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Science, Strategy and War

John Boyd is often known exclusively for the so-called ‘OODA’ loop model he
developed. This model refers to a decision-making process and to the idea that
military victory goes to the side that can complete the cycle from observation to
action the fastest. This book aims to redress this state of affairs and re-examines
John Boyd’s original contribution to strategic theory. By highlighting diverse
sources that shaped Boyd’s thinking, and by offering a comprehensive overview
of Boyd’s work, this volume demonstrates that the common interpretation of the
meaning of Boyd’s OODA loop concept is incomplete. It also shows that
Boyd’s work is much more comprehensive, richer and deeper than is generally
thought. With his ideas featuring in the literature on Network Centric Warfare, a
key element of the US and NATO’s so-called ‘military transformation’ pro-
grammes, as well as in the debate on Fourth Generation Warfare, Boyd con-
tinues to exert a strong influence on Western military thinking. Dr Osinga
demonstrates how Boyd’s work can help us to understand the new strategic
threats in the post-9/11 world, and establishes why John Boyd should be
regarded as one of the most important (post)modern strategic theorists.

Frans P.B. Osinga is a serving officer of the Royal Netherlands Air Force, a
former F-16 pilot, and a graduate of the Royal Netherlands Military Academy
and the Netherlands Defence College. He completed this study while serving as
the Netherlands MoD Senior Research Fellow at the Clingendael Institute of
International Relations in The Hague. He is currently stationed at the NATO
Headquarters of the Supreme Allied Command Transformation in Norfolk,
Virginia.
Series: strategy and history
Edited by Colin Gray and Williamson Murray
ISSN: 1473-6403

This new series will focus on the theory and practice of strategy. Following
Clausewitz, strategy has been understood to mean the use made of force, and the
threat of the use of force, for the ends of policy. This series is as interested in
ideas as in historical cases of grand strategy and military strategy in action. All
historical periods, near and past, and even future, are of interest. In addition to
original monographs, the series will from time to time publish edited reprints of
neglected classics as well as collections of essays.

1 Military Logistics and Strategic Performance

Thomas M. Kane

2 Strategy for Chaos

Revolutions in military affairs and the evidence of history
Colin Gray

3 The Myth of Inevitable US Defeat in Vietnam

C. Dale Walton

4 Astropolitik
Classical geopolitics in the space age
Everett C. Dolman

5 Anglo-American Strategic Relations and the Far East, 1933–1939

Imperial crossroads
Greg Kennedy

6 Pure Strategy
Power and principle in the space and information age
Everett C. Dolman

7 The Red Army, 1918–1941

From vanguard of world revolution to US ally
Earl F. Ziemke
 8 Britain and Ballistic Missile Defence, 1942–2002
Jeremy Stocker

9 The Nature of War in the Information Age

Clausewitzian future
David J. Lonsdale

10 Strategy as Social Science

Thomas Schelling and the nuclear age
Robert Ayson

11 Warfighting and Disruptive Technologies

Disguising innovation
Terry Pierce

12 The Fog of Peace and War Planning

Military and strategic planning under uncertainty
Edited by Talbot C. Imlay and Monica Duffy Toft

13 US Army Intervention Policy and Army Innovation

From Vietnam to Iraq
Richard Lock-Pullan

14 German Disarmament After World War I

The diplomacy of international arms inspection 1920–1931
Richard J. Shuster

15 Strategy and History

Essays on theory and practice
Colin S. Gray

16 The German 1918 Offensives

A case study in the operational level of war
David T. Zabecki

17 Special Operations and Strategy

From World War II to the War on Terrorism
James D. Kiras

18 Science, Strategy and War

The strategic theory of John Boyd
Frans P.B. Osinga

19 US Defense Strategy from Vietnam to Operation Iraqi Freedom

Military innovation and the new American way of war, 1973–2003
Robert R. Tomes
Science, Strategy and War
The strategic theory of John Boyd

Frans P.B. Osinga

First published 2007
by Routledge
2 Park Square, Milton Park, Abingdon, Oxon OX14 4RN
Simultaneously published in the USA and Canada
by Routledge
270 Madison Ave, New York, NY 10016
Routledge is an imprint of the Taylor & Francis Group, an informa business
This edition published in the Taylor & Francis e-Library, 2006.
“To purchase your own copy of this or any of Taylor & Francis or Routledge’s
collection of thousands of eBooks please go to www.eBookstore.tandf.co.uk.”
© 2007 Frans P.B. Osinga
All rights reserved. No part of this book may be reprinted or reproduced or
utilized in any form or by any electronic, mechanical, or other means, now
known or hereafter invented, including photocopying and recording, or in
any information storage or retrieval system, without permission in writing
from the publishers.
British Library Cataloguing in Publication Data
A catalogue record for this book is available from the British Library
Library of Congress Cataloging in Publication Data
A catalog record for this book has been requested
ISBN 0-203-08886-7 Master e-book ISBN

ISBN10: 0-415-37103-1 (hbk)

ISBN10: 0-203-08886-7 (ebk)

ISBN13: 978-0-415-37013-2 (hbk)

ISBN13: 978-0-203-08886-9 (ebk)
For Ankie, Timon, Femmeke and Minke.
Who else?
We are survival machines.
Richard Dawkins

Strategy is the mode of survival of a society.

Henry Kissinger

He who can handle the quickest rate of change survives.

John Boyd
Contents

List of figures xi
List of boxes xii
Acknowledgments xiii

1 Introduction 1
Introducing A Discourse 1
‘A towering figure’ 3
‘Deeply flawed’ 5
Aim and argument 6
A note on strategy 8
A note on strategic theory 11
The formative factors of strategic theory 15
Organization of this study 18

2 The seeds of a theory and the fertile soil 20

The seed of a theory; Boyd’s military life 20
Reading history 28
Fertile soil: the US Military after Vietnam 42
Experience, curiosity and challenges 50

3 Science: Boyd’s fountain 52

Boyd and science 52
Shifting foundations 54
Paradigm shift 64
The emerging systems view of the world 69
Systems everywhere 74
Boyd and the first stage of the paradigm shift 81
Conclusion 85
x Contents
4 Completing the shift 86
Riding the wave 86
Beyond open and chaotic systems: complexity theory 95
The postmodern turn 106
Chaos everywhere 112
A Discourse and the scientific Zeitgeist 121
Concluding words 127

5 Core arguments 128

A Discourse in prose 128
Boyd’s ‘Abstract’ of A Discourse 130
Destruction and Creation 131
Patterns of Conflict 139
Concluding words 188

6 Exploration and refinement 189

Introduction 189
Organic Design for Command and Control 189
The Strategic Game of ? and ? 201
Revelation 218
The Conceptual Spiral 219
The Essence of Winning and Losing 229
Conclusion 233

7 Completing the loop 234

Beyond the rapid OODA idea 234
The continuing relevance of A Discourse 243
Analysis/synthesis 255

