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Functional Discourse Grammar
This page intentionally left blank
Functional
Discourse Grammar

A typologically-based theory of
language structure

K E E S H E N G EV E L D A N D
J. LACHLAN MACKENZIE

1
3
Great Clarendon Street, Oxford ox2 6dp
Oxford University Press is a department of the University of Oxford.
It furthers the University’s objective of excellence in research, scholarship,
and education by publishing worldwide in
Oxford New York
Auckland Cape Town Dar es Salaam Hong Kong Karachi
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With offices in
Argentina Austria Brazil Chile Czech Republic France Greece
Guatemala Hungary Italy Japan Poland Portugal Singapore
South Korea Switzerland Thailand Turkey Ukraine Vietnam
Oxford is a registered trade mark of Oxford University Press
in the UK and in certain other countries
Published in the United States
by Oxford University Press Inc., New York
© Kees Hengeveld and J. Lachlan Mackenzie 2008
The moral rights of the authors have been asserted
Database right Oxford University Press (maker)
First published 2008
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
without the prior permission in writing of Oxford University Press,
or as expressly permitted by law, or under terms agreed with the appropriate
reprographics rights organization. Enquiries concerning reproduction
outside the scope of the above should be sent to the Rights Department,
Oxford University Press, at the address above
You must not circulate this book in any other binding or cover
and you must impose the same condition on any acquirer
British Library Cataloguing in Publication Data
Data available
Library of Congress Cataloging in Publication Data
Data available
Typeset by SPI Publisher Services, Pondicherry, India
Printed in Great Britain
on acid-free paper by
CPI Antony Rowe, Chippenham, Wiltshire
_________________
ISBN 978–0–19–927810–7 (Hbk.)
ISBN 978–0–19–927811–4 (Pbk.)

1 3 5 7 9 10 8 6 4 2
 Contents

Preface xi
List of tables and figures xiv
Abbreviations xvi

1 Introduction 1
1.1 Functional Discourse Grammar 1
1.2 Basic properties 1
1.2.1 Introduction 1
1.2.2 Top-down organization 1
1.2.3 Discourse grammar 3
1.2.4 Levels of representation 4
1.2.5 Conceptual Component, Contextual Component,
and Output Component 6
1.3 The architecture of FDG 12
1.3.1 Overall organization 12
1.3.2 Levels and Layers 14
1.3.3 Primitives 19
1.3.4 Levels and primitives 22
1.3.5 Implementation 23
1.4 FDG in its broader context 25
1.4.1 Introduction 25
1.4.2 Functionalism 26
1.4.3 Typology 31
1.4.4 Language modelling 37
1.4.5 On using FDG 41
1.5 Notational conventions 43
1.6 Structure of the book 45
2 The Interpersonal Level 46
2.1 Introduction 46
2.2 The organization of the Interpersonal Level 48
2.3 The Move 50
2.3.1 Introduction 50
2.3.2 Heads 52
vi contents

2.3.3 Modifiers 58
2.3.4 Operators 59
2.3.5 Frames 60
2.4 The Discourse Act 60
2.4.1 Introduction 60
2.4.2 Heads 63
2.4.3 Modifiers 64
2.4.4 Operators 65
2.4.5 Frames 68
2.5 Illocution 68
2.5.1 Introduction 68
2.5.2 Heads 69
2.5.3 Modifiers 81
2.5.4 Operators 83
2.5.5 Frames 83
2.6 The Participants 84
2.6.1 Introduction 84
2.6.2 Heads 84
2.6.3 Modifiers 85
2.6.4 Operators 85
2.6.5 Frames 87
2.7 The Communicated Content 87
2.7.1 Introduction 87
2.7.2 Heads 88
2.7.3 Modifiers 102
2.7.4 Operators 104
2.7.5 Frames 106
2.8 Subacts 107
2.8.1 Introduction 107
2.8.2 Ascription 108
2.8.3 Reference 113
2.9 Building up the Interpersonal Level 124
3 The Representational Level 128
3.1 Introduction: semantics in FDG 128
3.2 The organization of the Representational Level 130
3.2.1 Semantic categories 130
3.2.2 Subclasses of semantic categories 135
3.2.3 The structure of representational layers 138
3.3 Propositional contents 144
3.3.1 Introduction 144
 contents vii

3.3.2 Heads 145

3.3.3 Modifiers 151
3.3.4 Operators 153
3.3.5 Frames 156
3.4 Episodes 157
3.4.1 Introduction 157
3.4.2 Heads 160
3.4.3 Modifiers 162
3.4.4 Operators 163
3.4.5 Frames 165
3.5 States-of-Affairs 166
3.5.1 Introduction 166
3.5.2 Heads 166
3.5.3 Modifiers 171
3.5.4 Operators 172
3.5.5 Frames 180
3.6 Configurational Properties 181
3.6.1 Introduction 181
3.6.2 Heads 182
3.6.3 Modifiers 208
3.6.4 Operators 210
3.6.5 Frames 214
3.7 Lexical Properties 215
3.7.1 Introduction 215
3.7.2 Heads 215
3.7.3 Modifiers 230
3.7.4 Operators 233
3.7.5 Frames 236
3.8 Individuals 236
3.8.1 Introduction 236
3.8.2 Heads 237
3.8.3 Modifiers 241
3.8.4 Operators 245
3.8.5 Frames 247
3.9 Location 248
3.9.1 Introduction 248
3.9.2 Heads 249
3.9.3 Modifiers 254
3.9.4 Operators 256
3.9.5 Frames 257
viii contents

3.10 Time 257

3.10.1 Introduction 257
3.10.2 Heads 258
3.10.3 Modifiers 260
3.10.4 Operators 262
3.10.5 Frames 262
3.11 Manner 263
3.11.1 Introduction 263
3.11.2 Heads 265
3.11.3 Modifiers 267
3.11.4 Operators 267
3.11.5 Frames 268
3.12 Quantity 268
3.12.1 Introduction 268
3.12.2 Heads 269
3.12.3 Modifiers 271
3.12.4 Operators 271
3.12.5 Frames 272
3.13 Reason 272
3.13.1 Introduction 272
3.13.2 Heads 272
3.13.3 Modifiers 274
3.13.4 Operators 274
3.13.5 Frames 274
3.14 Reflexive language 275
3.15 Building up the Representational Level 277
4 The Morphosyntactic Level 282
4.1 Introduction 282
4.1.1 Purpose and scope of the chapter 282
4.1.2 Relation to Interpersonal Level and
the Representational Level 283
4.1.3 Relation to Phonological Level 287
4.2 The organization of the Morphosyntactic Level 291
4.2.1 Introduction 291
4.2.2 Hierarchical structure 291
4.2.3 Grammatical morphemes and secondary operators 301
4.3 Linguistic Expressions 308
4.4 Clauses 309
4.4.1 Introduction 309
4.4.2 Ordering of hierarchically related units 311
 contents ix

4.4.3 Alignment 316

4.4.4 Ordering of non-hierarchically related units 333
4.4.5 Templates 341
4.4.6 Dummies and support 347
4.4.7 Agreement 350
4.4.8 Subordination 352
4.4.9 Coreference 372
4.5 Phrases 376
4.5.1 Introduction 376
4.5.2 Ordering of hierarchically related units 377
4.5.3 Alignment 383
4.5.4 Ordering of non-hierarchically related units 387
4.5.5 Templates 389
4.5.6 Dummies and support 391
4.5.7 Agreement 394
4.5.8 Subordination 396
4.6 Words 400
4.6.1 Introduction 400
4.6.2 Ordering of hierarchically related units 405
4.6.3 Alignment 406
4.6.4 Ordering of non-hierarchically related units 409
4.6.5 Templates 411
4.6.6 Dummies and support 412
4.6.7 Agreement 414
4.6.8 Subordination 414
4.7 Building up the Morphosyntactic Level 416
5 The Phonological Level 421
5.1 Introduction 421
5.2 The organization of the Phonological Level 427
5.3 Utterances 430
5.4 Intonational Phrases 432
5.5 Phonological Phrases 436
5.6 Phonological Words 443
5.7 Phonemes, Syllables, and Feet 449
5.8 Building up the Phonological Level 455

