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Advances in Experimental Medicine and Biology
Volume 669
Editorial Board:
NATHAN BACK, State University of New York at BuffaloIRUN R.
COHEN, The Weizmann Institute of ScienceABEL LAJTHA, N.S. Kline
Institute for Psychiatric ResearchJOHN D. LAMBRIS, University of
PennsylvaniaRODOLFO PAOLETTI,
For further volumes :http://www.springer.com/5584
University of Milan
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Ikuo Homma · Hiroshi Onimaru ·Yoshinosuke FukuchiEditors
New Frontiers in RespiratoryControl
XIth Annual Oxford Conference on Modelingand Control of
Breathing
123
-
ISBN 978-1-4419-5691-0 e-ISBN 978-1-4419-5692-7DOI
10.1007/978-1-4419-5692-7Springer New York Dordrecht Heidelberg
London
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c© Springer Science+Business Media, LLC 2010All rights reserved.
This work may not be translated or copied in whole or in part
without the writtenpermission of the publisher (Springer
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ISSN 0065-2598
EditorsIkuo HommaDepartment of PhysiologyShowa University School
of Medicine1-5-8 Hatanodai, Shinagawa-kuTokyo 142-8555,
[email protected]
Yoshinosuke FukuchiDepartment of Respiratory MedicineJuntendo
University School of Medicine2-1-1 Hongo, Bunkyo-kuTokyo 113-8421,
[email protected]
Hiroshi OnimaruDepartment of PhysiologyShowa University School
of Medicine1-5-8 Hatanodai, Shinagawa-kuTokyo 142-8555,
[email protected]
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Preface Breathing is performed by the rhythmic contraction of
respiratory muscles. It main-tains homeostasis of the organism by
taking in the oxygen necessary to live and work and by controlling
the level of CO2 within the organism. At first glance, breathing
seems simple; however, it is produced by a complex system in the
brain with various afferents and efferents. The control of
breathing is of the utmost importance in sus-taining life, and
although more than 150 years have passed since research on
breath-ing control was first begun, many unsolved mysteries still
remain. Breathing is like watching the tides at a beach that are
created by the vast, complex open sea.
The first Oxford Conference on Modeling and Control of Breathing
was held 30 years ago in September of 1978 at the University
Laboratory of Physiology in Oxford, England. During this first
conference, the participants engaged in a hot dis-cussion on the
problem of whether breathing rhythm was produced by pacemaker cells
or a neural network. This was before the discovery of the Bötinger
complex in the medulla, and at the time, central chemoreceptive
areas were still the focus of research. This conference was an
especially unforgettable moment in the dawning of the new age of
respiratory research. It has since been held every 3 years in
various countries around the globe and is widely appreciated as the
best respiratory meeting in the world.
We were very much honored to organize the XIth Oxford Conference
in Nara, the ancient capital of Japan, and it was timely that the
city was just beginning the celebration of its founding as the
capital 1300 years ago. Nara is famous for its an-cient temples,
ruins, and forests, which collectively form the Historic Monuments
of Ancient Nara, a UNESCO World Heritage Site. The conference was
held at the Noh Theater in Nara Park and included 43 oral and 71
poster presentations focussing mainly on the control of breathing.
More than 140 participants from 15 countries attended, and we
believe that the participants could feel the tradition of old Japan
and at the same time discover new frontiers in respiratory control
in Nara. At the busi-ness meeting of the Oxford Conference, the
international committee decided that the next Oxford Conference
will be held
We are very grateful to all the participants who attend the
Conference and to the local program committees. The Editors
especially offer their heartfelt thanks to Lena Akai, Michiko
Iwase, and the members of Department of Physiology, Showa
University School of Medicine, for editorial assistance in the
production of this book. Publication of this book was supported by
Private University High Technology Research Center Project.
v
in Groningen, Holland.
The Editors
-
Conference Co-Chairs: Ikuo Homma (Tokyo, Japan) Yoshinosuke
Fukuchi (Tokyo, Japan)
vii
Conference Proceedings
-
ix
Dr. Gila Benchetrit Laboratoire de Physiologie
Respiratoire Expérimentale Université Joseph Fourier Université
de Grenoble (PRETA-TIMC, UMR CNRS 5525) 38700 La Tronche,
France
Dr. Jean Champanat Neurobiologie Génétique et
Intégrative Institut de Neurobiolo-gie Alfred Fessard, bât.
