Skip to main content
Advertisement

Main menu

  • Home
  • Articles
    • Accepted manuscripts
    • Issue in progress
    • Latest complete issue
    • Issue archive
    • Archive by article type
    • Special issues
    • Subject collections
    • Interviews
    • Sign up for alerts
  • About us
    • About JEB
    • Editors and Board
    • Editor biographies
    • Travelling Fellowships
    • Grants and funding
    • Journal Meetings
    • Workshops
    • The Company of Biologists
    • Journal news
  • For authors
    • Submit a manuscript
    • Aims and scope
    • Presubmission enquiries
    • Article types
    • Manuscript preparation
    • Cover suggestions
    • Editorial process
    • Promoting your paper
    • Open Access
    • Outstanding paper prize
    • Biology Open transfer
  • Journal info
    • Journal policies
    • Rights and permissions
    • Media policies
    • Reviewer guide
    • Sign up for alerts
  • Contacts
    • Contact JEB
    • Subscriptions
    • Advertising
    • Feedback
  • COB
    • About The Company of Biologists
    • Development
    • Journal of Cell Science
    • Journal of Experimental Biology
    • Disease Models & Mechanisms
    • Biology Open

User menu

  • Log in

Search

  • Advanced search
Journal of Experimental Biology
  • COB
    • About The Company of Biologists
    • Development
    • Journal of Cell Science
    • Journal of Experimental Biology
    • Disease Models & Mechanisms
    • Biology Open

supporting biologistsinspiring biology

Journal of Experimental Biology

  • Log in
Advanced search

RSS  Twitter  Facebook  YouTube  

  • Home
  • Articles
    • Accepted manuscripts
    • Issue in progress
    • Latest complete issue
    • Issue archive
    • Archive by article type
    • Special issues
    • Subject collections
    • Interviews
    • Sign up for alerts
  • About us
    • About JEB
    • Editors and Board
    • Editor biographies
    • Travelling Fellowships
    • Grants and funding
    • Journal Meetings
    • Workshops
    • The Company of Biologists
    • Journal news
  • For authors
    • Submit a manuscript
    • Aims and scope
    • Presubmission enquiries
    • Article types
    • Manuscript preparation
    • Cover suggestions
    • Editorial process
    • Promoting your paper
    • Open Access
    • Outstanding paper prize
    • Biology Open transfer
  • Journal info
    • Journal policies
    • Rights and permissions
    • Media policies
    • Reviewer guide
    • Sign up for alerts
  • Contacts
    • Contact JEB
    • Subscriptions
    • Advertising
    • Feedback
Research Article
Dynamic pressure maps for wings and tails of pigeons in slow, flapping flight, and their energetic implications
James R. Usherwood, Tyson L. Hedrick, Craig P. McGowan, Andrew A. Biewener
Journal of Experimental Biology 2005 208: 355-369; doi: 10.1242/jeb.01359
James R. Usherwood
  • Find this author on Google Scholar
  • Find this author on PubMed
  • Search for this author on this site
Tyson L. Hedrick
  • Find this author on Google Scholar
  • Find this author on PubMed
  • Search for this author on this site
Craig P. McGowan
  • Find this author on Google Scholar
  • Find this author on PubMed
  • Search for this author on this site
Andrew A. Biewener
  • Find this author on Google Scholar
  • Find this author on PubMed
  • Search for this author on this site
  • Article
  • Figures & tables
  • Info & metrics
  • PDF
Loading

Article Figures & Tables

Figures

  • Table 2.

    Morphology, kinematics and results for acceleration-compensated experiments

    Loaded mass m (kg)0.483±0.027
    Wing span b (m)0.660±0.01
    Frequency f (Hz)8.02±0.69
    Flight velocity V (m s-1)4.46±0.27
    Tail angle of attack Embedded Image (deg.)47.6±6.4
    Inclination of wing stroke plane β (deg.)35.1±2.6
    Tail area Stail (m2)0.0204±0.0020
    Differential pressure: inner tail (Pa)18.2±1.0
    Differential pressure: outer tail (Pa)32.9±4.5
    Differential pressure: mean Embedded Image (Pa)25.6±8.5
    Weight support %7.91±2.5
    Direct power (W)1.70±0.2
    Body mass specific power (W kg-1)3.68±0.58
    Muscle mass specific power (W kg-1)19.3±3.0
    Embedded Image 2.14
    Embedded Image 1.44
    Embedded Image 1.59
    • Values are means ± s.d. (N=3).

      The muscle mass used for the muscle mass specific power calculation is that of the pair of pectoralis muscles (18% body mass); while the power costs here are due to the tail, they are largely `paid' for by the wing-flapping musculature.

  • Table 3.