Annexes 258
Notes 260
Select bibliography 292
Index 304
Figures

1.1 Simplified drawing of the OODA loop 2

3.1 Example of a simple feedback loop 72
3.2 Example of a cybernetic model of the brain 75
3.3 Model of a double loop learning process 76
3.4 Refined model of generative learning 78
4.1 Bifurcation diagram 91
4.2 Functioning of schemata according to Murray Gell-Mann 99
5.1 Boyd’s starting idea 141
5.2 Dynamics of attrition warfare 167
5.3 The essence of maneuver conflict (1) 168
5.4 Causal chain of effects in maneuver conflict 168
5.5 The essence of maneuver conflict (2) 169
5.6 The essence of moral conflict (1) 171
5.7 The essence of moral conflict (2) 172
5.8 Influencing the enemy’s strategic behavior 176
5.9 Theme for disintegration and collapse 178
5.10 Theme for vitality and growth 182
6.1 The real OODA loop 231
Boxes

2.1 Chronology of Boyd’s life 21

2.2 Sun Tzu’s stratagems 38
3.1 Characteristics of creative people 79
3.2 Definitions of organizational learning 81
3.3 Positive and negative indicators of learning organizations 82
4.1 Boyd’s reading list at the time of his death 87
4.2 Traditional versus emerging worldview 88
4.3 Features of complex adaptive systems 97
4.4 Boyd’s list of sources of uncertainty 104
4.5 A deconstructionist’s view on reality and cognition 111
4.6 The new planning paradigm as defined by the new sciences 116
4.7 Pentland’s set of potential centers of gravity 120
4.8 Pentland’s description of non-linear systems dynamics 121
4.9 List of CAS informed stratagems 126
6.1 Scene setter for command and control 191
6.2 Negatively valued features of linkages 192
6.3 Contrast of command and control, and appreciation and
leadership 200
6.4 Boyd’s view on isolation and interaction 214–15
6.5 The art of success 216
6.6 Examples from engineering (1) 221
6.7 Examples from engineering (2) 222
6.8 Examples from science 223
6.9 Sources of uncertainty 228
7.1 Domains of Unrestricted Warfare 254
Acknowledgments

This book took off during my stay at the School of Advanced Airpower Studies
in 1998–99. Thanks in particular to professor Dennis Drew, my mentor at SAAS
who said ‘I think you’re on to something’, thus providing me with the necessary
dose of self-confidence for embarking on the academic trail. The Clingendael
Institute of International Relations in The Hague has been my home from 2000
till 2005. A special word of appreciation for Professor Alfred van Staden, then
director of the Clingendael Institute, for maintaining faith in the project, for his
broad perspective, as well as for his patience. From the head of the research
department, Professor Jan Rood, I received more latitude to conduct research in
a wide variety of topics than I could have wished for. Professor Rob de Wijk,
director of the Clingendael Center for Strategic Studies, has assisted me
throughout the development of this book with frequent pointed remarks. I have
also sincerely benefited from the personal encouragement and support I received
from the senior leadership of the Royal Netherlands Air Force in the past
decade. I am deeply indebted to LtGen Droste, LtGen Berlijn, LtGen Starink,
MGen Hilderink, MGen Melker, and MGen Meulman. I am grateful also to
LtCol. Paul Ducheine and Col. Peter Wijninga, who have been brothers in intel-
lectual arms in the past years. I would like to thank Barry Watts, Chet Richards
and Dick Safranski for providing me with very valuable comments and sugges-
tions on various drafts of this book, based on their personal acquaintance with
John Boyd. Mary Holton-Boyd was very kind in permitting me to incorporate
slides from Boyd’s work, which greatly help in explaining his ideas. The final
word of gratitude goes to professor Grant Hammond, of the Center for Strategy
and Technology of the Air War College, and author of the first Boyd biography.
He has been the mental and moral driving force in the development of this book
since 1999. Indeed, I even owe the title of the book to him.
1 Introduction

To flourish and grow in a many-sided uncertain and ever changing world that
surrounds us, suggests that we have to make intuitive within ourselves those
many practices we need to meet the exigencies of that world. The contents that
comprise this ‘Discourse’ unfold observations and ideas that contribute towards
achieving or thwarting such an aim or purpose.
John Boyd, A Discourse, p. 1

Introducing A Discourse
This book aims to provide a better understanding of the strategic thought
developed by John Boyd. He has exerted a very substantial influence on recent
military thinking in the western world, and continues to do so. On the other
hand, his work has invited dismissive critique. Despite this situation, however,
there is as of yet no comprehensive study concerning his ideas.
Most people associate Boyd with the so called ‘OODA loop’, where ‘OODA’
is generally understood to stand for observation, orientation, decision and action.
The idea has gained currency that the OODA loop is the equivalent of a decision
cycle. Subsequently, war can be construed of as a collision of organizations
going through their respective OODA loops, or decision cycles. In the popular-
ized interpretation, the OODA loop suggests that success in war depends on the
ability to out-pace and out-think the opponent, or put differently, on the ability
to go through the OODA cycle more rapidly than the opponent. In simplified
form, it looks like Figure 1.1.
Few are familiar with the source of the OODA loop: A Discourse on Winning
and Losing. A Discourse consists of four briefings and an essay. The set has also
been labelled as The Green Book. It was completed in 1987, although subse-
quently frequently the specific wording on slides was revised. The essay
Destruction and Creation was written in 1976. It is a window to Boyd’s mind,
according to Robert Coram, one of his two biographers.1 In it Boyd states that
uncertainty is a fundamental and irresolvable characteristic of our lives, no
matter how good our observations and theories for explanation are.
Patterns of Conflict forms the historical heart of the work. First draft com-
pleted in 1977, it has turned into the opus of Boyd’s research on conflict and
2 Introduction

Action Observation

Decision Orientation

Figure 1.1 Simplified drawing of the OODA loop.

warfare, containing 193 slides. It is a historical analysis of warfare and theories

for victory and represents, in Boyd’s own words, ‘a compendium of ideas and
actions for winning and losing in a highly competitive world’.2 It also contains
an introduction to the OODA loop or the ‘Boyd Cycle’.
In the presentations Organic Design for Command and Control (first draft
1982) and the one intriguingly titled The Strategic Game of ? and ? (first draft
1986), he uses insights and conclusions from Patterns of Conflict but now in
abstract form. He employs these abstractions to develop arguments about leader-
ship and about the essence of strategy, or in Boyd’s own description: Organic
Design for Command and Control ‘surfaces the implicit arrangements that
permit cooperation in complex, competitive, fast moving situations’, while The
Strategic Game of ? and ? emphasizes ‘the mental twists and turns we undertake
to surface appropriate schemes or designs for realizing our aims or purposes’.3
The last very brief presentation, Revelation ‘makes visible the metaphorical
message that flows from this Discourse’.
In addition to these briefings he developed three pieces that need to be con-
sidered as well for a proper understanding of Boyd’s work. While working his
essay he also finished a presentation titled A New Conception of Air to Air
Combat, showing a close relation with the essay and foreshadowing several
ideas he was to explore in Patterns of Conflict. It is the conceptual bridge between
his fighter pilot background and his maturation as a strategic thinker. He also
developed two other briefings that are not an integral part of The Green Book but
are fully in line with, and an elaboration on previous arguments. The Conceptual
Spiral was completed in 1992. It is a different rendition of arguments, themes
and insights he advanced earlier in Destruction and Creation now employed to
 Introduction 3
explain how and why innovation occurs in science, engineering and technology.
Boyd argues that the dynamics at play here hold universal validity for all types of
organizations that strive to survive under conditions of fundamental and unavoid-
able uncertainty. The final briefing is titled The Essence of Winning and Losing,
which is a very condensed rendering of Boyd’s core ideas. Completed in 1995,
only in this short presentation does Boyd offer a picture of the OODA loop, and in
a much more elaborated rendition than shown in Figure 1.1.