33, C.N.R.S. C.N.R.S. - U.P.R. 2216 91198, Gif-sur-Yvette,
France Dr. Ikuo Homma Department of Physiology Showa University
School
of Medicine 1-5-8 Hatanodai, Shinagawa-ku Tokyo 142-8555, Japan
Dr. Richard L. Hughson Cardiorespiratory and Vascular
Dynamics Laboratory, University of Waterloo
Waterloo, Ontario, Canada Dr. Homayoun Kazemi Pulmonary and
Critical Care Unit Harvard Medical School Bulfinch 148
Massachusetts General Hospital Boston, MA, 02114, USA
Dr. Chi-Sang Poon Harvard-MIT, Division of Health
Science & Technology, Massachusetts Institute of Technology
77 Massachusetts Avenuse- Bldg. E25-501 Cambridge, MA
02139, USA Dr. Marc J. Poulin Department of Physiology &
Biophysics Faculty of Medicine, University
of Calgary 3330 Hospital Drive NW Calgary, Alberta T2N 4N1,
Canada Dr. Peter Sheid Institute für Physiologie Ruhr-Universität
Bochum D-44780 Bochum, Germany Dr. John W. Severinghaus Department
of Anesthesiology, University of California Medical
School San Francisco, California 91143-0542, USA Dr. Peter A.
Robbins University Laboratory
of Physiology University of Oxford, Parks Road Oxford Ox1 3PT,
United Kingdom
The XIth Oxford Conference: International Organizing
Committee
-
x The XIth Oxford Conference Dr. Susan A Ward Human
Bio-Energetics Research
Centre Crickhowell Powys, NP8 1AT, United Kingdom Dr. Brian J.
Whipp Human Bio-Energetics Research
Centre Crickhowell Powys, NP8 1AT, United Kingdom
Dr. Richard Wilson Department of Physiology &
Biophysics Faculty of Medicine, University
of Calgary 3330 Hospital Drive NW Calgary, Alberta T2N 4N1,
Canada
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President Dr. Ikuo Homma Showa University School
of Medicine 1-5-8 Hatanodai, Shinagawa-ku, Tokyo142-8555,
Japan
Dr. Yoshinosuke Fukuchi Juntendo University School
of Medicine 2-1-1 Hongo, Bunkyo-ku, Tokyo 113-8421, Japan
Committee Dr. Akiko Arata Division of Physiome Department
of Physiology, Hyogo College of Medicine
1-1 Mukogawa, Nishinomiya Hyogo 663-8501, Japan
Dr. Makito Iizuka Center for Medical Sciences,
Ibaraki Prefectural University of Health Sciences
4669-2 Ami Ibaraki 300-0394, Japan
Dr. Naofumi Kimura Department of Pharmacology (II),
Jikei University School of Medicine
3-25-8 Nishi-Shinbashi, Minato-ku, Tokyo 105-8461, Japan
Dr. Hajime Kurosawa Department of Internal Medicine
and Rehabilitation Science, Tohoku University School of
Medicine
1-1 Seiryo-machi, Aoba-ku, Sendai, 980-8574, Japan
Dr. Tomoyuki Kuwaki Department of Physiology, Kagoshi-
ma University, Graduate School of Medical & Dental
Sciences
8-35-1 Sakuragaoka, Kagoshima-city Kagoshima, 890-8520,
Japan
ix
Dr. Hiroshi Kimura Second Department of Internal
Medicine, Nara Medical University School of
Medicine 840 Shijo-cho, Kashihara, Nara
634-8522, Japan
The XIth Oxford Conference: Local Organizing Committee
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The XIth Oxford Conference xii
Dr. Shun-ichi Kuwana Department of Physiology, Faculty
of Health Sciences, Uekusa Gakuen University
0639-3 Ogura-cho, Wakaba-ku, Chiba 264-0007, Japan
Dr. Yasumasa Okada Department of Medicine, Keio University
Tsukigase
Rehabilitation Center 380-2 Tsukigase, Izu City 410-3215 Japan
Dr. Yoshitaka Oku Division of Physiome, Department
of Physiology, Hyogo College of Medicine
1-1 Mukogawa, Nishinomiya Hyogo 663-8501, Japan Dr. Hiroshi
Onimaru Department of Physiology, Showa University School of
Medicine 1-5-8 Hatanodai, Shinagawa-ku
Tokyo 142-8555, Japan
Dr. Eiji Takahashi Department of Physiology, Yamagata University
School
of Medicine Yamagata 990-9585, Japan Conference Assistants Ms.
Yuki Kuwayama Showa University School
of Medicine 1-5-8 Hatanodai, Shinagawa-ku
Tokyo 142-8555, Japan Ms. Emi Kato Convention Academia Inc.
3-35-3 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan
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Conference Overview Title: The 11th Oxford Conference on
Modeling and Control of Breathing Theme: New Frontiers in
Respiratory Control Date: July 23 (Thursday) - 26 (Sunday), 2009
Venue: Nara Prefectural New Public Hall, Nara City, JAPAN
Presidents: Ikuo Homma, M.D., Ph.D. Showa University, School of
Medicine, Tokyo Yoshinosuke Fukuchi M.D., Ph.D. Juntendo
University, School of Medicine, Tokyo Web-site:
http://www.oxford-conference.com Secretariat: Head Office: Showa
University, School of Medicine, Tokyo, Japan Administration Office:
c/o Convention Academia Inc.