References 463
Language Index 489
Name Index 492
Subject Index 496
This page intentionally left blank
 Preface

At the Ninth International Conference on Functional Grammar (ICFG9), held

in Madrid in September 2000, Kees Hengeveld proposed the idea of a Func-
tional Discourse Grammar. At the time it was conceived of as a revised version
of Functional Grammar (FG; Dik 1997a, 1997b), a theory of the organization
of natural languages developed by the late Simon C. Dik and his colleagues
from 1978 onwards, the year in which the first book bearing the title Functional
Grammar appeared (Dik 1978). The addition of the word Discourse in the
denomination of the model was meant to reflect the awareness that the impact
of discourse features on linguistic form should be given greater prominence in
the theory. A number of the features of FDG as presented in this book were
already present in Hengeveld’s (2000) presentation, notably: the distinction
between an interpersonal, a representational, and a morphosyntactic level of
analysis, all of them with hierarchical layering; the actional status of ascription
and reference; the top-down operation of the model; the interaction of the
grammatical component with the conceptual and contextual components;
and the analysis of reflexive language use.
The ideas were taken up in lively discussion in the years following that
conference, not least at ICFG10 in Amsterdam (The Netherlands), ICFG11 in
Gijón (Spain), and ICFG12 in São João do Rio Preto (Brazil). This led to such
collections as Mackenzie and Gómez-González (2004, 2005), de Groot and
Hengeveld (2005), and García Velasco and Rijkhoff (2008), in which a range
of scholars from various countries contributed to the development of a new
architecture of the theory, with a strong desire to retain the best of FG while
increasing the scope and ambition of the model.
Just like FG, FDG seeks to reconcile the patent fact that languages are
structured complexes with the equally patent fact that they are adapted to
function as instruments of communication between human beings. FDG has
also inherited from its precursor the desire to achieve maximum typological
neutrality: the theory is designed to be equally applicable to languages of all
types, and indeed this book presents and analyses data from a very wide range
of languages, resorting to exemplifying from more familiar languages only
where the comprehensibility of the presentation makes this advisable.
However, FDG diverges from FG in so many ways that by now it should be
considered a theory in its own right, and it has been recognized as such, as evi-
denced by encyclopaedia entries such as Hengeveld and Mackenzie (2006, fc.)
and special issues of journals on FDG (van Staden and Keizer fc.; Hengeveld
xii preface

and Wanders fc.; Hattnher and Hengeveld 2007). FDG represents a significant
advance on FG in separating out the Interpersonal Level and the Represen-
tational Level and investigating the full complexity of the former as well as
the complex interaction between the two in determining linguistic form. It
also differs from its predecessor in regarding the Morphosyntactic Level and
the Phonological Level as more than mere expressions of the other Levels,
but as having their own principles of organization; these are fully elaborated
for the first time in this book. And finally, it differs crucially from FG in
being a top-down rather than a bottom-up model. All in all, then, FDG has
outgrown its intellectual origins and now offers an autonomous and balanced
account of the systematic impact of pragmatic, semantic, morphosyntactic,
and phonological phenomena on linguistic form.
The present book began life during a joint sabbatical of both authors
in Amsterdam in 2004. It then continued to grow in the form of e-mail
correspondence, with drafts being sent back and forth between Amsterdam
and Lisbon when other commitments permitted. In the last phase Mackenzie
was awarded a Visitor’s Scholarship from the Netherlands Organization for
Scientific Research (NWO) for the first months of 2007 under project number
B30-664, when he was able to work together with Hengeveld on a daily basis in
Amsterdam again. Mackenzie also wishes to acknowledge support received in
the early days from the Spanish Ministry of Education, the European Regional
Development Fund and the Xunta de Galicia under project number BFF2002-
02441 (PGIDIT03PXIC20403PN), and Hengeveld is grateful for support pro-
vided by the Amsterdam Center for Language and Communication and the
Department of Theoretical Linguistics of the University of Amsterdam.
We also wish to recognize the contribution of innumerable colleagues and
students who over the past years have contributed their ideas, encouragement,
and criticism to our enterprise of developing FDG. We were fortunate enough
to be given the opportunity to present FDG at conferences, in postgradu-
ate courses, and in guest lectures at many different places, and though we
cannot name all those who contributed with their questions, remarks, and
criticisms (for there have been so many), we would like to thank in gen-
eral terms our audiences at ICFG10 (Amsterdam, The Netherlands, 2002),
International Conference on Role and Reference Grammar (Logroño, Spain,
2002), LOT Winter School (Amsterdam, The Netherlands, 2003), Journées de
Linguistique Fonctionelle (Agadir, Morocco, 2003), ACLC/ILLC-Colloquium
(Amsterdam, The Netherlands, 2003), Københavns Universitet (Copenhagen,
Denmark, 2003), Århus Universitet (Århus, Denmark, 2003), Örebro Uni-
versitet and Södertörn Högskola (Stockholm, Sweden, 2003), Universidad
de Castilla La Mancha (Cuenca, Spain, 2003), Workshop on Grammar and
Discourse (Ghent, Belgium, 2003), Universidade Estadual Paulista (São José
 preface xiii

do Rio Preto, Brazil, 2003, 2004), SIL International Training Programme

(High Wycombe, UK, 2004), ESSE-7 (Zaragoza, Spain), 52o Seminário do
GEL (Campinas, Brazil, 2004), ICFG11 (Gijón, Spain, 2004), ICFG12 (São
José do Rio Preto, Brazil, 2006), Universidade Federal de Goiás (Goiânia,
Brazil, 2006), TWIST Student Conference (Leiden, 2007), Workshop on the
Representational Level in Functional Discourse Grammar (Zaandijk, 2007),
Università degli Studi del Molise (Campobasso, Italy, 2007), Universidade Fed-
eral do Rio de Janeiro (Rio de Janeiro, Brazil, 2007), Universidade Federal Flu-
minense (Niterói, Brazil, 2007), Moulay Ismail University (Meknès, Morocco,
2007) and Universiteit van Amsterdam (Amsterdam, The Netherlands, 2007).
Portions of this book were furthermore discussed at various occasions within
the context of the Functional Grammar Colloquium at the University of Ams-
terdam, and these discussions have led to considerable improvements of the
relevant parts. To all colleagues and students who participated in these events,
our deepest gratitude.
We are grateful to John Davey of Oxford University Press for his constant
support, his interest, and his advice.
Finally, we would like to give our special thanks to Inge Genee, Daniel
García Velasco, and Gerry Wanders, who read the entire pre-final manuscript
and generously gave us their detailed and invaluable comments. We hope they
will find their highly appreciated feedback reflected in the current book.
Kees Hengeveld
Amsterdam
J. Lachlan Mackenzie
Lisbon
 List of tables and figures

Tables
1. Semantic categories 131
2. Derived nominal expression of basic semantic categories 132
3. Semantic categories 2 136
4. Basic semantic functions 199
5. The Locatives of Avar 253
6. The constitution of a Linguistic Expression 309
7. Classification of complement Clauses 363
8. Correspondences between Lexical and Grammatical Word classes 401
9. Examples of correspondences between Grammatical Words and
elements of the Interpersonal Level and the Representational Level 402
10. Morpheme classes 404

Figures
1. FDG as part of a wider theory of verbal interaction 6
2. General layout of FDG 13
3. The Interpersonal Level 15
4. The Representational Level 15
5. The Morphosyntactic Level 17
6. The Phonological Level 18
7. Implicational relations between Illocutions 75
8. The representation of semantic functions 195
9. Flexible, differentiated, and rigid languages 227
10. Parts-of-speech systems 228
11. Nominative-Accusative alignment 325
12. Absolutive-Ergative alignment 325
13. Primative-Secundative alignment 327
14. Directive-Indirective alignment 327
15. Role-based alignment 328
16. Subjects in Nominative-Accusative/Directive-Indirective systems 328
 list of tables and figures xv

17. Subjects in Absolutive-Ergative/Directive-Indirective systems 328

18. Subjects and Objects in Nominative-Accusative/Primative-Secundative
systems 328
19. Subjects in Absolutive-Ergative/Primative-Secundative systems 329
20. Alignment within Phrases 387
21. Alignment of arguments with respect to incorporation
in Southern Tiwa 409
22. Alignment of arguments with respect to incorporation in Nivkh 410
 Abbreviations and symbols

