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Essential System
Administration
 THIRD EDITION

Essential System
Administration

Æleen Frisch

Beijing • Cambridge • Farnham • Köln • Paris • Sebastopol • Taipei • Tokyo

Essential System Administration, Third Edition
by Æleen Frisch

Copyright © 2002, 1995, 1991 O’Reilly Media, Inc. All rights reserved.
Printed in the United States of America.

Published by O’Reilly Media, Inc., 1005 Gravenstein Highway North, Sebastopol, CA 95472.

O’Reilly Media, Inc. books may be purchased for educational, business, or sales promotional use.
Online editions are also available for most titles (safari.oreilly.com). For more information contact
our corporate/institutional sales department: (800) 998-9938 or corporate@oreilly.com.

Editor: Michael Loukides

Production Editor: Leanne Clarke Soylemez
Cover Designer: Edie Freedman
Interior Designer: David Futato

Printing History:
August 2002: Third Edition.
September 1995: Second Edition.
October 1991: First Edition.

Nutshell Handbook, the Nutshell Handbook logo, and the O’Reilly logo are registered
trademarks of O’Reilly Media, Inc. Essential System Administration, Third Edition, the image of an
armadillo, and related trade dress are trademarks of O’Reilly Media, Inc. Many of the designations
used by manufacturers and sellers to distinguish their products are claimed as trademarks. Where
those designations appear in this book, and O’Reilly Media, Inc. was aware of a trademark claim,
the designations have been printed in caps or initial caps.

While every precaution has been taken in the preparation of this book, the publisher and author
assume no responsibility for errors or omissions, or for damages resulting from the use of the
information contained herein.

Library of Congress Cataloging-in-Publication Data

Frisch, AEleen
Essential System Administration/by AEleen Frisch.--3rd ed.
p. cm.
Includes index.
ISBN 0-596-00343-9
ISBN13 978-0-596-00343-2
1. UNIX (Computer file) 2. Operating systems (Computers) I. Title.
QA76.76.063 F75 2002
005.4'32--dc21 2002023321

[M] [05/07]
 For Frank Willison

“Part of the problem is passive-aggressive

behavior, my pet peeve and bête noire, and I don’t
like it either. Everyone should get off their high
horse, particularly if that horse is my bête noire.
We all have pressures on us, and nobody’s
pressure is more important than anyone else’s.”
***
“Thanks also for not lending others your O’Reilly
books. Let others buy them. Buyers respect their
books. You seem to recognize that ‘lend’ and ‘lose’
are synonyms where books are concerned. If I
had been prudent like you, I would still
have Volume 3 (Cats–Dorc) of the
Encyclopedia Britannica.”
 Table of Contents

Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi

1. Introduction to System Administration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Thinking About System Administration 3
Becoming Superuser 6
Communicating with Users 12
About Menus and GUIs 14
Where Does the Time Go? 31

2. The Unix Way . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

Files 33
Processes 53
Devices 61

3. Essential Administrative Tools and Techniques . . . . . . . . . . . . . . . . . . . . . . . . 74

Getting the Most from Common Commands 74
Essential Administrative Techniques 90

4. Startup and Shutdown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127

About the Unix Boot Process 127
Initialization Files and Boot Scripts 151
Shutting Down a Unix System 169
Troubleshooting: Handling Crashes and Boot Failures 173

5. TCP/IP Networking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180

Understanding TCP/IP Networking 180
Adding a New Network Host 202
Network Testing and Troubleshooting 219

vii
 6. Managing Users and Groups . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
Unix Users and Groups 222
Managing User Accounts 237
Administrative Tools for Managing User Accounts 256
Administering User Passwords 277
User Authentication with PAM 302
LDAP: Using a Directory Service
for User Authentication 313

7. Security . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 330
Prelude: What’s Wrong with This Picture? 331
Thinking About Security 332
User Authentication Revisited 339
Protecting Files and the Filesystem 348
Role-Based Access Control 366
Network Security 373
Hardening Unix Systems 387
Detecting Problems 391

8. Managing Network Services . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 414

Managing DNS Servers 414
Routing Daemons 452
Configuring a DHCP Server 457
Time Synchronization with NTP 469
Managing Network Daemons under AIX 475
Monitoring the Network 475

9. Electronic Mail . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 521

About Electronic Mail 521
Configuring User Mail Programs 532
Configuring Access Agents 537
Configuring the Transport Agent 542
Retrieving Mail Messages 596
Mail Filtering with procmail 599
A Few Final Tools 614

10. Filesystems and Disks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 616

Filesystem Types 617
Managing Filesystems 621

viii | Table of Contents

 From Disks to Filesystems 634
Sharing Filesystems 694

11. Backup and Restore . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707

Planning for Disasters and Everyday Needs 707
Backup Media 717
Backing Up Files and Filesystems 726
Restoring Files from Backups 736
Making Table of Contents Files 742
Network Backup Systems 744
Backing Up and Restoring
the System Filesystems 759

12. Serial Lines and Devices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 766

About Serial Lines 766
Specifying Terminal Characteristics 769
Adding a New Serial Device 776
Troubleshooting Terminal Problems 794
Controlling Access to Serial Lines 796
HP-UX and Tru64 Terminal Line Attributes 797
The HylaFAX Fax Service 799
USB Devices 807

13. Printers and the Spooling Subsystem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 814

The BSD Spooling Facility 818
System V Printing 829
The AIX Spooling Facility 848
Troubleshooting Printers 858
Sharing Printers with Windows Systems 860
LPRng 864
CUPS 874
Font Management Under X 878