TEL. +81 (0)3 5808 5261 FAX. +81 (0)3 3815 2028 E-MAIL:
[email protected] Acknowledgements The Organizing
Committee would like to thank the following parties for their
generous contribution towards the success of the 11th Oxford
Conference (2009). This Conference has been supported by: – The
Commemorative Organization for the Japan World Exposition (‘70) –
The Federation of Pharmaceutical Manufacturer’s Associations of
JAPAN – Nara Prefecture – Nara City – Association for Commemorative
Events of the 1300th Anniversary of Nara
Heijokyo Capital – Nara Medical Association – Nara Medical
University – Showa University Special Thanks to: – Abbott Japan
Co., Ltd. – Otsuka Pharmaceutical Co., Ltd. – Teijin Pharma Ltd.
Corporate Sponsors – Brainvision Inc. – CHEST M.I., Inc. – FUJI
RESPIRONICS Co., Ltd. – FUKUDA DENSHI
– MINATO MEDICAL SCIENCE Co., Ltd.
xiii
4F Hongo UC Building, 3-35-3 Hongo, Bunkyo-ku, Tokyo 113-0033
JAPAN
– Inter-Reha
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9)
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Past Oxford Conferences Ist Oxford Conference University
Laboratory of Physiology, Oxford, United Kingdom (September l978)
Publication: Modelling of a Biological Control System: The
Regulation of Breathing. E.R. Carson, D.J. C. Cunningham, R.
Herczynski, D.J. Murray-Smith and E.S. Peterson, eds., Oxford:
Institute of Measurement and Control, 1978 IInd Oxford Conference
University of California Conference Centre at Lake Arrowhead
California, USA (13-16 September, 1982) Publication: Modelling and
the Control of Breathing. B.J. Whipp and D.M. Wiberg, eds.,
Elsevier Press, New York, 1983. IIIrd Oxford Conference Medieval
Abbey of Solignac, Solignac, France (September l985) Publication:
Concepts and Formalizations in the Control of Breathing. G.
Benchetrit, P. Baconnier and J. Demongeot, eds., Manchester
University Press, 1987. IVth Oxford Conference Shadow Cliff Life
Centre at Grand Lake, Grand Lake, Colorado, USA (September 1988)
Publication: Respiratory Control: A Modeling Perspective, G.D.
Swanson, F.S. Grodins, and R.L. Hughson, eds., Plenum Press, New
York, 1989. Vth Oxford Conference Fuji Institute, Fuji, Japan
(199l) Publication: Control of Breathing and its Modelling
Perspective, Y. Honda, Y. Miyamoto, K. Konno and J. Widdicombe,
eds., Plenum Press, New York, 1992.
xvii
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Pass Oxford Conference xviii
Publication: Advances in Modeling and Control of Ventilation
(Advances in Expe-rimental Medicine and Biology series, Vol. 450).
R.L. Hughson, DA. Cunningham, and J. Duffin, eds., Plenum Press,
New York, 1998. VIIIth Oxford Conference North Falmouth, Cape Cod,
Massachusetts, USA (1 1-l5 October, 2000) Publication: Frontiers in
Modeling and Control of Breathing: Integration at Mole-cular,
Cellular, and Systems Levels (Advances in Experimental Medicine and
Biolo-gy series, Vol. 499). C.-S. Poon and H. Kazemi, eds., Kluwer
Academic/Plenum Publishers, New York, 2001. IXth Oxford Conference
Paris, France (September, 2003) Publication: Post-Genomic
Perspectives in Modeling and Control of Breathing (Advances in
Experimental Medicine and Biology series, Vol. 551). J. Champagnat,
M. Denavit-Saubie', G. Fortin, A.S. Foutz. M. Thoby-Brisson, eds.,
Kluwer Academic/Plenum Publishers, New York, 2004. Xth Oxford
Conference Chateau Lake Louise, Lake Louise, Alberta, Canada (l9-24
September, 2006)
VIth Oxford Conference Royal Holloway College, Egham, Surrey,
United Kingdom (September, l994) Publication: Modeling and Control
of Ventilation (Advances in Experimental Medi-cine and Biology
series, Vol. 393). S.J.G. Semple, L. Ahms, and B.J. Whipp, eds.,
Plenum Press, New York, 1995. VIIth Oxford Conference Grandview
Inn, Huntsville, Ontario, Canada (September, 1997)
Publication: Integration in Respiratory Control, From Genes to
Systems (Advances in Experimental Medicine and Biology series, Vol.