    Morphology and kinematics for non-acceleration-compensated experiments

       Loaded mass m (kg)0.460±0.040
       Wingspan b (m)0.664±0.02
       Frequency f (Hz)7.98±0.46
       Wing length R (m)0.302±0.022
       Single wing area (m2)0.0296±0.0054
    Kinematic means for 3 middle flaps
       Peak upstroke angle φx-z,max (deg.)86.2±7.5
       Angular velocity φx-z (deg. s-1)–2023±60
       Downstroke time Tds (s)0.0673±0.0023
    •    Values are means ± s.d. (N=3).

  • Fig. 3.
    • Download figure
    • Open in new tab
    • Download powerpoint
    Fig. 3.

    Average acceleration-compensated pressure measurements for inner and outer tail positions (red lines) and two positions along the wings (blue lines), with non-acceleration compensated results (black lines) underlying. The synchronising accelerometer trace (green lines throughout; black lines indicate ±1 s.d.; N=3 pigeons) from an accelerometer placed at the base of secondary S3 allows signals from separate flights to be combined; each cycle is defined by the peak in accelerometer signal, relating to rapid upwards acceleration of the wing towards the end of downstroke. The time-base for each raw signal is normalised to the wing stroke cycle (thus slight differences in frequency do not result in increasing variation through the cycle). In each trace, the signal is split into a take-off flap followed by three full flaps, three full flaps at the middle of the flight, and three full flaps prior to landing followed by a landing flap. Underlying grey bars indicate periods of downstroke throughout.

  • Fig. 4.
    • Download figure
    • Open in new tab
    • Download powerpoint
    Fig. 4.

    An expansion of Fig. 3 for the middle three flaps only, with traced kinematics of the wing stroke included. The effect of acceleration is highlighted (i) for the distal wing position, and relates to the high acceleration (up to 80 g) at the `clap' at the upstroke/downstroke transition. The effects of acceleration-compensation are generally slight for both tail and wing differential pressure signals.

  • Fig. 5.
    • Download figure
    • Open in new tab
    • Download powerpoint
    Fig. 5.

    Mean acceleration-compensated differential pressures for inner and outer tail positions, and distal and proximal wing positions. Also included are acceleration signals for inner and outer tail positions (second pair of traces) and the proximal wing position - the synchronising accelerometer. Tracings of the pigeon through the wing stroke cycle match the timing of the graph. Values are mean ±1 s.d. (shown in black; N=3 pigeons). The vertical red broken lines show the relationship between a peak in both inner and outer tail pressure signal, an upwards acceleration of the wings towards the end of downstroke, and a concurrent downwards acceleration of the tail.

  • Fig. 6.
    • Download figure
    • Open in new tab
    • Download powerpoint
    Fig. 6.

    Mean non-acceleration-compensated differential pressure signals for eight positions (P8, P7, P4, P3, S1, S2, S6, S7) along and across the wing. The traces are ordered from distal sites at the top towards proximal sites nearer the bottom, and the synchronising accelerometer trace again appears as the last trace, in green. Values are mean ±1 s.d. (shown in black; N=3 pigeons). Signals for the five sites near the leading edge of the wing are shown in red. Signals for the three pressure sites towards the trailing edge are shown in blue. Periods of downstroke defined by kinematic observations of the wrist are shown as grey for two of the middle three flaps. This shows the close relationship between downstroke timing and the development of `positive' (ventral-to-dorsal) differential pressures, which are used to define downstroke periods shown with underlying grey boxes for take-off and landing. LE, leading edge; TE, trailing edge.

  • Fig. 7.
    • Download figure
    • Open in new tab
    • Download powerpoint
    Fig. 7.

    Expansion of Fig. 6 for the middle three flaps: non-acceleration-compensated pressures for eight sites (P8, P7, P4, P3, S1, S2, S6, S7) along and across the wing. Six pigeon outlines relate to the same timing as the plots, and are labelled a-f. Each tracing is separated in time by an interval of approximately 16 ms (although the rest of the timing remains normalised to the cycle as defined by the synchronising accelerometer peaks). Positions (a) and (f) are the upstroke/downstroke transition, and occur just after the trough in synchronising accelerometer signal. Position (b) indicates mid downstroke, a period of little wing acceleration, coinciding with peak pressure for sites along the leading edge. Position (c) is towards the end of downstroke, when the synchronising accelerometer is being accelerated upwards. Position (d) relates to a downwards acceleration of the proximal wing, after which the distal wing opens up (position e), completing upstroke. LE, leading edge; TE, trailing edge.

  • Fig. 1.
    • Download figure
    • Open in new tab
    • Download powerpoint
    Fig. 1.