Abbreviations used in glosses

1 first person assv associative
2 second person attr attributive
3 third person aug augmentative
a actor aux auxiliary
abil ability av actor voice
abl ablative avol avolitional
abs absolutive ben benefactive
acc accusative caus causative
acq acquired cert certain
actnr action nominalizer cl class
adh adhortative clf classifier
adjr adjectivalizer cntrl control
admon admonitive coll collective
advr adverbializer comm commissive
aff affected comp complementizer
ag agent compl completive
agr agreement compv comparative
all allative cond conditional
anaph anaphoric conj conjunction
anim animate conn connective
anr agent nominalizer cont continuative
ant anterior contg contingent
aor aorist contr contrastive
appl applicative cop copula
approx approximative corr correlative
art article cv conveyance voice
asp aspect cvb converb
ass assertive dat dative
 abbreviations and symbols xvii

decl declarative foc focus

def definite fut future
deict deictic gen genitive
dem demonstrative genr general tense
dep dependent ger gerund
det determiner hab habitual
detrans detransitivizer hest hesternal (happening
dim diminutive yesterday)
dir direction hod hodiernal (happening
dishort dishortative today)
disp dispensative hon honorific
distr distributive hort hortative
dox doxastic hum human
drct direct imm immediate
ds different subject imp imperative
du dual impf imperfect
dub dubitative impr imprecative
dum dummy inabil inability
dur durative inan inanimate
dyn dynamic inch inchoative
emph emphasis incl inclusive
erg ergative ind indicative mood
ergr ergativizer indf indefinite
ess essive indftns indefinite tense
ex existential indep independent
ex.prev previously existing iness inessive
exact exact ascription inf infinitive
excl exclusive infer inferential
exclam exclamative ingr ingressive
exct exactly ins instrument
f feminine int intentive
fact factual intens intensifier
fam familiar inter interrogative
fin finite interp interpellative
xviii abbreviations and symbols

intr intransitive nonperc non-perceived

intr.abil intrinsic ability nonpst non-past
inv inverse nonsbj non-subject
irr irrealis nontop non-topic
irrat irrational nonvis non-visual sensory
iter iterative evidence
itive itive nucl nucleus
ld locative direction obj object
lk linker objp objective paradigm
loc locative objresp object of respect
locnr locative nominalizer objv objective
log logophoric obl oblique
lv locative voice oblg obligation
m male, masculine obv obviative
mann manner opt optative
mann.nr manner nominalizer part particle
mir mirative partv partitive
mit mitigation pass passive
ms male speaker paus pause
n neuter pc paucal
narr narrative perc perceived
neg negative, negation perm permanent
newtop new topic pfv perfective
nf non-feminine pl plural
nfact non-factual plup pluperfect
nh non-human pm predicate marker
nml nominal pol polite
nmlz nominalizer pos positive
nom nominative poss possessive
non.a/s non-nominative post posterior
non.inter non-interrogative pot potential
nonattr non-attributive potv potentive voice
nonf non-finite pred predicate
nonfut non-future prep preposition
 abbreviations and symbols xix

pres presentative rls realis

presup presupposed s argument of
prf perfect intransitive verb
priv privative sbj subject
proc procrastinative sbjv subjunctive mood
prof profession seq sequence
sg singular
prog progressive
sgltv singulative
proh prohibitive
sim simultaneous
prone prone (to negative
characteristic) sit situational
propr proprietive spec specific
prox proximate srdir superdirective
prs present tense ss same subject
pst past tense stat stative

ptcp participle sub subordinator

subdir subdirective
punct punctual
subess subessive
q question marker
subl sublative
quot quotative
subs subsequent
realnr realis nominalizer
suppl supplicative
reasnr reason nominalizer
tam tense-aspect-mood
rec recipient
tmpnr temporal
recpst recent past nominalizer
red reduplication tns tense
refl reflexive top topic
refr referential tr transitivity marker
reinf reinforcement u undergoer
rel relativizer uv undergoer voice
relr relator val validator
rem remote vfin finite verb
rep reportative vis visible
repv repetitive vis.evid visual evidence
res resultative voc vocative
rev reversative vol volitive
xx abbreviations and symbols

Abbreviations used in representations

General h higher social status
⊂ is entailed by HORT hortative
ILL variable for an illocution
Interpersonal Level IMP imperative illocution

lexeme IMPR imprecative
[±A] ± involving the INTER interrogative
addressee INTERP interpellative
[±S] ± involving the speaker M1 move
± for ± formal MIR mirative
± id ± identifiable mit mitigation
±s ± specific Motiv motivation
A addressee OPT optative
A1 discourse act Orient orientation
ADMON admonitive P1 speech-act participant
approx approximative PROH prohibitive
Aside aside R1 subact of reference
C1 communicated rep reportative
content
S speaker
COMM commissive
SA subact
Conc concession
SUPPL supplicative
Cor corrective
T1 subact of ascription
DECL declarative
V1 any interpersonal
DISHORT dishortative variable
DISP dispensative Y/N yes-no
emph emphasis
operator
exact exact ascription
M
operator on a move
Expl explanation (etc.)
F1 illocution  modifier
Feedb feedback  M
modifier of a move (etc.)
Foc focus  function
H head  M
function of a move (etc.)
 abbreviations and symbols xxi

Representational Level magn magnitude

m

lexeme x1 mass individual
∀ universal quantifying p1 propositional content
operator past past
∃ existential quantifying pc paucal
operator perc perception
∅ zero p
f1 permanent property
1 singular post posterior
2 dual pres present
3 trial prog progressive
A actor prox proximal
Abl ablative q1 quantity
ant anterior r1 reason
C comitative Ref reference
c
f1 contingent property rem remote
Circ circumstance s
e1 stative state-of-affairs
Cons consequence sim simultaneous
c
x1 collective individual s
x1 set individual
d
e1 dynamic state-of-affairs t1 time
distr distributive Temp temporal
e1 state-of-affairs U undergoer
ep1 episode v1 any representational
Ess essive variable
f1 property x1 individual
h head  operator
hab habitual Û modifier
infer inferential Û f
modifier of property
Ins instrument Û l
modifier of location
In inessive Û m
modifier of manner
L locative Û t
modifier of a time
l1 location  function
m plural, more than one p function of a propositional
m1 manner content (etc.)
xxii abbreviations and symbols

Morphosyntactic Level PM+N position situated N

Aff1 affix places after the medial
position
Adp1 adpositional phrase M-N
P position situated N
Advp1 adverb phrase
places before the medial
Advw1 adverbial word position
Ap1 adjective phrase Ppost postclausal position
As1 adjectival stem P pre
preclausal position
Aw1 adjectival word Vp1 verb phrase
Cl1 clause Vr1 verbal root
bal
Cl1 balanced clause Vs1 verbal stem
dep
Cl1 dependent clause Vw1 verbal word
der
Cl1 deranked clause f
Vw1 finite verb(al word)
Gw1 grammatical word Xm1 morpheme (of type x)
Le1 linguistic expression Xp1 phrase (of type x)
Np1 noun phrase Xr1 root (of type x)
Nr1 nominal root Xs1 stem (of type x)
Ns1 nominal stem Xw1 word (of type x)
Nw1 nominal word
P 2
second position Phonological Level
P2+N position situated N CAP characteristic accent
places after the second position
position f falling
P centre
position of clause with f foot
respect to pre- and h high
postclausal positions
ip1 intonational phrase
PF final position
l low
PF-N position situated N
m mid
places before the final
position pp1 phonological phrase
PI initial position pw1 phonological word
PI+N position situated N r rising
places after the initial s stressed
position s1 syllable
PM medial position u1 utterance
 abbreviations and symbols xxiii

Parts-of-speech Pro pronoun

A adjective V verb
Ad adposition
Grammatical models
Adv adverb
FDG Functional Discourse
Cont contentive
Grammar
DAdv degree adverb
FG Functional Grammar
Det determiner
RRG Role and Reference
Intj interjection Grammar
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question presents itself as to the connections between vital process
and the effect of stimulation.
When we study the motile flagellate infusorium Peranema swimming
undisturbed in water, we observe that the swimming movements are
absolutely regular in character. The elongated cell body remains
unaltered in shape. The long flagellum is extended in a perfectly
straight line in the axis of the body and only the extreme end lashes
with regularity through the water (Figure 8, A). There is majestic
grace in this perfect uniformity of motion. The picture suddenly
alters the moment the Peranema is influenced by the slightest jar.
The whole flagellum at once executes a few violent movements
(Figure 8, B), the body draws together, soon stretches itself again
and swims immediately after, in another direction, with the same
majestic calm as before.
 Fig. 8.
Peranema. A—
Swimming in non-
stimulated
condition. B—
Mechanically
stimulated at the
end of the
flagellum.