605) M.J.Poulin, R.J.A Wilson, eds., Springer, New York, 2008. XIth
Oxford Conference Nara Prefectural New Public Hall, Nara City,
JAPAN (23-26 July, 2009) Publication: New Frontiers in Respiratory
Control (Advances in Experimental Med-icine and Biology Series), I.
Homma, H. Onimaru, and Y. Fukuchi, eds., Springer, New York,
2010.
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xix
Contents
Part I Comparative Aspects ………………………………………………… 1 1 Evidence for
a Distributed Respiratory Rhythm
Generating Network in the Goldfish (Carassius auratus) ………………. 3
Maryana Duchcherer, Andrew Kottick, and R.J.A. Wilson
2 Fictive Lung Ventilation in the Isolated Brainstem
Preparation of the Aquatic Frog, Xenopus Laevis ……………………… 9
Naofumi Kimura
Part II Development …………………………………………………........... 13 3 Loss of
Pre-Inspiratory Neuron Synchroneity in Mice
with DSCAM Deficiency ………………………………………………… 15 Kenji Amano,
Morimitsu Fujii, Satoru Arata, Masaharu Ogawa,
Kazuhiro Yamakawa, and Akiko Arata
4 Central Respiratory Failure in a Mouse Model Depends on the
Gentic Background of the Host ………………………………….. 21
Satoru Arata, Kenji Amano, Kazuhiro Yamakawa, and Akiko
Arata
5 Adrenaline Modulates on the Respiratory Network Development ……
25 Morimitsu Fujii and Akiko Arata
6 Ontogeny of Homeostasis in Mouse Hypoglossal Nucleus ………… 29
Akihito Okabe, Akiko Arata, Yoshitaka Oku, Chitoshi Takayama, and
Atsuo Fukuda 7 Anatomical Changes of Phrenic Motoneurons During
Development ………………………………………………………33 Yasumasa Okada, Shigefumi
Yokota, Yoshio Shinozaki, Fumikazu
Miwakeichi, Yoshitaka Oku, and Yukihiko Yasui
Cl-
-
Contents xx 8 Postnatal Changes in Morphology and Dendritic
Organization of
Neurones Located in the Area of the Kölliker-Fuse Nucleus of Rat
….. 37 Julia Reuter, Miriam Kron, and Mathias Dutschmann
Part III Modeling ……………………………………………………………43 9 Geometrical
Analysis of Bursting Pacemaker Neurons
Generated by Computational Models: Comparison to In Vitro
pre-Bötzinger Complex Bursting Neurons ………………………………45
Juan M. Cordovez, Christopher G. Wilson, and Irene C. Solomon 10
Origami Model for Breathing Alveoli ……………………………………49 Hiroko
Kitaoka, Carlos A. M. Hoyos, and Ryuji Takaki 11 Biologically
Variable Respiration as a Stochastic Process
in Ventilation – a Stochastic Model Study
………………........................ 53 Kyongyob Min, Keita Hosoi,
Masayuki Degami, and Yoshinori
Kinoshita 12 Future Perspectives - Proposal for Oxford Physiome
Project …..…...... 57 Yoshitaka Oku 13 Homeostatic Competition:
Evidence of a Serotonin-Gated
Spinopara Brachial Pathway for Respiratory and Thermoregulatory
Interaction ………………………………………61
Chi-Sang Poon 14 A Simplified Model for Explaining Negative
Feedback
to Beginners in Life Sciences ……………………………………………. 67 Masato
Shibuya, Yoshitaka Oku, and Ikuo Homma 15 Paradoxical Potentiation
of Exercise Hyperpnea in Congestive
Heart Failure Contradicts Sherrington Chemoreflex Model and
Supports a Respiratory Optimization Model ………………69
Chung Tin, Karlman Wasserman, Neil S. Cherniack, and Chi-Sang
Poon
Part IV Respiratory rhythm generation …………………………………..73 16
Indirect Opioid Actions on Inspiratory pre-Bötzinger
Complex Neurons in Newborn Rat Brainstem Slices ………………….75 Klaus
Ballanyi, Bogdan Panaitescu, and Araya Ruangkittisakul
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Contents xxi 17 Multiphoton/Confocal Ca2+-Imaging of
Inspiratory
pre-Bötzinger Complex Neurons at the Rostral or Caudal Surface
of Newborn Rat Brainstem Slices …………………. 81
Nicoleta Bobocea, Araya Ruangkittisakul, and Klaus Ballanyi 18
Phox2b Expressing Neurons in the Most Rostral Medulla of Newborn
Rats ……………………………………………….. 87 Hiroshi Onimaru, Keiko Ikeda, and
Kiyoshi Kawakami 19 Depression by Ca2+ and Stimulation by K+ of
Fictive Inspiratory
Rhythm in Newborn Rat Brainstem Slices ………………………………91 Bogdan
Panaitescu, Araya Ruangkittisakul, and Klaus Ballanyi 20
Glycinergic Interneurons in the Respiratory Network
of the Rhythmic Slice Preparation …………………………………........ 97
Stefan M. Winter, Jens Fresemann, Christian Schnell, Yoshitaka Oku,
Johannes Hirrlinger, and Swen Hülsmann
Part V Neuromodulation …………………………………………………. 101 21 Cholinergic
Sensitivity of the Developing Bullfrog (Rana catesbeiana) Does Not
Explain Vulnerability
to Chronic Nicotine Exposure …………………………………………. 103 Cord M.