    Positioning of accelerometers (rectangles) and differential pressure transducers (circles) across (A) tail, (B) wing for acceleration-compensated trials, and (C) the non-compensated 8-position map. Following convention we code our feathers R for rectrices (tail feathers), P for primaries and S for secondaries, with numbering for the wing feathers counted from the primary/secondary boundary. The pressures measured at each of five positions along the leading edge are taken as representative for the wing sections demarcated by straight lines in C.

  • Table 1.

    Transducer placements

    Accelerometers
    Differential pressure sensors± 250 g± 50 g
    ABABAB
    Acceleration-compensated experiments
       Inner tailLeft inner tail (R2), mid-featherRight inner tail (R2), mid-featherXRight S3* baseLeft R2 baseRight R2 base
       Outer tailLeft outer tail (R5), mid-featherRight outer tail (R5), mid-featherXRight S3* baseLeft R5Right R5
       Right wingP8, 25 mm from baseS3, 25 mm from baseRight P8 baseRight S3* baseXX
    Non-acceleration-compensated experiments
       P8P7Left P8 midLeft P7 base(Right P8 base)(Right S3 base)Left S3* baseX
       P4P3Left P4 baseLeft P3 mid(Right P8 base)(Right S3 base)Left S3* baseX
       S1S2Left S1 baseLeft S2 mid(Right P8 base)(Right S3 base)Left S3* baseX
       S6S7Left S6 midLeft S7 base(Right P8 base)(Right S3 base)Left S3* baseX
    • R indicates a rectrice (tail feather), with numbers indicating feathers (of 6) from the centre. P indicates a primary feather, S indicates a secondary feather, and numbering counts from the primary/secondary boundary (thus, given there are 10 visible primaries in an adult pigeon wing, P8 is the third visible primary from the wing tip).

      S3* indicates the synchronising accelerometer. X indicates the sensor was not used for a given set-up. Values from sensors shown in parentheses (those on the right wing for non acceleration-compensated experiments) were measured but are not discussed here.

  • Fig. 2.
    • Download figure
    • Open in new tab
    • Download powerpoint
    Fig. 2.

    The axes and measured angles for side view (A) and front view (B) during level flight between two perches. For an explanation of symbols, see List of symbols.

  • Fig. 8.
    • Download figure
    • Open in new tab
    • Download powerpoint
    Fig. 8.

    The implications of the differential pressure measurements of the five sites towards the leading edge of the wing (P8, P7, P4, S1 and S7), and the downstroke kinematics, during the earlier two of the middle three flaps (complete flap cycles are required). (A) Differential pressures. (B) The variation in wing stroke angle during downstroke within the x-z plane, i.e. when observed from front-on (φx-z), is shown in black, the angle within the stroke plane (φ) in red. (C) The calculated geometric velocity (including forward flight speed) for each wing section. Mean section forces assuming that the measured point pressures can be taken as mean pressures for relevant wing sections of known area (D) are combined, and their orientations to the vertical are taken into account when calculating contribution to weight (BW) support (E). The effective moment arm for the aerodynamic force on the wing (Reff), or the effective centre of pressure, acts approximately half way along the wing (F). Instantaneous muscle-mass specific powers (G) are calculated assuming that the pectoralis dominates downstroke power, and constitutes 18% of body mass. Means± s.d. for weight support, effective moment arm and powers, are calculated from appropriate individual average measurements of differential pressure, kinematics and morphology; averaging is performed as the final step.

  • Table 4.

    Muscle-mass specific aerodynamic powers – wing results

    Cycle powers (weight support 74.5±7.7%)
    For both wings For a single wing or pectoralis
    Aerodynamic power (W)Body-mass specific power (W kg-1)Muscle-mass specific power (W kg-1)Peak force on a wing (N)Peak force on a pectoralis (N)Peak pectoralis stress (kNm2)
    Method for power calculation
       Direct measurements: mean kinematics and 5-site pressure map applied to 3 pigeons25.6±3.851.2±5.1272.7±26.75.759.074.1
       Induced power6.5914.379.5
    • Induced power is calculated following Pennycuick (1989) with the mean wing span and mass from Table 3 and flight speed from Table 2. The induced power factor k is taken as its default value of 1.2. Muscle-mass specific powers are calculated assuming that the pectoralis dominates downstroke power and constitutes 18% of body mass. Muscle force and stress are calculated using the wing geometry (specifically the shoulder to delto-pectoral crest distance and the cross section of the pectoralis) from pigeons used in other studies (Dial and Biewener, 1993; Biewener et al. 1998; Soman et al., 2005) scaled to the mean body mass of the wild-type pigeons used here, for which a value of 7.385 cm2 was used for the physiological cross-sectional area of the muscle. These values, therefore, should be considered approximate.

Previous ArticleNext Article
Back to top
Previous ArticleNext Article

This Issue

 Download PDF

Email

Thank you for your interest in spreading the word on Journal of Experimental Biology.