Another instance. A number of fertilized eggs of the sea urchin are

placed in a watch glass in sea water. The temperature of the water
should correspond with the mean temperature in which the animals
live in the sea, averaging about 15° C. The eggs begin to form
grooves and to develop slowly by progressive division. In another
glass we observe a second sample of fertilized eggs of the same kind
and under the same conditions, but in this case we increase the
temperature to 25° C. The increased temperature brings about a
decided increase of segmentation and the same stage of
development is reached in less than half the time. The increased
temperature, therefore, increases the development. Further we take
a third sample of the same urchin eggs in a watch glass with sea
water of 15° C. and add a little sea water mixed with ether. The
development of the eggs now comes to a standstill. The narcotic has
produced an inhibition of development.
To quote another instance. Bacterium phosphorescens having been
bred upon a putrid fish are exposed in the culture fluid to the air. In
the dark the bacteria give forth a phosphorescent light. Then the
culture fluid containing the bacteria is put into a glass receptacle,
which can be rendered air-tight and all oxygen excluded. After a
short time the light formation ceases completely. The absence of
oxygen has here had a depressing effect and it is only after air has
been again introduced that light is once more produced.
Lastly, an example from the group of mammals may be cited. The
metabolism of a dog in complete rest is examined for a prolonged
length of time and we ascertain the values of the oxygen
consumption, the carbon dioxide production, and the nitrogen
elimination in the urine. Under the same nutritive conditions the
animal is then allowed to work from time to time in a treadmill.
During these working periods impulses of excitation are continually
conducted to the muscles from the nervous system. It is now found
that under the influence of the constantly recurring stimuli the
quantity of nitrogen in the urine has only very slightly augmented,
whereas the consumption of oxygen and the production of carbon
dioxide has markedly increased.
What conclusions can be drawn from these instances of response to
stimuli, of which any number could still be quoted? They show us,
first of all, that a state or process existing under given conditions, is
altered by the influence of the stimulus. This is a fact, however,
which could be expected from the beginning and is self-evident, for
stimuli are alterations in the vital conditions, and when these are
altered the state of the system or the happenings thereof must also
alter. The question with which we are here more closely concerned,
however, is a somewhat more detailed characterization of the state
or process itself, as well as that of alterations produced by the
influence of the stimulus. The instances of response to stimuli
already cited furnish us with information in both kinds.
In all these examples, the living processes occur with equal
constancy and unaltered rapidity, provided a stimulus is not
operative. Here, however, the gradual alterations, the result of
development, must not be overlooked. An excellent example of this
is seen in the eggs of sea urchin, where the development is readily
perceptible. In all these instances, however, the condition is
immediately changed by the influence of the stimulus. The previous
state of constancy in the vital process is disturbed. The rapidity of its
course is changed, being either increased or decreased, and the
specific vital manifestations concerned are, therefore, augmented or
diminished. We will now study the vital process with the methods of
chemical investigation and consider the problem from the standpoint
of metabolism. It may be noted here, that other methods, such as
the transformation of energy or changes of form of the living
system, would serve equally well as indicators for this purpose. In
every instance there is a uniformity of the processes; the difference,
however, is in the nature of the indicators and the terms used. The
methods and the terms used in chemical investigation and
description reach proportionately much deeper than those employed
when the transformation, energy or the variations of form of the
organisms are studied, and permit of the finest differentiation of the
processes. The atomistic terminology is, for this reason,
preëminently fitted for the description of vital processes. When we
study the vital process metabolically, we can, as shown in the above-
mentioned instance, divide the processes into a metabolism of
stimulation in contradistinction to a metabolism of rest.
The comprehension of the metabolism of rest demands a closer
consideration. On closer observation we must say that this much-
used conception is merely an abstraction nowhere realized in a strict
sense. In truth, there is nowhere in nature a metabolism of rest. No
cell exists which in a mathematical sense remains for even two
successive moments under absolutely the same external conditions.
If we imagine a single living cell of the simplest kind living in a fluid
nutritive medium, and if we suppose its body and surroundings so
magnified that the single molecules and atoms were respectively of
the size of cannon and rifle balls, the boundary between cell and
medium would represent a battlefield, on which a heavy
bombardment is constantly taking place. The rain of shot of food
and oxygen molecules penetrating into the cell from the medium,
would produce an explosion in the existing ammunition depots, now
at one point, now at another, creating great breaches through which
new masses of shot would reach the interior. The fragments of these
exploding molecules would be flung out here and there into the
medium and would stem, now at this, now at that point the
besieging masses of shot. In this wild confusion on the whole
boundary line between cell and medium there can be no question of
rest or even equilibrium at any point. The human mind, superior to
the material world as we may deem it, is yet always dependent upon
the results of experience, and even in its highest flights cannot
become wholly emancipated from the concrete objects. For this
reason it is of great purport to conceive processes whose dimensions
cannot be observed even microscopically, as enlarged and
transformed to that method of expression most familiar to the
human mind, namely, in the field of optical presentation. This
method is of great help in aiding our understanding, and likewise
here, even in the resting state, the cell is constantly exposed to local
effects of stimulation, now at one point, now at the other. The
conception of the metabolism of rest is, therefore, in a strict sense
fiction.
Nevertheless, the conception of the metabolism of rest as an
abstraction can be of value provided always that it is strictly and
definitely limited. It must, for instance, not be applied to short
periods of time. The continued local and temporary responses to
stimulation constitute a mean value which, although composed of
numberless small sub-threshold responses, we can still call a
metabolism of rest. Weak stimuli have, however, as already seen,
the property, provided their influence is constant, of effecting an
adaptation to the stimulus on the part of the living organism, so that
the stimulus becomes a vital condition for this state of the organism.
Hence the continued existence of a vital process resulting from the
constant action of stimulation is made possible. That which we are in
the habit of calling metabolism of rest, would, therefore, be
metabolism of stimulation, but one that is characterized by a
constantly existing metabolic equilibrium.
This “equilibrium of metabolism” distinguishes the metabolism of
rest from that form which is developed in response to temporary
stimulation, in that every temporary stimulation has the effect that it
disturbs the existing metabolic equilibrium for a longer or shorter
time. This disturbance of the equilibrium of metabolism can in
contrast to the metabolism of rest be termed “metabolism of
stimulation.” In this, but only in this sense, can these two
conceptions be placed in opposition and used to characterize the
processes in the living organism. The conception of the metabolism
of stimulation must always stand in relation to that of an equilibrium
of metabolism characterized by a constantly existing metabolism of
rest, just as the conception of stimulus can likewise only be defined
relatively to that of vital conditions.
Nevertheless, the conception of the equilibrium of metabolism
requires a somewhat more accurate definition before we can feel
justified in using this term. Definitions are always trite, nevertheless
they are the basis of all our thinking and a definite understanding is
impossible unless we first clearly fix their contents. The history of
theology and philosophy even to the most recent times furnishes a
long line of instances in which the most eminent minds, for the want
of fixed definitions of the conceptions which they made use of, failed
to find a mutual basis for their ideas. Without a sharp definition
every conception is a mere word, which each individual, according to
his personal experiences and views, endows with a different
meaning. To such conceptions we may apply Mephisto’s ironical
comment to his pupil:
“Mit Worten lässt sich trefflich streiten,
Mit Worten ein System bereiten.”