Brundage, Carla A. Nelson, and Barbara E. Taylor 22 Modulation of
Respiratory Activity by Hypocretin-1 (Orexin A)
In Situ and In Vitro ……………………………………………………. 109 Andrea Corcoran,
George Richerson, and Michael Harris 23 Effect of JM-1232(-), a New
Sedative on Central Respiratory
Activity in Newborn Rats ………………………………………………. 115 Junya
Kuribayashi, Shun-ichi Kuwana, Yuki Hosokawa, Eiki Hatori, and
Junzo Takeda 24 PACAP Modulates the Respiratory Rhythm
Generated
in the Brainstem Slice Preparation ……………………………………. 119 Fernando
Peña
25 Caffeine Reversal of Opioid-Evoked and Endogenous
Inspiratory Depression in Perinatal Rat En Bloc Medullas and
Slices …………………………………………………………………123
Araya Ruangkittisakul, Bogdan Panaitescu, Junya Kuribayashi, and
Klaus Ballanyi
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Contents xxii
26 Acute Morphine Effects on Respiratory Activity in Mice with
Target Deletion of the Tachykinin 1 Gene (Tac1-/-) ……………. 129 Yuri
Shvarev, Jonas Berner, Andras Bilkei-Gorzo, Hugo Lagercrantz, and
Ronny Wickström
Part VI Respiratory rhythm and motor pattern generation ……………133
27 Active Inspiratory-Expiratory Phase Switching Mechanism Exists
in the Neonatal Nucleus Parabrachialis ……………………….. 135 Akiko Arata,
Ikuko Tanaka, Morimitsu Fujii, and Kazuhisa Ezure 28 Influence of
5-HT2A Receptor Blockade on Phrenic Nerve Discharge at Three Levels
of Extracellular K+ in Arterially-Perfused Adult Rat
…………………………………….... 139 Tejus A. Bale and Irene C. Solomon 29 The
Generation of Post-Inspiratory Activity in Laryngeal Motoneurons: A
Review ……………………………………………...... 143 Tara G. Bautista, Peter G.R.
Burke, Qi-Jian Sun, Robert G. Berkowitz, and Paul M. Pilowsky 30
Plasticity of Respiratory Rhythm-Generating Mechanism in Adult
Goats …………………………………………………………….. 151 Hubert V. Forster, Katie L.
Krause, Tom Kiner, Suzanne E. Neumueller, Josh M. Bonis, Baogang
Qian, and Lawrence G. Pan 31 Abdominal Respiratory Motor Pattern in
the Rat …………………… 157
Makito Iizuka 32 What Does the Multi-peaked Respiratory Output
Pattern Tell Us About the Respiratory Pattern Generating Neuronal
Network? …………………………………………………….. 163 Makio Ishiguro, Shigeharu
Kawai, Yasumasa Okada, Yoshitaka Oku, Fumi kazu Miwakeichi,
Yoshiyasu Tamura, and Amit Lal 33 The Diaphragm: a Hidden but
Essential Organ for the Mammal and the Human ………………………………………. 167
Hiroko Kitaoka and Koji Chihara
34 Upper Airway and Abdominal Motor Output During Sneezing: Is
the In Vivo Decerebrate Rat an Adequate Model? …………………173 Kenichi
Ono, Tabitha Y. Shen, Hyun Hye Chun, and Irene C. Solomon
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Con tents xxiii 35 Laudanosine has No Effects on Respiratory
Activity but Induces Non-Respiratory Excitement Activity in
Isolated Brainstem-Spinal Cord Preparation of Neonatal Rats ……………….
177 Shigeki Sakuraba, Yuki Hosokawa, Yuki Kaku, Junzo Takeda, and
Shun-ichi Kuwana 36 Influence of Extracellular [K+]o on Inspiratory
Network Complexity of Phrenic and Hypoglossal Nerve Discharge in
Arterially-Perfused Adult Rat …………………………………....... 181 Tabitha Y.
