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As a fish or a bird
moves through the water
fins/wings create lift forces --
forces that tend to raise or lower
the animal. These forces
“originate from [fluid] viscosity
and are caused by asymmetries
in the flow. As fluid moves past
an object [such as a fin or a
wing], the pattern of flow may
be such that the pressure on one
…side is greater than on the
opposite. Lift is then exerted on
the object in a direction
perpendicularto the flow
direction” (Sfakiotakis 1999).
Ladybird beetle
Linden Gledhill
Katayoon Taghizade et al. Designing a Mobile Facade Using Bionic Approach. American Journal of Materials Engineering and
Technology, 2013, Vol. 1, No. 2, 22-29. doi:10.12691/materials-1-2-2
Load of wing is
taken up by the
tendon of the
supracoracoideus. Direction of muscle
Stress (force) is pull is changed by
created by the the rope-like nature
contracting muscle of the tendon, i.e., it
and tension is a pulley.
develops in the
tendon which
stretches,
Sternum moved down (and up) by supracoracoideus and pectoralis major operating
during flight: flight power movement integrated with ventilation.
Birds have pneumatic bones (air-filled) and air sacs
• Lungs of human occupy about 5% body volume. Lungs of bird only occupy 2% of body volume. But
lungs plus air sacs of a duck occupy about 20% of its body volume. Nevertheless bird air sacs are not
involved in gas exchange: they are not vacularized (no blood vessels): but they are involved in air
circulation. Air sacs are reconnected to the mesobronchi by recurrent bronchi.
• Multiple functions of air sacs: cooling system: flying produces heat and air sacs are well placed to
remove this excess heat (bird has an air-cooled engine). Shock absorber: gannets, pelicans diving into
the ocean from a great height. Resonance: sound box areas in bird sound production. Egg laying:
abdominal air sacs help in egg laying (and in defecation?).
Tiny air capillaries within the parabronchi walls, in
the (fixed-volume) lungs, are the site of actual gas
exchange.
Tracheae, which
conduct the low
density fluid (air)
into the body need
cartilaginous rings to
support their lumen
against the denser
tissues (water-
based) that
surround.
Convergence of a
sort with the
tracheae of insects.
• To select forelimbs for flight -- to alter the neuromuscular system and forelimb
form to create a powering aerofoil -- involves many complex body changes. A
‘protobird’ ancestor had to change gradually from a ground-walking reptile to a
bird capable of “level flapping flight” (i.e., from a lizard to a soaring eagle).
Needed is a plausible hypothesis of locomotion-related selection pressures: one
that envisages gradual changes to forelimbs that are adaptive throughout the
evolutionary process, i.e., are adaptive at each stage in the historical process.
• The authors of a recent paper propose an hypothesis based upon study of the
development (ontogeny) of young partridge, the ontogenetic-transitional wing
hypothesis. They studied ‘wing-stroke dynamics’ of maturing partridges and
traced a developmental pattern of forelimb movement that is consistent with
‘incline flap running’.
• Dial, K.P. et al. 2008. Nature. A fundamental avian wing-stroke provides a new
perspective on the evolution of flight. Nature 451: 985-989.
• Read this paper to understand their hypothesis, how they arrived at it and what
experimental evidence they present that tests the hypothesis.
• “…we propose that explorations of the ontogeny of post-natal behaviour and
morphology among extant taxa provide insight into ecological and evolutionary
locomotor transitional stages” (jargon destroys interest)
Chukar: Alectoris chukar
“Blue and black outlines represent the positions of the bird and wing at the start and end
of downstroke, respectively. a, In the vertebral space, the mean wing-stroke plane angle
shifts more than 30° from a more antero-posterior orientation during flap-running to
dorso-ventrally in flight, implying different wing-strokes are used to execute different
locomotor modes. The wing-stroke path is consistently oriented, however, in both the (b)
global and (c) gravitational coordinate spaces over diverse locomotor behaviours,
illustrating a simplified wing-stroke that is multi-functional. Data for juveniles are presented
from 8- to 10-day olds.”