‘A towering figure’
Some regard Boyd as the most important strategist of the twentieth century, or
even since Sun Tzu.4 James Burton claims that ‘A Discourse on Winning and
Losing will go down in history as the twentieth century’s most original thinking
in the military arts. No one, not even Karl von Clausewitz, Henri de Jomini, Sun
Tzu, or any of the past masters of military theory, shed as much light on the
mental and moral aspects of conflict as Boyd.’5 Colin Gray has ranked Boyd
among the outstanding general theorists of strategy of the twentieth century,
along with the likes of Bernard Brodie, Edward Luttwak, Basil Liddell Hart and
John Wylie, stating that

John Boyd deserves at least an honorable mention for his discovery of the
‘OODA loop’ . . . allegedly comprising a universal logic of conflict. . . .
Boyd’s loop can apply to the operational, strategic, and political levels of
war. . . . The OODA loop may appear too humble to merit categorization as
grand theory, but that is what it is. It has an elegant simplicity, an extensive
domain of applicability, and contains a high quality of insight about stra-
tegic essentials. . . .6

Boyd’s influence first became apparent during the late 1970s and 1980s in the
development of what later turned out to be the AirLand Battle concept.7 Later,
the US Marine Corps incorporated Boyd’s ideas into their new fighting manuals.
In the US Joint Chiefs of Staff Publication, JP 3–13.1, Joint doctrine for
Command and Control Warfare (C2W) the OODA loop is included in Appendix
A (without however mentioning Boyd’s name anywhere). Outside the US too,
his influence is demonstrable. For instance, the UK military doctrine description
of the doctrinally preferred method of war fighting, ‘the maneuvrist approach’ is
also pure Boyd (and fully in line with the US Marines doctrine):

The maneuvrist approach to operations is one in which shattering the

enemy’s overall cohesion and will to fight, rather than his materiel is para-
mount [. . .] significant features are momentum and tempo, which in combi-
nation lead to shock and surprise. Emphasis is on the defeat and disruption
of the enemy – by taking the initiative, and applying constant and unaccept-
able pressure at the times and places the enemy least expects – rather than
attempting to seize and hold ground for its own sake. It calls for an attitude
4 Introduction
of mind in which doing the unexpected and seeking originality is combined
with ruthless determination to succeed. A key characteristic of the maneu-
vrist approach is to attack the enemy commander’s decision process by
attempting to get inside his decision making cycle. This involves presenting
him with the need to make decisions at a faster rate than he can cope with,
so that he takes increasingly inappropriate action or none at all, thereby par-
alyzing his capability to react. Clearly any degradation of the overall
command system which can be achieved by physical or other means will
hasten the onset of paralysis.8

His influence extends into weapon systems and recent military operations.
The 1991 Gulf War air campaign employed F-16, F-18 and F-15 aircraft,
fighters that Boyd helped create during the 1960s and 1970s. The war itself is by
some considered a validation of the innovation in operational theory and praxis
that matured in AirLand Battle.9 In fact, Boyd has been credited with directly
influencing the design of the military ground campaign through his association
with Dick Cheney, then US Secretary of Defense, a former member of the so-
called Military Reform Group, who was well versed in Boyd’s military thinking.
In the May 6, 1991 issue of US News & World Report Boyd was mentioned,
together with two officers who were directly influenced by Boyd, as the persons
who determined the tactics employed during the Gulf War.10
In the aftermath of the terrorist attacks of September 11 2001, US Secretary
of State and former chairman of the US Joint Chiefs of Staff Colin Powell
implicitly honored Boyd by talking of a response involving multiple thrusts and
getting inside the adversary’s decision cycle.11 In the 1990s his ideas were incor-
porated in various European military doctrines, while the concept of Network
Centric Warfare, a key theme of US and NATO military transformation initi-
atives ongoing since 2002, surfaces many themes central to Boyd’s work.
Looking back on the stunning victory of Operation Iraqi Freedom (2003) against
substantial Iraqi armed forces, the commander of the coalition troops, General
Tommy Franks also referred explicitly to Boyd’s idea of getting inside the
enemy’s decision cycle.12 This indicates that Boyd’s concepts and terminology
have become mainstream in the Western militaries, and will also be employed
for the security challenges of the twenty-first century.13
And his fame has not been confined to military strategy. The OODA Loop
has even been discussed in Forbes and Harvard Business Review.14 In fact,
Figure 1.1 and a discussion of the OODA loop appear in Competing Against
Time, a book on management.15 In addition, Tom Peters, author of Thriving on
Chaos, a book that revolutionized management theories in America, talks of cre-
ating and exploiting chaos, of shaping the marketplace and of mutual trust.
Peters admitted that his book had been shaped by Boyd’s ideas. Since then
Boyd’s ideas have been applied by consultants and have been taught at business
schools, with the active endorsement of Boyd, who considered this an affirma-
tion of the fact that his intellectual legacy encompassed more than war fighting;
his ideas were universal, timeless, and could be applied to any form of conflict.16
 Introduction 5
The tribute written two days after Boyd’s death by General C.C. Krulak, then
Commandant of the US Marine Corps, reflects the view shared by many that
Boyd was

a towering intellect who made unsurpassed contributions to the American art

of war. Indeed, he was one of the central architects in the reform of military
thought which swept the services, and in particular the Marine Corps, in the
1980’s. From John Boyd we learned about the competitive decision making
on the battlefield – compressing time, using time as an ally. Thousands of
officers in all of the services knew John Boyd by his work on what was to be
known as the Boyd Cycle or OODA loop. His writings and his lectures had a
fundamental impact on the curriculum of virtually every professional mili-
tary education program in the United States – and many abroad [. . .] he was
the quintessential soldier-scholar – a man whose jovial outgoing exterior
belied the vastness of his knowledge and the power of his intellect.17

‘Deeply flawed’
Controversy and misperception nevertheless surround Boyd’s work. Like
Clausewitz and Sun Tzu, his work is more heard of than read or understood.
Very few people have actually worked their way through the presentations.
Instead, the neat graphical depiction of the OODA loop has become the symbol
of Boyd’s entire work, indeed, it is often regarded as the concise representation
of his ideas. One expert, for instance, asserts that ‘Boyd’s theory claims that the
key advantage to success in conflict is to operate inside the opponent’s decision
cycle. Advantages in observation and orientation enable a tempo in decision
making and execution that outpaces the ability of the foe to react effectively in
time.’18 And in 1996 this interpretation of the OODA loop was incorporated in,
and elevated to, long-term US defense policy when the Joint Chiefs of Staff
1996 document Joint Vision 2010 stated that US Forces will gain ‘OODA-loop
dominance’, being able to ‘observe, orient, decide, and act much more quickly
than our opponents’.19
This particular view on Boyd’s work has inspired critique. To be sure, some
offer assessments based on a somewhat wider appreciation of Boyd. Colin Gray
regards the ideas of Boyd as constituting a general theory of conflict. Others
discuss his ideas in particular within the framework of operational level doctrine
of warfare, regarding his work as the conceptual foundation of maneuver
warfare.20 Some others dismiss Boyd’s ideas on the grounds that they are rather
underdeveloped and too theoretical, pointing to the fact that, unlike Baron de
Jomini or, more recently, air-power theorist John Warden, Boyd’s notions
remain too vague to amount to anything other than a moving target of little use
in structuring a debate or attempting to educate one’s mind on the nature of war
before arriving at the battlefield.21
But frequently, discussions concerning the merits of Boyd’s work focus
solely on the merits of the OODA cycle idea, with one school suggesting that
Exploring the Variety of Random
Documents with Different Content
The Project Gutenberg eBook of Physiology
and histology of the Cubomedusæ
This ebook is for the use of anyone anywhere in the United States
and most other parts of the world at no cost and with almost no
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Title: Physiology and histology of the Cubomedusæ