Shen, Kenichi Ono, and Irene C. Solomon 37 Bilateral Lesions of
Pontine Kölliker-Fuse Nuclei Provoke Apnea instead of Apneusis in
Anesthetized Adult Rats ……………… 185 Gang Song, Chung Tin, and
Chi-Sang Poon 38 Vesicular Glutamate Transporter 2-immunoreactive
Synapses onto Phrenic Motoneurons in the Neonatal Rat ……………………… 189
Shigefumi Yokota, Yoshio Shinozaki, Yoshitaka Oku, Yasumasa Okada,
and Yukihiko Yasui Part VII Hypoxic sensing ………………………………………………….
193 39 Hypoxic Responses of Arterial Chemoreceptors in Rabbits are
Primarily Mediated by Leak K Channels ………………………… 195 40 Halothane
and Sevoflurane Exert Different Degrees of Inhibition on Carotid
Body Glomus cell Intracellular Ca2+ Response to Hypoxia
…………………………………………….... 201 Jaideep J. Pandit and Keith J Buckler 41
Differential effects of Halothane and Isoflurane on Carotid Body
Glomus Cell Intracellular Ca2+ and Background K+ Channel Responses
to Hypoxia ………………………………………… 205 Jaideep J. Pandit, Victoria Winter,
Rebecca Bayliss, and Keith J. Buckler 42 ‘Hypoxic Ventilatory
Decline’ in the Intracellular Ca2+ Response to Sustained Isocapnic
Hypoxia in Carotid Body Glomus Cells …….. 209 Jaideep J Pandit,
Josie Collyer, and Keith J Buckler
43 Intracellular Diffusion of Oxygen and Hypoxic Sensing: Role
of Mitochondrial Respiration…………………………………… 213 Eiji Takahashi and
Michihiko Sato
N. Kobayashi and Y. Yamamoto
...
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Contents xxiv
Part VIII Integrative aspect of control of breathing
……………………..219 44 Measuring the Hypoxic Ventilatory Response
…………………………221
A. Battisti, J.A. Fisher, and J.Duffin 45 Multiple Pathways to
Long-Lasting Phrenic Motor Facilitation …..... 225 Erica A.
Dale-Nagle, Michael S. Hoffman, Peter M. MacFarlane,
46 Phase Relations Between Rhythmical Movements and Breathing in
Wind Instrument Players ……………………………………………. 231 D. Ebert, E.
Georgas, D. Rosenthal, C. Wibowo, T. Massing, T. Barth, and H.
Hefter 47 Circadian Changes in Respiratory Responses to Acute
Hypoxia and Histamine H1 Receptors in Mice ………………………………...... 235
Michiko Iwase, Yasuyoshi Ohshima, Masahiko Izumizaki, and Ikuo
Homma 48 Chemical Control of Airway and Ventilatory Responses
Mediated Via Dorsomedial Medullary 5-HT2 Receptors…………………….......
239 Mitsuko Kanamaru and Ikuo Homma 49 Hypothalamic Modulation of
Breathing ………………………………. 243 Tomoyuki Kuwaki 50 Rapid Increase to
Double Breathing Rate Appears During REM Sleep in Synchrony with
REM-A Higher CNS Control of Breathing? - ……………………………………………………………249
Shinichi Sato, Takashi Kanbayashi, Hideaki Kondo, Namiko
Matsubuchi, Kyoichi Ono, and Tetsuo Shimizu 51 The Diaphragmatic
Activities During Trunk Movements……………. 253 Minako Uga, Masatoshi
Niwa, Naoyuki Ochiai, and Sei-Ichi Sasaki Part IX Sleep apnea
………………………………………………………. 257 52 GABAergic and Glycinergic Control of
Upper Airway Motoneurons in Rapid Eye Movement Sleep …………………………..