NOTE: We only request your email address so that the person you are recommending the page to knows that you wanted them to see it, and that it is not junk mail. We do not capture any email address.

Enter multiple addresses on separate lines or separate them with commas.
Dynamic pressure maps for wings and tails of pigeons in slow, flapping flight, and their energetic implications
(Your Name) has sent you a message from Journal of Experimental Biology
(Your Name) thought you would like to see the Journal of Experimental Biology web site.
CAPTCHA
This question is for testing whether or not you are a human visitor and to prevent automated spam submissions.
Share
Research Article
Dynamic pressure maps for wings and tails of pigeons in slow, flapping flight, and their energetic implications
James R. Usherwood, Tyson L. Hedrick, Craig P. McGowan, Andrew A. Biewener
Journal of Experimental Biology 2005 208: 355-369; doi: 10.1242/jeb.01359
del.icio.us logo Digg logo Reddit logo Twitter logo CiteULike logo Facebook logo Google logo Mendeley logo
Citation Tools
Research Article
Dynamic pressure maps for wings and tails of pigeons in slow, flapping flight, and their energetic implications
James R. Usherwood, Tyson L. Hedrick, Craig P. McGowan, Andrew A. Biewener
Journal of Experimental Biology 2005 208: 355-369; doi: 10.1242/jeb.01359

Citation Manager Formats

  • BibTeX
  • Bookends
  • EasyBib
  • EndNote (tagged)
  • EndNote 8 (xml)
  • Medlars
  • Mendeley
  • Papers
  • RefWorks Tagged
  • Ref Manager
  • RIS
  • Zotero
Alerts

Please log in to add an alert for this article.

Sign in to email alerts with your email address

Article navigation

  • Top
  • Article
    • SUMMARY
    • Introduction
    • Materials and methods
    • Results
    • Discussion
    • ACKNOWLEDGEMENTS
    • References
  • Figures & tables
  • Info & metrics
  • PDF

Related articles

Cited by...

More in this TOC section

  • Sex-specific microhabitat use is associated with sex-biased thermal physiology in Anolis lizards
  • Early developmental stages of native populations of Ciona intestinalis under increased temperature are affected by local habitat history
  • Three auditory brainstem response (ABR) methods tested and compared in two anuran species
Show more RESEARCH ARTICLE

Similar articles

Other journals from The Company of Biologists

Development

Journal of Cell Science

Disease Models & Mechanisms

Biology Open

Advertisement

Meet the Editors at SICB Virtual 2021

Reserve your place to join some of the journal editors, including Editor-in-Chief Craig Franklin, at our Meet the Editor session on 17 February at 2pm (EST). Don’t forget to view our SICB Subject Collection, featuring relevant JEB papers relating to some of the symposia sessions.


2020 at The Company of Biologists

Despite 2020's challenges, we were able to bring a number of long-term projects and new ventures to fruition. As we enter a new year, join us as we reflect on the triumphs of the last 12 months.


The Big Biology podcast

JEB author Christine Cooper talks to Big Biology about her research. In this fascinating JEB sponsored podcast she tells us how tough zebra finches adjust their physiology to cope with extreme climate events. 


Developmental and reproductive physiology of small mammals at high altitude

Cayleih Robertson and Kathryn Wilsterman focus on high-altitude populations of the North American deer mouse in their review of the challenges and evolutionary innovations of pregnant and nursing small mammals at high altitude.


Read & Publish participation extends worldwide

“Being able to publish Open Access articles free of charge means that my article gets maximum exposure and has maximum impact, and that all my peers can read it regardless of the agreements that their universities have with publishers.”

Professor Roi Holzman (Tel Aviv University) shares his experience of publishing Open Access as part of our growing Read & Publish initiative. We now have over 60 institutions in 12 countries taking part – find out more and view our full list of participating institutions.

Articles

  • Accepted manuscripts
  • Issue in progress
  • Latest complete issue
  • Issue archive
  • Archive by article type
  • Special issues
  • Subject collections
  • Interviews
  • Sign up for alerts

About us

  • About JEB
  • Editors and Board
  • Editor biographies
  • Travelling Fellowships
  • Grants and funding
  • Journal Meetings
  • Workshops
  • The Company of Biologists
  • Journal news

For Authors

  • Submit a manuscript
  • Aims and scope
  • Presubmission enquiries
  • Article types
  • Manuscript preparation
  • Cover suggestions
  • Editorial process
  • Promoting your paper
  • Open Access
  • Outstanding paper prize
  • Biology Open transfer

Journal Info

  • Journal policies
  • Rights and permissions
  • Media policies
  • Reviewer guide
  • Sign up for alerts

Contact

  • Contact JEB
  • Subscriptions
  • Advertising
  • Feedback

 Twitter   YouTube   LinkedIn

© 2021   The Company of Biologists Ltd   Registered Charity 277992