Author: E. W. Berger

Contributor: Franklin Story Conant

Release date: March 3, 2017 [eBook #54276]

Most recently updated: October 23, 2024

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*** START OF THE PROJECT GUTENBERG EBOOK PHYSIOLOGY

AND HISTOLOGY OF THE CUBOMEDUSÆ ***
Transcriber’s Note: The images contained within black borders are clickable for a larger
version, if you are using a browser/device that supports this functionality.

Memoirs from the Biological Laboratory

OF THE
JOHNS HOPKINS UNIVERSITY
IV, 4
WILLIAM K. BROOKS, EDITOR

PHYSIOLOGY AND
HISTOLOGY
OF
THE CUBOMEDUSÆ

INCLUDING
Dr. F. S. Conant’s Notes on the Physiology

A DISSERTATION PRESENTED TO THE BOARD OF UNIVERSITY

STUDIES OF THE JOHNS
HOPKINS UNIVERSITY FOR THE DEGREE OF DOCTOR OF
PHILOSOPHY

BY
E. W. BERGER
 BALTIMORE
The Johns Hopkins Press
1900

PRINTED BY
The Lord Baltimore Press
THE FRIEDENWALD COMPANY
BALTIMORE, MD., U.S.A.

This Memoir is a continuation of the work upon the Cubomedusæ

which was begun by the late Dr. Franklin Story Conant, and it
contains his notes of physiological experiments, as well as new
results which have been obtained by Dr. E. W. Berger from the study
of material which had been collected by Dr. Conant, who had hoped
to make it the object of further study.
In order that this work may be made public as a continuation of
Dr. Conant’s researches, his sister, Grace Wilbur Conant, has, with the
coöperation of other members of his family, made an adequate and
generous provision for its publication.
For this gift, which is at once a contribution to science and a
memorial of an able and promising investigator, lately student and
fellow in this institution, the Johns Hopkins University returns its
grateful acknowledgments.
DANIEL C. GILMAN, President.
W. K. BROOKS, Professor of Zoölogy.
 CONTENTS.
PAGE
INTRODUCTION.
History 1
Epitome of Anatomy 2
PHYSIOLOGICAL.
Charybdea.
Light and Darkness 5
Concretions 8
Sensory Clubs 9
Velarium and Frenula 11
Pedalia, Interradial Ganglia, Tentacles 12
Stomach, Suspensoria, Proboscis, Subumbrella 13
Margin, Radial Ganglia, Nerve 15
Stimulation 17
Activity of Charybdea 17
Temperature 17
Food and Feeding 18
Occurrence of Charybdea 18
Aurelia and Polyclonia (Cassiopœa) 19
Summary 22
DR. CONANT’S NOTES.
Charybdea.
Light and Darkness 24
Sensory Clubs 26
Nerve 29
Side, Subumbrella 30
Pedalia, Velarium, Ganglia 31
Tentacles 32
 Proboscis, Stomach, Phacelli 33
Temperature 33
Food and Feeding 33
Occurrence of Charybdea 33
Activity of Charybdea 34
Aurelia and Polyclonia 35
Cassiopœa 39
Aurelia 39
HISTOLOGICAL.
Method 40
Anatomy 41
Distal Complex Eye—
General 41
Cornea 42
The Lens 42
The Capsule 44
The Retina 45
(a) The Prism Cells 46
(b) The Pyramid Cells 48
(c) The Long Pigment Cells 50
(d) Subretinal Nerve Tissue 53
(e) Discussion of Literature 53
(f) Function of the Retinal Cells, Patten’s Theory, 56
and further Literature
The Proximal Complex Eye 60
The Simple Eyes 61
Lithocyst and Concretion 63
The Epithelium of the Clubs 64
Network and Multipolar Ganglion Cells 67
The Nerve Tissue 67
The Supporting Lamella 68
Epithelium of Ampulla and Floating Cells 68
The Endothelium of the Peduncle 73
 The Tentacles and Pedalia—
The Ectoderm 74
(a) Thread Cells 74
(b) Muscle Fibers 74
(c) Ganglion Cell 75
The Endoderm 75
Summary 77
LITERATURE 78
REFERENCE LETTERS 80
DESCRIPTION OF FIGURES 81
 INTRODUCTION.
This paper may be regarded as a continuation of the
Cubomedusan studies pursued by Dr. F. S. Conant while in Jamaica,
in 1896 and 1897, with the Johns Hopkins Marine Laboratory. His
systematic and anatomical results have since been published as his
Dissertation (“The Cubomedusæ”) by this University. Conant
described this paper as Part I, hoping soon to add a second part on
the physiology and the embryology, for which he had some notes
and material at hand. Returning, however, to Jamaica with the
laboratory, in 1897, he continued his physiological experiments, and
preserved material for histological purposes. Upon the untimely
death of Conant, his material and notes were placed in my hands by
Professor Brooks, to whom I here take the opportunity of expressing
my appreciation and sincere thanks for the honor thus conferred and
for the many favors received.
In this paper I shall note at some length Conant’s physiological
results and append his notes. I shall also add my results on the
histology of the eyes and the sensory clubs in general, with some
few facts on the histology of the tentacles. The embryology will be
reserved for a future paper.
The forms used in the physiological experiments were Charybdea
Xaymacana, one of the two species (see Literature V, a and b) first
found and described by Conant; Aurelia aurita; Polyclonia and
Cassiopœa. The greater number of Conant’s notes are on
Charybdea, and were left by him just as taken at the time of
experimenting. Many of these notes are highly interesting and in the
main fit in well with Romanes’[I] and Eimer’s[IV] results.
Dr. Conant’s work on Charybdea, in 1897, was wholly done at Port
Antonio, Jamaica. At first Conant had only varying success in
obtaining Charybdea, scouring the harbor and neighboring water at
all hours, only to obtain but few specimens. It was on the forenoon
of August 7th, while we were dredging at the head of East Harbor
with a steam launch, that many Charybdeæ were brought up in the
dredge. This gave Conant a clue to their whereabouts and to the
means of obtaining them, and from that time on he was able to
obtain them in abundance. His first physiological experiments were
begun on August 4th and continued thereafter at intervals of several
days until his departure from Jamaica on September 6th.
Dr. Conant usually performed his experiments during the second
half of the forenoon, after the animals had stood for a few hours in
the laboratory.