259 Patricia L. Brooks and John H. Peever 53 Antioxidant Treatment
Does Not Prevent Chronic Hypoxia-Induced Respiratory Muscle
Impairment in Developing Rats …………………263 Jayne Carberry, Aidan
Bradford, and Ken D. O Halloran
and Gordon S. Mitchell
’
-
Con tents xxxxv 54 Respiratory Plasticity in the Behaving Rat
Following Chronic Intermittent Hypoxia …………………………………………. 267
Deirdre Edge, J. Richard Skelly, Aidan Bradford, and Ken D. O
Halloran 55 Cardiorespiratory Alterations Induced by Intermittent
Hypoxia in a Rat Model of Sleep Apnea…………………………………………. 271 Rodrigo
Iturriaga, Esteban A. Moya, and Rodrigo Del Rio 56 Model-Based
Studies of Autonomic and Metabolic Dysfunction in Sleep Apnea
………………………………………………………….. 275 Michael C.K. Khoo 57 Noradrenergic
Control of Trigeminal Motoneurons in Sleep: Relevance to Sleep
Apnea ……………………………………………… 281 Peter B. Schwarz and John H. Peever 58
Intermittent Hypoxia Impairs Pharyngeal Dilator Muscle Function in
Male But Not Female Rats ………………………………………….. 285 J. Richard Skelly,
Aidan Bradford, and Ken D. O'Halloran 59 Sleep Loss Reduces
Respiratory Motor Plasticity ……………………. 289 Arash Tadjalli and John
Peever
60 Role of Neurotrophic Signaling Pathways in Regulating
Respiratory Motor Plasticity ……………………………… 293 Arash Tadjalli and
John Peever 61 Repeated Obstructive Apneas Induce Long-term
Facilitation
of Genio lossus Muscle Tone……………………………………………. 297 Arash
Tadjalli, James Duffin, and John Peever 62 Mouse Models of Apnea:
Strain Differences in Apnea Expression and its Pharmacologic and
Genetic Modification …………………….. 303 Motoo Yamauchi, Hiroshi Kimura,
and Kingman P. Strohl
Part X Muscle and exercise ……………………………………………..... 309 63
Influence of Cycling History on the Ventilatory Response to Cycle-
Ergometry in Humans: A Role for Respiratory Memory? ………….. 311
Andrew J. Cathcart, Brian J. Whipp, Anthony P. Turner, John Wilson,
and Susan A. Ward
’
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Contents xxvi
64 Low pH Enhances Response of Thin Muscle Afferents to
Mechanical Stimuli ……………………………………………………315 Norio Hotta, Toru
Taguchi, and Kazue Mizumura 65 Effects of Deconditioning on the
Initial Ventilatory and Circulatory Responses at the Onset of
Exercise in Man ………………………….... 319 K. Ishida, K. Katayama, H.
Akima, S. Iwase, K. Sato, N. Hotta, and M. Miyamura 66 Kinetics of
the Ventilatory and Metabolic Responses to Moderate- In tensity
Exercise in Humans following Prior Exercise-Induced Metabolic
Acidaemia ……………………………………………………. 323 Susan A. Ward and Brian J.
Whipp
Part XI Higher brain function and dyspnea ……………………………. 327 67
Characteristics of Respiratory Pattern and Anxiety in Rhythmic
Gymnasts …………………………………………………. 329 68 Effects of Hypocapnia on
Spontaneous Burst Activity in the Piriform- Amygdala Complex of
Newborn Rat Brain Preparation In Vitro ……333 69 Breathing and Noh
:Emotional Breathing ……………………………. 337 Ikuo Homma 70 Patterns of
Brain Activity in Response to Respiratory Stimulation in Patients
with Idiopathic Hyperventilation (IHV) …………………. 341 71 Respiratory
Response toward Olfactory Stimuli might be an Index for
Odor-Induced Emotion and Recognition ………………….. 347 72
Periaqueductal Gray Control of Breathing ………………………….... 353 Hari H.
Subramanian and Gert Holstege Author Index
………………………………………………………………... 359 Subject Index
………………………………………………………….......... 367
Lena Akai, Sakuko Ishizaki, Masao Matsuoka, and Ikuo Homma
T. Fujii, H. Onimaru, M. Suganuma, and I. Homma
S. Jack, G.J. Kemp, W.E. Bimson, P.M.A. Calverley, and D.R.
Corfield
Yuri Masaoka and Ikuo Homma
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Part IComparative Aspects
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1 Evidence for a Distributed Respiratory Rhythm Generating
Network in the Goldfish (Carassius auratus)
1 Department of Physiology and Biophysics, University of
Calgary, Calgary, Canada, [email protected]
2 Department of Physiology and Biophysics, University of
Calgary, Calgary, Canada, [email protected]
3 Department of Physiology and Biophysics, University of
Calgary, Calgary, Canada, [email protected]
Abstract Central pattern generators located in the brainstem
regulate ventilatory behaviors in verte-brates. The development of
the isolated brainstem preparation has allowed these neural
networks to be characterized in a number of aquatic species. The
aim of this study was to explore the architec-ture of the
respiratory rhythm-generating site in the goldfish (Carassius
auratus) and to determine the utility of a newly developed isolated
brainstem preparation, the Sheep Dip. Here we provide evidence for
a distributed organization of respiratory rhythm generating neurons
along the rostro-caudal axis of the goldfish brainstem and outline
the advantages of the Sheep Dip as a tool used to survey neural
networks.
1 Introduction
Water-breathing fish use a buccal force pump to produce
unidirectional flow of water over the gills. Central pattern
generators in the brainstem regulate this activity and have been
characterized in a number of lower vertebrates (Kawasaki 1979;
Wilson et al. 2000). The evolutionary transition from water
ventilation to air breathing likely required a func-tional
reorganization of some primitive respiratory CPG (Milsom 2008). In
order to fully understand the mechanisms by which ventilatory drive
is generated and maintained in modern air-breathers, it may be
important to characterize the neural networks that govern water
ventilation in our fish ancestors.
