exp. Biol. 192: 107–118, 1994.
 536. Zwaan, A. d., and G.v.d. Thillart. Low and high power output modes of anaerobic metabolism: invertebrate and vertebrate strategies. In: Circulation, Respiration, and Metabolism, edited by R. Gilles. Berlin: Springer‐Verlag, 1985, p. 166–192.
 537. Zwaan, A. d., R. J. Thompson, and D. R. Livingstone. Physiological and biochemical aspects of the valve snap and valve closure responses in the giant scallop Placopecten magellanicus II. Biochemistry. J. Comp. Physiol. [B] 137: 105–114, 1980.

Contact Editor

Submit a note to the editor about this article by filling in the form below.

* Required Field

How to Cite

Robert J. Full. Invertebrate Locomotor Systems. Compr Physiol 2011, Supplement 30: Handbook of Physiology, Comparative Physiology: 853-930. First published in print 1997. doi: 10.1002/cphy.cp130212