The building that was rented at Port Antonio for a laboratory had,
in the basement, a photographer’s dark-room, which was of great
service to Conant in his experiments.
The experiments on Aurelia, in 1897, were also performed at Port
Antonio, between August 6th and 9th. The experiments on
Cassiopœa were probably made at Port Antonio, where specimens
were occasionally obtained.
The notes on Aurelia and Polyclonia, in 1896, were taken at Port
Henderson, between May 12th and June 27th.
In his notes Conant speaks of Polyclonia and Cassiopœa. It is at
present undetermined whether he really had both forms or whether
he uses the two names for the same form. It seems likely that in
1896 he thought the form to be Polyclonia, while for some reason, in
1897, he supposed it to be Cassiopœa. I have examined several
specimens of these medusæ brought from Port Antonio and find that
they all have twelve marginal bodies and twenty-four radial canals,
according to which (V, Haeckel’s System), they should be Polyclonia.
Conant, however, speaks of removing sixteen marginal bodies, which
seems to indicate that he had Cassiopœa. A careful classification of
this form of medusæ found about Jamaica seems to be a
desideratum. I suppose, however, that for our purpose in this paper
it will make little difference which name is used, the two forms being
so similar in form and structure. I have, therefore, decided to retain
both the names used by Conant.
For the complete anatomy of Charybdea the reader is referred to
Dr. Conant’s dissertation, “The Cubomedusæ” (8b), or the Johns
Hopkins University Circulars (8a), both published by the Johns
Hopkins Press. But, for the convenience of those who may be less
familiar with Cubomedusan anatomy, the following brief summary of
the anatomy of Charybdea is given:
The Cubomedusæ, as the name implies, approximate cubes, with
their tentacles (four in Charybdea) arranged at the four corners of
the lower face of the cube. These tentacles are said to lie in the
interradii. Half way between any two points of attachment of the
pedalia (the basal portions of the tentacles) and a little above the
margin of the bell (cube), in a niche, hang the sensory clubs, one on
each side, four in all. Each sensory club hangs in a niche of the
exumbrella and is attached by a small peduncle whose axial canal is
in connection with one of the four stomach-pockets and in the club
proper forms an ampulla-like enlargement.
Each club is said to lie in a perradius, and, like the tentacles,
belongs to the subumbrella. This is shown by the course of the
vascular lamellæ, bands of cells that, stretching through the jelly
from the endoderm to the ectoderm all around the margin, form the
line of division between sub- and exumbrella.
Each club has six eyes. Two of these on the middle line of the club
facing inwards are called the proximal and distal complex eyes, to
distinguish them from the four simple eyes that are disposed
laterally, two on each side of the line of the two complex eyes. All of
these eyes look inwards into the bell cavity through a thin
transparent membrane of the subumbrella. Besides the eyes and the
ampulla already mentioned, a concretion fills the lowermost part of
the club, and a group of large cells, having a network-like structure
and called network cells by Conant, fill the uppermost part of the
club between the proximal complex eye and the attachment of the
club to its peduncle (Plate II, Fig. 13). What is evidently nerve
tissue, fibers and ganglion cells, fills the rest of the club, with two
groups of large ganglion cells disposed laterally from the network
cells. A sensory (flagellate) epithelium covers the club.
Most Cubomedusæ, among them Charybdea, have a velarium
(comparable to the velum of the Hydromedusæ), a membrane of
tissue that extends inwards at right angles all around the margin.
This velarium, like a velum, has a central opening through which the
water is expelled from the bell-cavity when the animal pulsates. In
the perradii and in the angle between the velarium and the body
wall, are the frenula, which give support to the velarium much like
brackets support a shelf, except that here the brackets are above the
shelf instead of below.
In the upper part of the bell is the stomach, with the phacelli in its
interradii, and continued ventrally into the manubrium, or the
proboscis. The cavity of the stomach is continued in the perradii
through the four gastric ostia into the four stomach pockets, which
occupy the sides of the bell and extend to the margin. Immediately
below the gastric ostia, and in the bell cavity, are the suspensoria,
one in each perradius. These support the floor of the stomach much
as the frenula support the velarium, except that the suspensoria are
placed under the shelf (to continue Conant’s figure) and not above it
as are the frenula.
A nerve ring, underneath the epithelium of the subumbrella,
passes from near the origin of each pedalium at the margin to the
origin of the peduncles of the sensory clubs, a little above the
margin, giving off a branch to each club. Eight ganglia are found in
the course of this nerve. The four pedal ganglia lie near the bases of
the pedalia, and are hence interradial; the four radial ganglia lie near
the bases of the peduncles of the clubs, and are perradial. A small
nerve, radial nerve, can be traced a short distance upwards from
each radial ganglion. Underlying the epithelium of the frenula and
the suspensoria are ganglion cells and nerve fibers in larger numbers
than elsewhere (excepting the ganglia mentioned) in the
subumbrella. Otherwise, ganglion cells and nerve fibers underlie the
epithelium of the subumbrella, including the inner surface of the
velarium, as also do muscle fibers, except in the perradii and in the
region of the nerve, where the latter become interrupted.
 PHYSIOLOGICAL.
Charybdea.