Adrian and Buytendijk (1931) pioneered the field of the isolated
fish brainstem prep-aration using the goldfish (Cyprinus
carassius). They were able to demonstrate the exis-tence of a
neural network capable of producing rhythmic output in the absence
of any peripheral feedback. Using similar preparations, respiratory
rhythm generators have been investigated in a number of aquatic
species including the water-breathing lamprey ( 1977), and the
facultative air-breathing gar (Wilson et al. 2000).
In early stages of development the respiratory network appears
to be diffuse, with each maturing rhombomere containing its own
respiratory rhythm generating circuit (Fortin et al.1995). In some
fish species such diffuse organization is thought to per-
3I. Homma et al. (eds.), New Frontiers in Respiratory Control,
Advances in Experimental Medicine and Biology 669, DOI
10.1007/978-1-4419-5692-7_1, © Springer Science+Business Media, LLC
2010
Maryana Duchcherer1, Andrew Kottick2, and R.J.A. Wilson3
Rovainen
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sist into adulthood, contrasting the tightly nucleated networks
often observed in amphibians and mammals (Rekling and Feldman 1998;
Vasilakos et al. 2006).
It is hypothesized that the fish central nervous system contains
a local rhythm ge-nerating area that drives gill ventilation. The
aim of this study was to investigate the architecture of the
respiratory rhythm generator in the goldfish using a novel,
verti-cally mounted, isolated brainstem preparation known as the
Sheep Dip.
2 Methods
This study was carried out on goldfish (C. auratus).
Experimental procedures were ap-proved by the Animal Care Committee
at the University of Calgary. Prior to experimen-tation, each
animal was anesthetized in 1% MS-222 solution until unresponsive to
touch. Once fully anesthetized, the fish were mounted horizontally
on a Sylgard™ platform and superfused with fish artificial
cerebrospinal fluid (aCSF) cooled to 2–5˚C, aerated with 98% O2/2%
CO2, and containing (in mM): 120 NaCl, 3.5 KCl, 1.3 MgCl, 11
D-Glucose, 13 NaHCO3, 1.25 NaH2PO4, and 2 CaCl. The brainstem was
dis-sected from the level of the rostral tectum to the caudal
spinal cord, isolated and trans-ferred to the Sheep Dip apparatus
(Fig. 1) where it was mounted vertically on a fixed stage and fully
immersed in chamber containing aCSF.
Fig. 1 The Sheep Dip isolated brainstem preparation
4 M. Duchcherer et al.
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Evidence for a Distributed Respiratory Rhythm Generating Network
in the Goldfish
Rhythmic bursts of activity, assumed to be respiratory, were
recorded from cranial nerve VII rootlet using an extracellular
glass microelectrode. The signal was amplified using a differential
AC amplifier (A-M Systems, Inc.). Data was acquired using the
Digi-data 1322A 16-bit data acquisition system (Axon
Instruments/Molecular Devices) at a sampling rate of 5 KHz.
Axoscope 10.1 software (Axon Instruments/Molecular Devices) was
used to visualize and analyze the collected data.
After an initial stabilization period (30–60 min) the chamber
containing aCSF was lowered: adequate superfused of the preparation
was maintained entirely by a drip system. The aCSF in the chamber
was replaced with aCSF containing high magnesium (40 mM) and low
calcium (0mM) intended to block synaptic activity. At the beginning
of each experiment the entire preparation was briefly exposed to
the high magnesium aCSF to ensure it was able to block rhythmic
output. Following a second stabilization period under the aCSF
drip, the chamber containing high mag-nesium was raised in 500 µM
increments at 10 min intervals in a caudal to rostral direction
beginning at the most caudal cranial nerve root. Upon complete
cessation of rhythmic output, a 2% neutral red solution was added
to the chamber for several minutes.
3 Results
Initially, when the level of high magnesium solution was below
the most caudal cranial nerve root, CN VII produced a rhythmic
motor output with a mean frequency of 44.1 +/− 7.5 min-1 (n = 6).
The pattern was characterized by rhythmic high frequency bursts
which in some preparations were occasionally interrupted by bursts
of slightly larger amplitude. Each experiment began with complete
submersion of the preparation in the high magnesium solution to
ensure that it was able to block rhythmic output. In every
experiment block of motor output was achieved within 10 seconds and
the preparation recovered when returned to control conditions.
Starting at the caudal most CN root, the chamber was raised in
500 µM intervals every 10 min submersing more and more of the
preparation (Fig. 2A). In all six preparations there was a
progressive detrimental effect on respiratory frequency leading to
burst cessation (Fig. 2B). The number of submersion steps necessary
to block output completely varied from 6–8 steps (3–4mm). The first
trial received an additional washout period after the sheep dip was
complete. During this washout, the frequency of respiratory motor
output returned to control levels.
5