The natural sciences, if they are to retain their reputation for

exactness and precision, require the strictest and clearest definitions
of all conceptions. If we seek to penetrate more deeply into the
varied happenings in concrete conditions, we must reconcile
ourselves to dry pedantic definitions. In the case of that of the
equilibrium of metabolism indeed we have before us one of the most
important conceptions in physiology.
The justification to speak of an equilibrium of metabolism arises
from investigations of metabolism in mammals. The classical
experiments of the previous century, as is well known, have shown
that in the adult mammal receiving a necessary quantity of
nourishment and in a state of rest, the intake and outgo of the
constituent elements are the same. The carbon, hydrogen, nitrogen,
oxygen, sulphur, phosphorus, etc., taken in during a lengthened
period in the form of food and respired air, appear again in equal
quantity, in other combinations, in the products of excretion of the
organisms. Calorimetric experiments likewise show an equilibrium of
the consumption and elimination of energy. If there thus exists an
equilibrium of metabolism for the whole cell community, it is clear
that the same must also apply to the individual cell, that is, for all
living substance. The quantitative relations of the foodstuffs taken
in, and the excreted metabolic products given off, are, however,
merely a standard of the metabolism. We know that the former are
used to build up new living substance and that the latter represent
the result of disintegration of that previously existing living
substance; for we find, as in the case of the plant, complicated
protein combinations, which are built up from comparatively simple
constituents of the food and are again broken down into
comparatively simple substances. And so the building up and
breaking down processes form the two great processes of
metabolism, which with Hering 41 we can briefly call “assimilation”
and “dissimilation.” In the terms assimilation and dissimilation are
comprised the sum of all processes of construction and
disintegration in the living organism. It is apparent that equilibrium
of metabolism occurs when assimilation and dissimilation are equal.
The formula A : D, that is, the relation of the sum of all assimilation
to the sum of that of all dissimilative processes, is a factor of
fundamental importance in the study of the course of the vital
processes, for upon its value depends individual vital manifestation,
and, in fact, the continuation of life. I have, therefore, designated
the formula A = D “Biotonus.” The equilibrium of metabolism would
then be characterized by the biotonus 42 of a living organism being
equal to one. This would be the metabolism of rest of a system,
whilst its metabolism of stimulation would consist in an alteration of
the biotonus. But is this state of living substance strictly speaking
ever realized?
In considering the nature of the equilibrium of metabolism one
factor has been disregarded which must be taken into account at
every point; this is growth. Growth changes, although varying more
or less, are never absent during the life of the organism. An
equilibrium of metabolism never exists in a strictly mathematical
sense, and here again we are working with a conception which is
faulty, because it is an abstraction, originating from experience with
rather too restricted boundaries. But an error of which one is aware
is not dangerous. In mathematics we also consciously reckon with
errors, without the result being altered. In the before mentioned
cases the equilibrium of metabolism was maintained, because the
investigations involved only a short time in an adult mammal. In the
adult mammal the growth processes occur very slowly, so that
alterations within a relatively short time are not demonstrated.
If it were possible to subject the adult mammal to metabolic or
calorimetric experiments, extending for years, it would be found that
the intake would be qualitatively and quantitatively different at the
end of the investigation and that the same would apply to the outgo.
In the growing egg cell this takes place with much more rapidity. In
the organism which rapidly grows, it can be seen at once that the
quantity of the outgo of the products of disintegration cannot be
equal to that of the intake of foodstuffs. If biotonus were equal to
one, the organism could not grow. Equilibrium of metabolism can
only be understood when we take into consideration a period of time
in which the alterations in growth take place with such imperceptible
slowness that the resultant error is inconsiderably minute. This
period of time is of greatly varying length in different living
organisms and this fact must be taken into account in every living
form. Only with this restriction can we justify the use of the term
“equilibrium of metabolism.” Then, however, its use is of great value.
The metabolism of stimulation is then a disturbance of the
metabolism of rest, that is, a disturbance of the equilibrium of
metabolism through the effect of stimuli.
The question here follows: Is there a constancy of this interruption
of the equilibrium of rest produced by the stimulus which can be
formulated into a general law? To begin with, the number of possible
responses are greater than the variety of forms of living substance,
for every living organism with its specific properties can undergo
alteration in its metabolism in various directions. Thereby results an
infinite number of manifold reactions to stimuli. However, in answer
to the question, in which direction the change in the specific
metabolism of rest in response to a stimulus takes place, we find a
comparatively simple scheme of general reaction. All phenomena can
change in their rapidity as well as in their nature. That is
quantitatively and qualitatively. In this way the specific vital process
of an organism can be altered by the stimulus, on the one hand, in
its rapidity; on the other, in the manner of its action.
The majority of all temporary responses to stimuli consist in
alterations of rapidity of the vital process, and form either a
quickening or retardation of its course. The former is manifested in a
strengthening or an increase, the latter in a decrease or repression
of the specific action of the living organism. The stimuli have the
same effect as in the case of the catalysers in chemical processes.
According to Ostwald’s 43 well-known definition of catalysis a
catalyser is a substance which, without appearing in the final
product of a chemical reaction, alters its rapidity. This group of
reactions can, therefore, be referred to as “catalytic stimulation and
response.” When the response consists in increase, we speak, in a
physiological sense, of an excitation, and when there is decrease in
the vital processes, we speak of a depression.
The conception of excitation and depression are purely empirical.
They are terms for real things, referring, in fact, simply to alterations
in rapidity of life process, which can be as readily observed as the
process itself. I wish to lay particular stress on this fact, for the
reason that Cremer 44 has recently made the extraordinary statement
that I have introduced hypothetical processes into the definition of
the conception of excitation. I have always considered excitation as
merely an increase or change of intensity of the specific actions of a
living system, and as such is an established process without a trace
of the hypothetical element. 45 If, however, the excitation process is
to be regarded as something absolute, as a mysterious state sui
generis, which is entirely independent and totally unlike the
metabolism of rest, then, of course, it would appear utterly
incomprehensible and would be without purpose. As an absolute
process excitation is merely a meaningless word. Excitation and
depression are relative conceptions and can only acquire meaning
when the process which is excitated or depressed is more closely
defined. This is the specific vital process of a given organism, and
the two conceptions only have meaning in relation to it. The
conception of the vital process, however, is one directly gained from
experience. However complex or difficult to analyze the process may
be, it still is as little hypothetical as that of the combustion of carbon
into carbon dioxide, or the revolving of the earth around the sun. It
can be looked upon as something positive and real. Quite another
question is the manner in which we are to consider the mechanism
of the vital process. In analyzing this mechanism we cannot, at least
in the present state of our knowledge, entirely dispense with
hypothesis. But these hypotheses are in no way involved in the
definition of the process of excitation. If we look upon every
excitation or depression produced by a stimulus as an alteration in
rapidity in the specific vital process of a given organism, we are
thereby expressing the same fact which Johannes Müller has termed
“specific energy.” We give, however, the doctrine of specific energy a
more general application in so far as it comprehends not only the
increase but likewise the decrease of activity in response to stimuli.
Johannes Müller’s doctrine of specific energy of the living substance
at all times has been the subject of most animated discussion. When
I refer here to the specific energy of living substance, it is with the
knowledge that Johannes Müller did not use this expression of “living
substance” in this connection. He was already acquainted, however,
as we have seen, with the fact of the existence of the specific
energy of all living structures. For appertaining to the muscle he
says: “This is universal in all organic reaction.” The reason why the
doctrine of sense energy has become of importance in the discussion
of the specific energy of the living substance, is in consequence of
the theoretical interest, resulting from its connection with the nature
of the specific energy of our sense substances. The controversies on
this subject are still far from settled. 46 Indeed, according to the
special philosophical standpoint taken by an observer, the existence
of a specific energy of the senses is acknowledged or disputed. For
any one acquainted with the general physiological reaction to
stimuli, such a discussion is wholly without purport. The sense
substances have as a matter of course in common with all living
substances their specific energy, that is, the influence of stimuli can
produce an increase or decrease of their specific vital processes.
“Specific energy” of “sense substance” in this sense is like that of all
other living substances, a fact. In that the psychical capability of
these sense substances, in which we include not only the peripheral,
but also the central portion, are dependent upon their specific vital
processes, it must be self-evident that the excitation and the
suppression of sense sensation can be brought about by adequate
and inadequate stimuli, no matter what one may think of the
relations between physical and psychical phenomena.