Light and Darkness—Experiments 1-9, 10, 33, 34.—As already

stated in the Introduction, a part of Conant’s experiments were
performed in a photographer’s dark-room, with the animals in a deep
glass jar. In the dark a fair proportion of the animals became nearly
quiescent on the bottom, but upon lighting a lamp many started up
immediately, while others took a longer time to come to the surface
and swim. These experiments were tried a number of times and on
different occasions with very similar results. Some medusæ,
however, tried immediately after being brought in, seemed not to
react so well upon being placed in the dark-room, nor would they
become quiescent. This, probably, was due to the fact that the
animals had not yet recovered from the effects of being caught and
placed in new surroundings. (Experiments 1, 2, 3.)
Other experiments (4-8, 33, 34) were tried by carrying the jar with
the animals from the weaker light of a room into the more intense
light of outdoors or into direct sunlight. The usual result was an
inhibition of pulsation and a settling to the bottom, while the
medusæ immediately became active again upon returning with them
to the room. These results were so marked that no doubts can be
entertained as to their cause, though some exceptions occurred in
which animals placed in the sun continued to swim on the surface or
soon recovered pulsation. In some experiments, too, no animals
responded to the inhibitory stimulus of the brighter light or all very
soon recovered. (See, however, Temperature.)
Reducing the light by placing a coat over the jar produced the
same effect in some experiments (8, 9, 10) as did reducing the light
in other ways, while removing the coat produced the same effect as
exposure to brighter light. In these instances it appears to be the
transition from weaker to stronger light that inhibits pulsation, rather
than the actual intensity of the light; and vice versa. It must be
noted, too, that when left for some time in any one place the
animals changed, some coming to the surface and others going to
the bottom.
These experiments show beyond doubt that Charybdea is sensitive
to light, and that it is moderate light that stimulates the animals to
activity, while darkness and strong light inhibit activity. While the
individual exceptions, as Conant himself suggests, are well explained
on the supposition of individual diversity, yet it appears that other
conditions, such as the time of day, temperature, etc., may have
been responsible for some of the exceptional experiments in which
no animals responded as expected.
While light of any intensity seems to have stimulated Romanes’[I]
Sarsia and Tiaropsis (Hydromedusæ) to activity, we note that it is
moderate light that stimulates Charybdea. This fact is evidently
correlated with the circumstance that Charybdea usually lives upon
or near the bottom.
It may further be added in regard to Romanes’ Tiaropsis
polydiademata, that when it was suddenly exposed to light it went
into a spasm preceded by a long latent period during which there
was a “summation of stimulating influence” in the ganglia. Sarsiæ
would congregate toward the source of light and in general were
more active in light than in the dark, while sudden darkness often
inhibited a swimming bout. Romanes proves for Sarsia that the
marginal bodies are the seat of luminous stimulation and that it is
the light rays and not heat rays that stimulate. He also remarks that
he has obtained similar results on the covered-eyed
(Scyphomedusæ) medusæ, namely, that they respond to luminous
stimulation.
It may here be of interest to note a few observations made by
myself at Wood’s Holl, Mass., on a beautiful Olindiad, which is
abundant in the Eelpond at the above place. I found that in a room,
in the ordinary light of evening, the animals swam actively; but the
moment the electric light was turned on they stopped swimming and
settled to the bottom or attached themselves to a branch of some
weed or stem suspended in the water. This was the result in every
trial. It is found, further, to be little active during the brighter parts
of the day, when one must dip quite deep with a net in order to
obtain it. A similar observation is also made by Murbach[II], who
further states that this medusa may be deceived into laying its eggs
by placing it in the dark.
One cannot help but remark how analogous is the behavior of
medusæ, in respect to light and darkness, to the behavior of many
of the higher animals,—and medusæ are among the most lowly
organized of the animal creation.
Were one to conclude from the behavior of Charybdea in light and
darkness in the laboratory, that it remained on or near the bottom in
the daytime but became more active near or at the surface
evenings, nights and early mornings, one would probably not be far
from the truth. Dr. Conant, while towing near the bottom with a
weighted net, in water four to five feet (1.2-1.5 m.) deep not far
from shore and deeper farther out, found Charybdea in abundance
mornings and afternoons, but very few in the evening. In the
evening some few were usually taken in the surface tow. (See
Introduction, Occurrence and Activity.)
Again, who knows but that Charybdea is active during the day, on
the bottom where it was dredged (the light there would only be
moderate), and quiet at night. This supposition would seem to be
true, at least, for those forms of Cubomedusæ that live in deep
water. We can hardly suppose that they should regularly rise to the
surface from great depths and become active. This much we do
know that bright light inhibits Charybdea’s activities, while it
probably would not be active in perfect darkness.
I do not know just what interpretation to put upon Conant’s
finding Charybdea at Port Henderson at the surface during the early
part of the forenoon, before the sea-breeze roughened the water
(“Cubomedusæ” p. 7). This fact hardly fits in with my conclusions
above. Perhaps Charybdea’s habits vary with its habitat.
Finally, while I find no experimental evidence in Conant’s notes
about what parts of Charybdea are sensitive to light, yet it would
seem preposterous, from histological evidence and from Romanes’
results on Sarsia, to doubt that the eyes of the marginal bodies are
the seat of this stimulation.
Dr. Conant further experimented by cutting off certain organs and
parts from the Cubomedusan bell. These excisions consisted chiefly
in cutting out the concretions of the sensory clubs, cutting off the
whole club, eliminating a part or whole of the margin and the
velarium, cutting the bell into sectors, excising the stomach and
parts connected with it, and other parts.

Concretions—Experiments 10, 11.—The four concretions were

removed from each of four animals. Two of these (Experiments 10,
and another (X), not appended, to save space) seemed to be little if
at all affected by the operation. One of the two (10) swam actively,
at first up and down more changeably than those intact, but later
mostly near the surface. The other one also swam actively and
showed nothing to indicate weakened sense-perception. The other
two (11) did not stand the operation well, as Conant remarks, and
immediately went to the bottom, where they remained, one
swimming, while eight hours later one was still in good condition.
Several attempts with stronger light by removing the coat from
the jar made no difference in the behavior of 10; it continued to
swim as heretofore. Upon a final trial, however, with removing the
coat, it went to the bottom, thus showing a possible reaction to
light; but when next seen it was keeping to the bottom.
That the concretions should function as organs of light sensation,
as the first of the above animals might seem to indicate, I believe is
out of the question.[a] The fact, too, that this same animal (10),
together with another (X), swam actively, immediately changing their
course upon coming to the surface, in reality behaving quite as
normal animals, hardly permits us to conclude from the behavior of
the other two (11) that the concretions function directly as organs of
equilibrium or space relations. May these concretions not function
simply as weights for keeping the sensory clubs with their eyes
properly suspended? Since these concretions lie at the lowermost
part of the clubs and in closed sacs and unsupported by cilia, it
would seem that the above suggestion as to their being weights is
not improbable. Direct observation (Experiment 20) by Conant
shows, furthermore, that the clubs always hang with a tendency for
the concretions to be lowermost, regardless of the position of the
animal.
Again, while they may function as weights, as just explained, the
fact that the epithelium of the clubs is flagellated (a flagellum,
continued as a nerve fiber, to each cell—see Histology), the
supposition lies near that these flagella are the ones influenced by
the concretions as the clubs bear against one side of the sensory
niche or the other. A somewhat similar view seems to be held by
other observers and is noted by Lang in his text-book (“The outer
epithelium of the auditory body carries the auditory hairs”). It
seems, then, that in functioning as weights for suspending the clubs,
they may also serve at the same time for making the pressure of the
club against the niche greater than if they were absent, and thus in
part serve in equilibrium. On this supposition we should expect,
furthermore, that after the removal of the concretions the animal
would be little, if at all, affected, since the clubs themselves, without
the concretions, would still be of sufficient weight to be influenced
by gravity and thus to bear against the walls of the sensory niche. It
must be noted, however, that Conant’s experiments upon
equilibration in Charybdea are negative. Also, that Charybdea has
any auditory sense is negatived by two attempts of Conant’s with a
violin—one attempt with the violin near the animals, and another
with it in contact with the dish. (From an unpublished note.) Hence,
some other word such as sensory or equilibrating should perhaps be
substituted for “auditory” in the above quotation.
Removing the concretions from Aurelia gave negative results very
similar to those on Charybdea. (Experiment 42.)