“a, When wing-stroke plane angles are viewed
side-by-side in both the vertebral and
gravitational frames of reference, the wing-
stroke is nearly invariant relative to gravity
whereas the body axis re-orients among
different modes of locomotion. Red lines
represent the wing-tip trace in WAIR (flap-
running) and blue lines represent the wing-tip
trace in level flight. b, Wing-strokes are
estimated to produce similar aerodynamic forces
oriented about 40° above the horizon during
WAIR, level flight and descending flight. Error
bars are s.e.m. c, Representative traces of AOA
through a wing-beat for an animal flap-running
vertically (red) and in horizontal flight (blue)
demonstrate the similarities of AOA among
behaviours. The similarities are further clarified
by examining wing cross-sections and mean
global stroke-planes in the first, middle and last
thirds of downstroke. Here, the orientation of
the aerodynamic force (Faero) is estimated from
the middle third.”
Conclusions about flight evolution
• “Our experimental observations show that proto-wings moving through a stereotypic and conserved wing-stroke
have immediate aerodynamic function, and that transitioning to powered flapping flight is limited by the relative
size of the wing and muscle power, rather than development of a complex repertoire of wing-beat kinematics.”
• In other words, wingstrokes were useful to protobirds even before they could take to the air. Wing beats (at the
particular narrow range of angles the authors discovered) could help a non-flying bird ancestor to run faster, up
or down inclines – faster than ‘birds’ that were not modifying and using their forelimbs in this way. Flapping
with forelimbs and running on hindlimbs was a better, faster means of escape from a predator than running on
all four limbs at the same time.
Turning now to flight in insects: some anatomy is necessary: tergum, sternum, pleuron
are ‘top’ ‘side’ and ‘bottom’ of each insect cuticular metamere.
• Insects are descended from metameric, i.e., segmented animals and the thorax is a
locomotory tagma comprised of three metameres.
• Segments of the abdomen contrast with those of the thorax. The thorax is a
tagma functioning as a locomotory ‘box’, giving a firm base on which muscles can
originate and pull, muscles for walking (legs below) and for flying (wings above).
The mesothorax + metathorax: the two segments specialized for bearing the wings
and for flight are together called the pterothorax.
• Muscles involved in flight in insects (with exceptions, e.g., in Odonata) insert on
the exoskeleton of the thoracic box and NOT on the wings directly; they move the
wings by distorting pterothorax shape and are referred to as indirect flight
muscles.
• LONGITUDINALS DOWNSTROKE; TERGOSTERNALS UPSTROKE
• Attitude of the wings (pronation, supination) is achieved by the elastic interplay of
the veins and membranes with the air flow. The wings don’t just go up and down
and maintain elevation they must scull through the fluid (air) just like fish fins.
Locust flight {Source: R.E. Snodgrass The thoracic mechanism of a
grasshopper, and its antecedents. Smithsonian Miscellaneous Collections 82,
pp. 111. } [This reference is given just for completeness; it is not something you
should try to obtain and read, but it is the source of much of the information
given here and in the lab.]
Locusts are strong fliers. The flight-powering muscles of the locust are indirect and
have their effect upon the wings by distorting the pterothorax and by tergal tipping of
the second axillary. (The pterothorax is the flight tagma (just segments 2 & 3, not the
prothorax.) There are two antagonistic muscle sets: longitudinals (downstroke), and
tergosternals (upstroke).
• The longitudinals are situated high up in the pterothorax. Partially obscured behind
them, arrayed against the pleuron, are the many tergosternals (83, 84, 89 etc.), running
between the sterna (S2,S3) and the terga (Sct2, Sct3). The axes of the tergosternals all
lean headward (the insect’s anterior is to the left). Notice how the upper end of the
tergosternals insert on the terga where their contraction can reduce the convexity of
this region. Reducing tergal convexity is associated with elevation of the wings.
More diagramatic views: Snodgrass drew the phragmata (Aph anterior phragma, Pph posterior phragma) of Fig. 129 purposely
distorted, so as to show their interconnecting longitudinal muscles both ahead and behind: notice the critical placing of the second
axillary, 2Ax, atop the pleural wing process, WP.
Euler buckling
dorsal
ventral
Tergosternals contract and forces would tend to buckle the pleuron; the
pleural ridge stiffens the side between the wing base and the coxa.
Function of the pleurosternal muscles: even more stiffness
Imparted to the thoracic sides (pleura): and VARIABLE.