The only debatable question is that concerning the limits of the
validity of the doctrine of the specific energy of living substances.
This question will involve our attention when we have analyzed
somewhat more closely the happenings in the living substance
taking place under the influence of stimuli. We will, therefore, return
later on to a more detailed consideration of the last question.
Nevertheless, we will here refer to a fact which, upon a superficial
observation, seems to restrict the validity of the conception of the
specific energy of living substance.
In contrast to those reactions to stimuli, which consist merely in the
changes of a rapidity of the specific vital process, are another group
of reactions in which the influence of stimuli leads to qualitative
alterations in the specific vital process. In these instances, the
influence of the stimulus directs the metabolism of rest into new
channels, so that chemical processes occur in the cell, which under
ordinary circumstances do not take place. This group of reactions,
which I wish to term “metamorphic stimulation and response,” are
chiefly observed where weak stimuli act continuously upon the living
substance. These are essentially weak chemical stimuli, which last
for a prolonged period or frequently reoccur in the life of the cell
community. Examples of this are found in the continual ingestion of
alcohol and other poisons by the human being, or in the formation
of metabolic products of bacteria, etc. The majority of chronic
diseases belong to this group of reactions; disease being simply
response to stimulation. Disease is life under altered vital conditions
and altered vital conditions are stimuli. This simple and self-evident
fact shows the immense importance which the knowledge of the
general laws of the physiology of stimulation has for pathology. The
pathologist, who does not wish to confine his observations to a
purely superficial symptomatology or a merely histological
morphology, must seek above all to penetrate as deeply as possible
into the nature of the general reactions to stimulation in the living
organism. It is the essential point which meets him everywhere. In
spite of their great interest for pathology, however, it is just these
qualitative alterations of the normal vital process produced by
continuous stimulation which have up to now been least analyzed. In
this field we expect much from pathological investigation which
alone has the immense amount of material at its command. This will
take place only when pathology adds to the almost exclusively
histological direction of investigation, that also of experimental
physiology. It is true that the problems of the qualitative alterations
of a vital process by chronic stimulation are much more complicated
than those of the rapid responses to temporary stimuli, consisting
simply in mere alterations of rapidity of the specific vital process. An
understanding of the nature of the former can only be expected
when a deeper knowledge of the latter is gained, for, as will be seen
presently, there is the closest relation between the two groups.
The reactions to catalytic stimuli of short duration, which produce
merely an alteration of rapidity in the specific phenomena of a living
organism, show on a closer analysis the interesting fact, that it is not
always the entire metabolic processes of the cell which are
perceptibly quickened, but that only certain constituent processes of
the same are affected by the action of excitation. This is the more
noticeable, as, considering the close correlation which all the
individual links of the chain of metabolism bear to each other, it is to
be expected that the alteration in rapidity of one would be followed
at once by a corresponding change in all the others. An example of
the case in question, in which a special constituent process may be
predominately affected, is that of the specific activity of a muscle
which is repeatedly stimulated by nervous impulses. Since the
classical investigation of Fick and Wislicenus 47 on themselves, and of
Voit 48 on the dog, we know that the nitrogen metabolism is
practically unaltered by the functional use of the muscle and there is
a remarkable increase only in the breaking down of the nitrogen-free
groups of the living substance. Sufficient importance has not as yet
been attached to this knowledge. This fact not only has a particular
interest for the much-discussed question of the source of muscle
energy, but also affords a deeper insight into the metabolic activity
of the living substance. It shows us that we must not imagine a
purely linear linking of the individual constituent metabolic
processes, but rather, at least at certain points, a branching
formation, the individual members spreading in various directions.
An alteration in an individual member can occur without an
immediate change in the other branches. This would not be the case
if there were only a linear connection of the constituent processes,
for the breaking of a single member of the chain would be followed
by a change in all the following members.
It shows us, further, that certain branches are more labile than
others. In the case referred to here, the branches of this system,
which bring about the nitrogen metabolism, are relatively firm and
stable, the branches, which are disturbed by the stimulus producing
functional activity of the muscle, are particularly labile. I should like
in passing to call here your attention to the fact that as is well
known, Ehrlich, 49 in another field involving other conditions and
other experiences and considerations, has arrived in analogous
manner at his “side chain theory.” In order to have an expression for
those stimuli which involve rapid alteration of the labile constituent
processes and which are connected with the specific action of the
particular organism, I have called them “functional stimuli,” and
contrasted with them the “cytoplastic stimuli.” In the latter the
alterations produced include all the constituent processes extending
even to the stable processes of nitrogen changes, and sometimes
extend to complete disintegration and rebuilding of living
substance. 50 To the first group belong all adequate stimuli within
certain limits of duration and intensity, and the greater part of
inadequate stimuli of brief duration so long as they do not exceed a
certain intensity. To the latter group belong in general all the
stronger adequate and inadequate stimuli of prolonged duration;
such as extreme temperature, the stronger electric currents,
constant alteration in the supply of food, water, oxygen, the
prolonged or stronger influence of extraneous chemical matter, etc.
Considering the close correlation of the individual part processes it
would appear very strange, however, if a single one of these could
undergo an alteration of its rapidity without the course of the rest of
the processes being in the least influenced. One cannot comprehend
such absolute independence of a process brought about by
functional stimulation from all the other constituent processes,
particularly when this is of prolonged duration and involves to a
considerable extent the alterations in rapidity, for the individual
constituent processes are dependent in a high degree upon the
quantity of the particular chemical substances of which the living
system is composed. The cycle of the individual constituent
processes of this system is determined in the most delicate manner
in its rapidity and extent, by the relative quantities of the individual
substances. Associated with an alteration in the rapidity of an
individual constituent process, there would also be a relative
alteration quantitatively of the substances. And with the increase in
the quantity of the disintegration products, and also the increase of
the substances for their replacement, there would result, during this
time, an alteration in the amount of interaction of the molecules of
the other constituent processes, so that these processes secondarily
suffer an alteration in rapidity which is perceptible after long
continued involvement of the functional part of metabolism.
In fact, in the previously mentioned case of the functional
stimulation of the muscle, the proof has been furnished that a long-
continued increase of the functional metabolism is followed,
although to a less extent, by an increase in the entire cytoplastic
metabolism. Argutinski showed this on himself in 1890 in Pflüger’s
laboratory. He found, namely, that after the exertion of a long walk
in a hilly district, a considerable increase of nitrogen excretion in the
urine took place, which extended over the succeeding two or three
days. This increase of the nitrogen metabolism in its totality is not
nearly as great as that of the breaking down of nitrogen-free
substances, but it is, nevertheless, present and shows us that
functional metabolism cannot experience a lasting excitation without
being followed by secondary results in the entire cytoplastic
metabolism. This fact is even more strikingly illustrated in the
alteration of the entire volume of a living organism as produced by
the lengthened duration of functional stimulation. It has been long
known, that the muscle as the result of frequent functional excitation
by means of adequate nerve impulses, that is, prolonged activity, is
considerably increased in size, whereas in the absence of such it
loses more and more in volume. A hypertrophy of activity, produced
by functional stimuli, and the atrophy of inactivity, the result of the
discontinuance of the functional excitation, is universal and can be
observed in the various tissues of our body. We see it, for example,
in the glands; we see it in the skin and we see it in the elements of
the nervous system. Berger, 51 for instance, established the fact that
the ganglion cells of the optic lobe in the cerebrum of newborn dogs
only reach their full development when functionally excitated by
adequate light stimuli (Figure 9, B), coming from the eye, whereas
they remain in the embryonic state when these light stimuli are
eliminated. (Figure 9, A.) The cytoplastic increase of volume of the
neurons under the influence of functional stimuli is a fact of
fundamental importance for the entire happenings of the nervous
system and forms the physiological basis for reinforcement of
reflexes, which, in its turn, is essential for all acts of memory and
intelligence. For the increase in volume of the ganglion cell body is,
when functionally activated, accompanied at the same time by an
increase of specific capabilities and the intensity of discharge. Its
excitation impulses can, therefore, be conducted through a greater
number of neurons, with which it is connected, than would be the
case if development of the volume of the ganglion cell increased to a
less extent.
 Fig. 9.
A—Undeveloped ganglia cells in the optic lobe of
a dog, the eyes of which have been sewn up
immediately after birth. B—Fully developed
ganglia cells in the same region of a normal dog
of the same age. (After Berger.)