Sensory Clubs—Experiments 12-19, 20, 24.—The entire sensory

clubs were removed from a number of animals. A paralysis of
pulsation followed by a rapid recovery was the usual result. In some
instances, however, there was no paralysis, while in others no
recovery followed paralysis. This is true in a general way whether
one club only or all were removed. While no permanent paralysis
followed the removal of one or two clubs, yet permanent paralysis
did occur after the removal of a third club, as, of course, also after
the removal of a fourth. It is evident, too, that as the removal of the
clubs progressed recovery seemed to be weaker after each cutting,
except in one case when pulsation seemed to be quickened after the
removal of a second club. The pulsations after recovery seemed to
be not so strong and regular, often quite feeble, and in one instance
in groups. Pieces of tissue with a club attached and pulsating
regularly, ceased pulsating after removal of the club, in one instance,
however, still giving occasional contractions.
These results are quite the same as those of Romanes[I] on
Aurelia, Cyanæa, etc., and of Eimer[IV] on Aurelia, Rhizostoma,
Cotylorhyza, etc.[b] In these forms Romanes sometimes obtained
complete paralysis after the removal of the sensory clubs only, as
also after the removal of the whole margin, though this was not
marked in Aurelia. In Cyanæa and other forms motor centers
seemed to be more abundant than in Aurelia, so that paralysis was
oftener followed by recovery. He concludes that while the principal
motor centers reside in the lithocysts, other centers doubtless exist
that may function vicariously, but that the centers of the margin are
more definitely limited to the marginal bodies in the Scyphomedusæ
than in the Hydromedusæ, in which the whole margin seems to be
replete with centers. He feels positive, furthermore, that no motor
centers exist in Aurelia’s margin outside of the marginal bodies
(lithocysts). Eimer’s results are essentially the same as Romanes’, so
that for a more detailed comparison of the two, Romanes’ works
should be consulted.
Romanes’ conclusion for the Hydromedusæ is that the motor
centers are not so definitely localized in the marginal bodies, but in
the margin generally, the excision of the marginal bodies alone
producing only partial paralysis, as would also the removal of the
margin from between the marginal bodies, but not so marked. For
the Hydromedusæ he concludes, then, that all the centers of
spontaneity are definitely localized in the margin, but not limited to
the marginal bodies. To this he mentions one exception, namely,
Staurophora laciniata, in which another center is found near the
margin and two others in two opposite arms of the proboscis.
I made the remark in an abstract (VI) on Conant’s notes that
Romanes did not obtain recovery of pulsation after removal of all the
lithocysts in Aurelia. As noted above, he did obtain recovery, so that
Conant’s results on Charybdea and also Aurelia (see Polyclonia and
Aurelia) are quite in agreement with Romanes.
The paralysis following the removal of the clubs in Charybdea is
evidently, primarily, the result of a loss of a part of its nervous
mechanism (motor centers), and, secondarily, of nervous shock, and
points to the existence of a definite nervous mechanism in the clubs.
The histological evidence is here, as usual, corroborative of the
physiological.
Another interesting phenomenon observed after the removal of
one or all of the clubs was the strange behavior of the proboscis.
This would reach from side to side, expanding and contracting its
lips as if trying to grasp something. This behavior is very similar to
that of the proboscis of Tiaropsis indicans when Romanes stimulated
any part of its subumbrella, or of Limnocodium sorbii, a little fresh-
water medusa, when he stimulated its margin or the region of the
radial canals. (Ib., p. 242.)
 I may add that I observed a very similar movement of the
proboscis of the Olindiad, before mentioned. When I pulled off
pieces of its gonads by means of quick jerks, with a small forceps, it
would continually reach toward the injured part of its subumbrella.
This medusa is generally quite active with its proboscis and can
occasionally be seen to reach with it.
Romanes states in one place that the proboscis is not affected by
the excision of the margin. This is evidently not the case in
Charybdea, in which excision of the sensory clubs (which really
belong to the margin—see “Cubomedusæ”) decidedly stimulated the
proboscis to active movements. This, furthermore, points to the
marginal bodies as being organs of considerable importance in giving
information in the life of Charybdea. In Romanes’ Sarsia and other
medusæ, however, the proboscis did respond to the stimulation of
the tentacles and the marginal bodies, as also would the bell
respond to a stimulation of the proboscis (manubrium), thus
showing a reflex nervous connection between these regions of the
bell, similar to that described for Charybdea.

Velarium and Frenula—Experiments 18, 29, 30, 41c.—“The power

of originating contractions” to use Conant’s own words, “evidently
resides in the velarium or in ganglion cells of the frenula, just as it
does in the proboscis and the floor of the stomach.” Isolated pieces
of the velarium contracted by themselves as did the whole velarium
when all other tissue had been removed. An isolated velarium with
the margin and the pedalia attached gave irregular contractions.
When the pedalia with the interradial ganglia were removed it still
contracted; and when all the other tissue was cut off contractions
continued.
Cutting the velarium caused the pedalia to be strongly contracted
inwards so that the tentacles were brought inside the bell. Cutting
away the velarium did not interfere with the pulsations of the bell,
but progress was much retarded.
 Cutting the frenula caused the pedalia to contract but seemed not
to affect the ability to swim. Comparing the velarium of the
Cubomedusæ with the velum of the Hydromedusæ, I recall no
observations similar to the ones here noted, though it seems that
the two may have quite similar functions. It seems somewhat
probable that the velum, and also the velarium, may function in
obtaining food,—and this besides their function in swimming. Their
probable function in swimming, as is well known, is evidently to
narrow the mouth of the bell and thus to cause the water to be
forced out in a smaller but more rapid stream, giving the animal a
steady and more prolonged movement through the water at every
contraction of the bell. In regard to taking food, I observed that a
small crustacean, in the process of being swallowed by an Olindiad,
seemed to be held by the velum being firmly contracted about it
while the proboscis was working itself over the crustacean. It would
seem, furthermore, that my supposition is supported for Charybdea
by the fact that the pedalia and tentacles were contracted so as to
be brought inside the bell when the velarium was cut. The stimulus
of cutting the velarium may be comparable to a stimulus from some
object touching it, and thus cause the pedalia and tentacles to come
reflexly to aid in capturing or holding the object, a fish, crustacean,
or such, to be captured.

Pedalia, Interradial Ganglia, Tentacles—Experiments 15, 23, 27-31,

41b.—When the pedalia were removed, the power of the animal to
guide itself was completely gone. When one pedalium was cut the
others contracted, while stroking the outer edge of the pedalia,
touching the sensory clubs, or sharply pricking the subumbrella,
often produced the same result. (See also Nerve.) The upper part of
the subumbrella seemed not so sensitive and more seldom produced
the reflex of the pedalia, while the base of the stomach did not give
it at all. Stroking the outer edge of the pedalia of Tripedalia
cystophora, the second of the two species of Cubomedusæ
described by Conant, also caused the pedalia to be contracted
inwards. I may note here that the muscle fibers under the ectoderm
of the pedalia are specially well developed at and near the inner and
outer edges, both in Charybdea and Tripedalia. On the flattened
sides of the pedalia the muscle fibers are fewer.
When the pedalia were cut off far enough up to remove the
interradial ganglia, coördination was not affected and the animal
could pulsate well enough but with little progress. (See above under
Velarium and Frenula.)
An isolated tentacle is capable of squirming contractions, and
when stimulated at either end, it would contract wholly or in part
only.
The pedalia, then, it would seem, serve also as a steering
apparatus, for which they are admirably fitted, considering their
blade-like thinness.
Considering, now, the reflexes noted under this head and the
preceding one, we find that there is an intimate nervous connection
between the velarium and frenula, subumbrella, sensory clubs,
nerve, and a single pedalium, on the one hand, and the pedalia on
the other hand. This is born out fully, furthermore, by the
histological evidence—(See Introduction and “Cubomedusæ”).
Considering the subumbral plexus of ganglion cells and fibers,
including the velarium and the frenula, which is in connection with
the nerve ring and this again with the sensory clubs and the
interradial ganglia at the bases of the pedalia, we have a basis for
these reflexes. While Conant failed to demonstrate nerves
(“Cubomedusæ”) from the interradial ganglia to the pedalia, yet,
that a nervous connection exists between the pedalia and the bell is
well shown by his physiological experiments. I have, furthermore,
demonstrated ganglion cells under the ectoderm of the tentacles
(see Histology).
Romanes obtained quite similar results in the Hydromedusæ. He
found that when a tentacle of Sarsia was slightly stimulated, it alone
would contract, but when it was more strongly stimulated the other
tentacles also would respond as also the manubrium. I find no
evidence in Conant’s notes of any such response of the manubrium
of Charybdea, except when the clubs were cut off.
The reflex obtained on stimulating the subumbrella of Charybdea,
when the pedalia would contract, is somewhat different from that
obtained by Romanes, who found that the most sensitive part of the
subumbrella in producing a reflex of the margin was at the junction
of the manubrium to the bell and that the subumbrella below this
point did not give the reflex.

Stomach, Suspensoria, Proboscis, Subumbrella—Experiments 12,

18, 19, 24-26, 29, 31.—The proboscis and the stomach with the
phacelli when cut out, contracted with or without the lips removed.
The isolated lips also contracted (twitched).
Pieces of the sides connected only with the stomach and
suspensoria, or with the margin (Experiment 47 (?)) twitched
spontaneously, but seldom did so when these were removed. In one
instance the whole side was cut out so as to exclude the radial
ganglion but still connected with a portion of the suspensorium. This
pulsated, or contracted, but on being halved transversely, the lower
half ceased to contract while the upper half connected with the
suspensorium, continued to contract.
Cutting off the whole stomach end of the animal excited to very
rapid pulsations of the remaining part, with the stream of water
stronger out the aboral end than past the velarium.
Conant says, “It seems I get no good evidence of the subumbrella
without connection with special nerve centers being able to contract
by itself.” The piece in which he did get contractions he suspects
may have been intimately associated with some part of the frenula
or the suspensoria. In Polyclonia no such doubt exists, for small
pieces of subumbrella were seen to contract. A small piece of
subumbrella of Charybdea with a sensory club attached could
contract by itself.
 From the above it would seem that a center capable of inciting to
contractions resided in the suspensoria as well as in the sensory
clubs, and this may be one of the centers that becomes potent upon
the removal of the clubs. This is further supported by Conant’s
observation (Introduction and “Cubomedusæ”) that an extra large
number of ganglion cells is found under the epithelium of the
suspensoria. A somewhat similarly located center of spontaneity
described by Romanes for Staurophora laciniata (Hydromedusa) has
already been noted.
As to the rapid pulsations of the bell after cutting out the stomach
end, this also is similar to Romanes’ results on Aurelia and other
Scyphomedusæ, when he cut off parts of the manubrium or an
aboral ring out of the bell. In these instances, however, Romanes
soon obtained a slackening of the rhythm following the temporary
acceleration. The temporary acceleration he attributes to the
stimulus of cutting, and the slackening to a lack of some afferent
stimulus from the removed tissue. Conant obtained the same results
on Polyclonia by removing the oral arms (see Polyclonia) but says
nothing about a slackening of the rhythm in Charybdea. I believe the
increased rhythm in Charybdea was in part due to the decreased
amount of labor necessary to force the water out of two openings
instead of one, namely, past the velarium. Just how much this
observation bears upon Romanes’ theory of rhythmic contraction,
that the rhythm is due to an alternate exhaustion and recovery of
the contractile tissue, as opposed to the ganglionic theory of rhythm
of physiologists, one does not wish to speculate much. Yet, I feel
that the observation rather supports this theory. The tissue having to
do less work, would become less exhausted at each contraction and
require less time for recovery and hence have a more rapid rhythm.
I here sum up Romanes’ theory in a few words. The ganglia
liberate a constant and comparatively weak stimulus, one perhaps
about minimal. This stimulus sets off the contractile tissue; but as
the tissue contracts and becomes exhausted the constant stimulus
becomes, in relation to it, sub-minimal, and it does not contract
again until it has recovered and the stimulus is again strong enough
to set it off. The ganglionic theory of rhythmic contraction supposes
that the ganglia liberate stimuli to the contractile tissue at successive
intervals. Romanes had this theory suggested to him by the rhythmic
contractions he succeeded in obtaining by subjecting deganglionated
bells to a continuous but weak faradic stimulus, or by placing them
into weakly acidulated water, or into 5 per cent. glycerine. Romanes
claims that his theory better explains muscular tonus and the
contraction of involuntary muscle. He does not, however, hold this
theory to the exclusion of the ganglionic theory, since only too often
does he speak in terms of the latter. He further brings in his support
the fact that the frog’s tongue, in which no ganglia have been
demonstrated, can be made to contract rhythmically when subjected
to a weak and continuous stimulus. He also calls attention to the
rhythmic contractions seen in the Protozoa, the snail’s heart, etc.
Finally, physiologists are much inclined to explain the rhythmic
contraction of the heart and other involuntary muscles, in part, at
least, as due to a property of the contractile tissue.

Margin, Radial Ganglia, Nerve—Experiments 18, 21-23, 30.—

Complete removal of the margin did not stop pulsation; but the
removal of the radial ganglia stopped it permanently. While this
experiment seems to have been tried only once, yet, taking into
consideration the results of other operations, it would seem that the
principal centers of spontaneity reside in these ganglia. (It should
here be remembered that the interradial ganglia were probably
removed at the removing of the margin.)
Cutting the nerve in the eight adradii caused the pedalia to bend
inwards at right angles to their normal position but did not in the
least affect the coördination of the sides. When, however, the sides
were cut in the eight adradii to the base of the stomach,
coördination for the main part ceased, and each side pulsated in its
own rhythm.
I have said that the principal centers of spontaneity reside in the
radial ganglia. Upon further thought this hardly seems warranted. No
doubt, among the principal motor centers must be placed the
ganglionic masses of the clubs, and the radial ganglia, together with
the homologous interradial ganglia, represent centers of equal value.
I speak of these two sets of ganglia as homologous, since strictly
speaking, they both belong to the margin, and the clubs at whose
bases they lie probably represent modified tentacles. Conant’s
experiments leave us in the dark as to the function of these ganglia.
Next in order, it would seem, are the ganglion cells in the
suspensoria, as is suggested by the contractions of an isolated side
with a portion of a suspensorium attached. (See previous head.)
While we have seen that the frenula and the velarium can contract
by themselves, yet, I find no evidence that these can impart their
contractions to any adjacent tissue.
Conant’s results on cutting the nerve eight times and then
continuing the cuts to the base of the stomach are quite the same as
Romanes and Eimer obtained upon Aurelia. Romanes, however,
concludes that in his Sarsia, Tiaropsis, etc., coördination was broken
when only short incisions were made in the margin. Charybdea
appears, then, to agree with Aurelia rather than with the
Hydromedusæ. Yet, since Romanes at first obtained similar results to
those of Charybdea on Sarsia, but on further experimenting
concluded that coördination had really been destroyed at the first
cutting, we cannot speak with certainty that coördination had not
been destroyed in Charybdea before the cuts had been continued to
the base of the stomach. I say not with certainty, because the injury
to the bell being slight, coördination may have been maintained on
the principle of a simultaneously (simultaneous for the octants)
alternate exhaustion and recovery of the contractile tissue on the
principle of Romanes’ theory.

Stimulation.—Romanes found when he stimulated a

deganglionated bell of a Hydromedusa, that it responded by a single
contraction, while that of a Scyphomedusa responded with several
quite rhythmic contractions. Charybdea in this respect agrees with
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