The increase in volume under the influence of stimuli further shows

the relation between the group of those solely catalytic effects of
stimulation consisting in mere alterations of rapidity of the specific
vital process, and that of the metamorphotic effects of stimulation,
which manifest themselves in qualitative alterations of the vital
process. Simple observation shows us that a qualitative change of
individual constituent processes must necessarily result from the
increase of volume of a cell, and that considering the close
correlation of all the individual processes a profound alteration of the
entire metabolism must be produced. I have already at another
place 52, 53 treated these conditions more in detail and will, therefore,
only briefly refer to them here. If we study the growth of a ball-
shaped cell, we find that the surface then increases as a square, and
the volume as the cube. It therefore follows that, by progressive
volume increase, the conditions for the interchange of substance
with the surrounding medium must become more and more
unfavorable for those cell portions situated in the interior, whereas
those at the exterior are at much greater advantage. This must lead
to a constantly increasing difference of the rapidity of the metabolic
processes between the peripheral and central portions. Accordingly,
the intricate interworkings of the individual constituent processes,
the rapidity of action of all which is intimately connected, are,
therefore, followed by corresponding alterations in the entire
metabolism. Sooner or later a stage is reached in which the
individual constituent processes become so limited that certain
metabolic products, which previously were broken down as soon as
formed, can be no longer eliminated and remain in the cell acting as
foreign bodies. In this way the relative quantity of the individual cell
substances become more and more altered, and as the course of
chemical processes occurs in accordance with the law of mass
action, the whole metabolism is directed into another channel, so
that finally new constituent processes take place, which were
formerly not possible. These in their turn produce deep-seated
alterations of the relations of the cell to its surrounding medium, etc.
Hence this mere increase of volume of the cell in growth forms the
source of an infinite mass of alterations in the activities of cell
metabolism, which we briefly term its “development,” and which by
constant progression, leads either to a process of cell division, and
with this to a correction of existing disorder, or finally to irreparable
disturbances ending in death. In this way an inseparable relation
exists between increase of volume and the development of living
substance. We have seen, however, that the catalytic reactions of
stimulation, which at first only produce an alteration of rapidity of
the individual constituent processes, if of prolonged duration or of
frequent recurrence, secondarily effect a change of volume of the
entire living organism. One can, therefore, hardly reject the
conclusion that seeing the close interworkings of the individual part
process of metabolism, every change of rapidity of a single member,
if of prolonged duration or of frequent occurrence, must finally lead
to qualitative alterations of the entire metabolism. In consequence
there results an important dependence between catalytic stimulation
and metamorphic reaction. Indeed, it is not unlikely that the
metamorphic reactions, which are especially seen in the continued
effect of weak stimuli, result from alterations of rapidity, which the
individual members of the vital processes have primarily undergone
from this influence.
It is perhaps expedient to cite a concrete instance in illustration. A
simple example is furnished by asphyxiation. If oxygen is withdrawn
from any living organism, the result is a depression of its oxydation
processes. Here there is primarily only a change in rapidity,
especially a retardation of oxydation processes. The metabolism,
however, proceeds, the disintegration of living substance continues,
although at a slower rate, but produces an accumulation of other
products. Whereas formerly during the existence of a sufficient
supply of oxygen an oxydative disintegration of nitrogen-free groups
into carbon dioxide and water took place, both of which could easily
be eliminated from the cell, the anaërobic disintegration furnishes
only complex products, having a higher carbon content, such as
lactic acid, fatty acids, aceton, etc. These, being more difficult to
excrete from the cell, accumulate. These asphyxiation products have
in their turn a depressing effect and so on. In this way the whole
metabolism is forced into a wrong course. The accumulation of fat in
those tissue-cells with an insufficient blood supply, as we have seen
in the case of the fat metamorphosis, is doubtless brought about in
the same manner by relative oxygen insufficiency. The fatty acids
accumulate as products of an incomplete combustion and combine
with glycerine to form neutral fats. In like manner it may be that the
accumulation of amyloid substance in amyloid metamorphosis, of
lime salts in arteriosclerosis, etc., is produced by a primary
depression of the individual constituent processes of the particular
cells.
The relation here described, of the catalytic stimuli to the production
of the metamorphic processes, leads us to the distinctions between
primary and secondary effects of stimulation. Should the general fact
be established, which has up to now only been pointed out in
individual cases, that all the metamorphic processes are merely
secondary results of primary alterations in rapidity of individual
metabolic constituent processes, then the primary reactions of every
stimulus would consist purely in the excitation or depression of the
directly concerned constituent. Whether or not, as may be assumed,
this primary effect of stimulation applies to all stimuli, is a question
which only the future can answer.
The metamorphic processes are not, however, the only secondary
effects of stimulation. The influence of long-continued excitation of
the functional constituent processes upon the entire cytoplastic
metabolism can be looked upon as a secondary response. Therefore,
they may be considered as a secondary effect of stimulation which,
in contrast to this primary excitation, may be called the secondary
excitation.
Further: While the secondary excitation and metamorphic processes
are generally produced by the continued existing effects of weak
stimulation, we also observe as the result of a stimulus of short
duration or frequently repeated at brief intervals, but otherwise not
exceeding the physiological limits of intensity, a secondary effect,
which plays a very important part in the activity of the organism. I
refer to fatigue. Here a secondary depression is developed in
connection with the primary excitation, for fatigue of a living
organism must be characterized as a depression of activity. This case
shows that we have to distinguish between a primary depression, as
for example, produced by temperature reduction, withdrawal of
food, deficiency of oxygen, etc., which occurs as a direct effect of
stimulation, and secondary depression, which as in fatigue is an
indirect result of primary excitation.
After the cessation of a briefly catalytic stimulus, not exceeding the
physiological limit of intensity, another secondary result is observed,
which is of the greatest importance for the continued existence of
the living substance. The catalytic stimulus brings about a
disturbance of the equilibrium of metabolism, which after cessation
of the stimulus is reestablished by the living substance. In other
words: recovery takes place. This fundamental principle has been
known for a long time as the result of observation. If a skeletal
muscle of our body has been activated for a prolonged period by
nerve impulses, until it has become completely fatigued and
incapable of work, a recovery takes place on the cessation of these
impulses and the muscle is again capable of action. Likewise, as the
result of strong mental activity during the day, we are mentally
fatigued in the evening; recovery, however, occurs during the night,
which results from the removal of the source of activity. The next
morning finds us refreshed. This restitution occurs in every cell, and
the return of its former capability of action, which had disappeared
under the influence of stimulation, shows that compensation has
taken place of the metabolism of rest, disturbed by the effects of the
stimulus. Hering 54 has aptly termed this restitution as “the internal
self-regulation of metabolism.” All recovery after disease is based on
this self-regulation. The physician simply provides, by means of
therapy, for the possibility of its taking place. Healing itself is
brought about by the organism. “Natura sanat, medicus curat.”
Finally, a third kind of secondary effect of stimulation claims our
interest. This is the secondary extension of the result of stimulation
from the part of a living organism directly and primarily affected by
the stimulus, to the surrounding structures. All living substance has
the capability of conducting an excitation, which is produced locally
through a catalytic stimulus, to a neighboring part, not directly
affected by the stimulus. It finds its highest development in the
nerve, but in no living structure is it completely absent. This
capability has been frequently termed “conductivity of stimulation.”
It is more precise, however, to speak of conductivity of excitation,
for it is not the primary influencing external stimulus which is
conducted in the living substance, but the excitation which it has
produced. I have intentionally considered only the excitating effects
of stimulation, and not those of the depressing reactions, as only
excitations, not depressions, are conducted by the living substance.
These questions, however, demand a closer analysis. Here we were
concerned only with a survey of the general effects of stimulation. If
I, therefore, once more summarize the results which have been
gained, this is most clearly demonstrated by the following scheme:
Primary Effects of Stimulation
Excitation Depression
Functional Cytoplastic Functional

Secondary Effects of Stimulation

Secondary excitation Secondary depression
Conduction of excitation, Metamorphic processes, Self-regulation of metabolism

This, however, is simply a scheme, like all other schemes, having for
its purpose a superficial survey of the subject.
It brings to some extent order into the overwhelming mass of
manifold effects of stimulation but tells us nothing of the mechanism
and genesis. Our further task must, therefore, be a more thorough
analysis of this field.
 CHAPTER V
THE ANALYSIS OF THE PROCESS OF EXCITATION
Contents: Indicators for the investigation of the process of
excitation. Latent period. The question of the
existence of assimilatory excitations. Dissimilatory
excitations. Excitations of the partial components of
functional metabolism. Production of energy in the
chemical splitting up processes. Oxydative and
anoxydative disintegration. Theory of oxydative
disintegration. Dependence of irritability on oxygen.
Experiments on unicellular organisms, nerve centers
and nerve fibers. Restitution after disintegration by
metabolic self-regulation. Organic reserve supplies of
the cell. The question of a reserve supply of oxygen
of the cell. Metabolic self-regulation as a form of the
law of mass effect, and metabolic equilibrium as a
condition of chemical equilibrium. Functional
hypertrophy.
If it is true that all primary effects of stimulation consist either in an
excitation or depression of the metabolism, and that all other effects
of stimulation secondarily follow this primary alteration of the
metabolism of rest, then every thorough analysis of the mechanics
of reaction must have its beginning in the investigation of these
primary processes. I desire to adopt this method here and will
analyze somewhat further the primary process of excitation and its
immediate and remote sequences. This will be followed later by the
analysis of the process of primary depression and its results.
The investigation of the more obscure processes in the living
substance places us in a difficult position, for their details cannot be
observed by the unaided senses. That which we can perceive is
merely the grosser vital action, consisting of a complex combination
of the individual processes, the total result of a multitude of different
components. For this reason the conception of excitation can only be
established by observations based upon the combined vital actions,
which are produced by the effect of stimulation upon the complex
system. In the beginning, the process of excitation was studied
exclusively on the muscle and nervous system. A physical factor
served as indicator, such as muscle contraction or production of
electricity. These showed, besides the direct and primary effect of
stimulation, the secondary process of conductivity. Even graphic
registration is merely an expression of the phenomena composed of
a great mass of individual elements. The visible course of the
phenomena, as shown, for instance, by the latent period by the
ascent and descent of the curve of contraction, represents as it were
a reflected picture of the actual excitation processes similar to an
object seen in a distorting mirror; the first and the last parts of the
process are not even perceptible. Later, when organ physiology was
extended into a cell physiology the processes of excitation were
studied in numerous simple organisms, such as the plant cell, the
rhizopoda, the infusoria, etc. Later, in this way, by the use of
comparative methods many essential facts were discovered.
However, even the single cell, in spite of its minuteness, is,
compared with the size of a molecule, a gigantic system, and it
would be a grave error if we should consider this system even in its
simplest aspect as homogeneous. In order, therefore, to analyze the
vital activities in the cell, cell physiology must endeavor to penetrate
into molecular conditions. For this purpose the indicators employed
must be essentially of a chemical nature, capable of magnifying the
processes of molecular dimension to such a degree that we are
enabled to base conclusions upon these not otherwise directly
perceptible phenomena. To obtain a sufficient magnification we must
necessarily place somewhat larger quantities of living substance
under observation and apply a stimulus of such frequency or length
of duration that the chemical alterations as a result of excitation are
so increased as to be plainly perceptible with the aid of our chemical
indicators. Unfortunately, we do not possess specific chemical
indicators for every individual molecular constituent process of the
cell and so cannot dispose with the help of indicators of the
combined happenings in a greater quantity of living substance. It
remains for us to obtain data concerning the cycle of excitation
processes in the living substances by the aid of the combined
employment of the most varied kinds of physical as well as chemical
indicators. If we use the most varied types of living substance of
widely differing properties, showing us the greatest variety of vital
manifestations, we may hope by the use of comparative
physiological methods, even though with difficulty, to separate more
and more the essential details of the general processes of excitation.
At present we are still at the very beginning of this task and vast
fields of unexplored regions are yet before us. But it is the unknown
which has a particular fascination, especially if we succeed from time
to time in making new advances.
If we suppose a living system in a state of metabolism of rest
influenced by an instantaneously excitating stimulus, the entire
course of excitation extends from the first alteration produced by the
stimulation until the complete restitution of the metabolic
equilibrium, and we will, therefore, differentiate individually the
successive stages of this whole process.
The very beginning of the chain of alterations produced by the
excitating stimulus cannot be studied by any indicator. The changes
must first reach a certain dimension by conduction from the point of
stimulation before they influence even the most delicate indicators.
The application of the stimulus is, therefore, followed at first by a
measurable “latent period,” in which the living substance remains
apparently at rest. This latent period has been particularly studied in
muscle. After its discovery by Helmholtz 55 it was made the object of
innumerable investigations and met with an interest which can only
be explained by the exactness of the methods employed. Among
others Tigerstedt 56 has made the most thorough study of the
influence of various factors on the duration of the latent period.
These experiments have established the fact that the duration of the
latent period varies according to the intensity of the stimulus,
temperature, loading or fatigue. This is apparent when it is
understood that the amount of the alterations produced by the
stimulus must ascend from the value zero to a certain height before
the changes are perceptible, and that under various conditions this
amount is, on the one hand, attained in different lengths of time
and, on the other, must reach a varying amount before it is
perceptible by means of the indicator.
The facts concerning the whole latent period and its dependence on
various factors would be incomprehensible if it were assumed that
no alterations whatever take place during the latent period although
the stimulus is already operative. In reality, the alterations following
a stimulus occur with imperceptible rapidity in the form of a
molecular interchange, and the latent period is simply an expression
of the fact that the primary alterations, being limited in nature, are
not registered by our indicators.
The question first arises, In what do these first imperceptible
alterations consist? Nernst 57 has evolved the theory for electric
stimulus, that the primary effect produced by the electric current is
an alteration in the ion concentration on the surface of the living
substance. In fact, we know that the surfaces of all protoplasm
possess the property of semi-permeable membranes and that
changes in the concentration of ions invariably occur when an
electric current flows through two electrolytes separated by a semi-
permeable membrane, in which the anions and cations have a
different rapidity of movement. It is apparent, therefore, that such
an alteration in the ion concentration must be followed by further
chemical processes in the living substance. According to the theory
of Nernst the first impetus for all further alterations, which the
electrical stimulus brings about in the metabolism of rest, is the
alteration in the concentration of the ions on both sides of the semi-
permeable membrane, which represents the surface of the
protoplasm. In view of the present findings of physical chemistry,
objections can hardly be made to this theory of Nernst’s. It is a
question, however, in how far this theory, especially established for
the electric stimuli, can be applied to other forms of stimuli and their
action. It cannot be denied that the degree of dissociation of an
electrolyte can be altered by very different factors, such as heat,
light, chemical processes, etc., and in that the surfaces of the
protoplasm, acting as semi-permeable membranes, bring about a
selective action on the passage of the ions, there arises the
opportunity for the development of difference of electrical potential
on both sides, and for further chemical alterations in the protoplasm.
These observations, however, require further experimental
investigations in many fields, before we are justified in extending the
Nernst theory of the manner of action of the electric stimuli to a
general explanation of the primary alterations produced by all stimuli
in the living substance. For the present we must confine our
observations to those alterations which are known to be responses
to an excitating stimulus; these are the chemical alterations in the
metabolism of rest in the living substance.
If it is asked, which members of the entire metabolic chain are
increased primarily by the stimulating excitation of a vital system, we
should not be able to answer this question generally for all living
systems. To begin with, it appears highly probable that the various
forms of vital substances in this respect act quite differently. It is to
be regretted that, up to the present, this question has not been
treated from a comparative standpoint. This inquiry should be