14. Automating Administrative Tasks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 885

Creating Effective Shell Scripts 886
Perl: An Alternate Administrative Language 899
Expect: Automating Interactive Programs 911
When Only C Will Do 919
Automating Complex Configuration Tasks with Cfengine 921

Table of Contents | ix
 Stem: Simplified Creation of Client-Server Applications 932
Adding Local man Pages 942

15. Managing System Resources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 945

Thinking About System Performance 945
Monitoring and Controlling Processes 951
Managing CPU Resources 963
Managing Memory 978
Disk I/O Performance Issues 1001
Monitoring and Managing Disk Space Usage 1007
Network Performance 1017

16. Configuring and Building Kernels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1024

FreeBSD and Tru64 1026
HP-UX 1031
Linux 1033
Solaris 1046
AIX System Parameters 1047

17. Accounting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1049

Standard Accounting Files 1051
BSD-Style Accounting: FreeBSD, Linux, and AIX 1052
System V–Style Accounting: AIX, HP-UX, and Solaris 1058
Printing Accounting 1066

Afterword: The Profession of System Administration . . . . . . . . . . . . . . . . . . . . . . . 1069

SAGE: The System Administrators Guild 1069
Administrative Virtues 1070

Appendix: Administrative Shell Programming . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1073

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1097

x | Table of Contents
 Preface

This book is an agglomeration of lean-tos and annexes

and there is no knowing how big the next addition will
be, or where it will be put. At any point, I can call the
book finished or unfinished.
—Alexander Solzhenitsyn

A poem is never finished, only abandoned.

—Paul Valery

This book covers the fundamental and essential tasks of Unix system administra-
tion. Although it includes information designed for people new to system administra-
tion, its contents extend well beyond the basics. The primary goal of this book is to
make system administration on Unix systems straightforward; it does so by provid-
ing you with exactly the information you need. As I see it, this means finding a mid-
dle ground between a general overview that is too simple to be of much use to
anyone but a complete novice, and a slog through all the obscurities and eccentrici-
ties that only a fanatic could love (some books actually suffer from both these condi-
tions at the same time). In other words, I won’t leave you hanging when the first
complication arrives, and I also won’t make you wade through a lot of extraneous
information to find what actually matters.
This book approaches system administration from a task-oriented perspective, so it
is organized around various facets of the system administrator’s job, rather than
around the features of the Unix operating system, or the workings of the hardware
subsystems in a typical system, or some designated group of administrative com-
mands. These are the raw materials and tools of system administration, but an effec-
tive administrator has to know when and how to apply and deploy them. You need
to have the ability, for example, to move from a user’s complaint (“This job only
needs 10 minutes of CPU time, but it takes it three hours to get it!”) through a diag-
nosis of the problem (“The system is thrashing because there isn’t enough swap
space”), to the particular command that will solve it (swap or swapon). Accordingly,
this book covers all facets of Unix system administration: the general concepts,

xi

This is the Title of the Book, eMatter Edition

Copyright © 2007 O’Reilly & Associates, Inc. All rights reserved.
underlying structure, and guiding assumptions that define the Unix environment, as
well as the commands, procedures, strategies, and policies essential to success as a
system administrator. It will talk about all the usual administrative tools that Unix
provides and also how to use them more smartly and efficiently.
Naturally, some of this information will constitute advice about system administra-
tion; I won’t be shy about letting you know what my opinion is. But I’m actually
much more interested in giving you the information you need to make informed
decisions for your own situation than in providing a single, univocal view of the
“right way” to administer a Unix system. It’s more important that you know what
the issues are concerning, say, system backups, than that you adopt anyone’s spe-
cific philosophy or scheme. When you are familiar with the problem and the poten-
tial approaches to it, you’ll be in a position to decide for yourself what’s right for
your system.
Although this book will be useful to anyone who takes care of a Unix system, I have
also included some material designed especially for system administration profes-
sionals. Another way that this book covers essential system administration is that it
tries to convey the essence of what system administration is, as well as a way of
approaching it when it is your job or a significant part thereof. This encompasses
intangibles such as system administration as a profession, professionalism (not the
same thing), human and humane factors inherent in system administration, and its
relationship to the world at large. When such issues are directly relevant to the pri-
mary, technical content of the book, I mention them. In addition, I’ve included other
information of this sort in special sidebars (the first one comes later in this Preface).
They are designed to be informative and thought-provoking and are, on occasion,
deliberately provocative.

The Unix Universe

More and more, people find themselves taking care of multiple computers, often
from more than one manufacturer; it’s quite rare to find a system administrator who
is responsible for only one system (unless he has other, unrelated duties as well).
While Unix is widely lauded in marketing brochures as the “standard” operating sys-
tem “from microcomputers to supercomputers”—and I must confess to having writ-
ten a few of those brochures myself—this is not at all the same as there being a
“standard” Unix.At this point, Unix is hopelessly plural, and nowhere is this plural-
ity more evident than in system administration. Before going on to discuss how this
book addresses that fact, let’s take a brief look at how things got to be the way they
are now.
Figure P-1 attempts to capture the main flow of Unix development. It illustrates a sim-
plified Unix genealogy, with an emphasis on influences and family relationships
(albeit Faulknerian ones) rather than on strict chronology and historical accuracy. It

xii | Preface

This is the Title of the Book, eMatter Edition

Copyright © 2007 O’Reilly & Associates, Inc. All rights reserved.
traces the major lines of descent from an arbitrary point in time: Unix Version 6 in
1975 (note that the dates in the diagram refer to the earliest manifestation of each
version). Over time, two distinct flavors (strains) of Unix emerged from its beginnings
at AT&T Bell Laboratories—which I’ll refer to as System V and BSD—but there was
also considerable cross-influence between them (in fact, a more detailed diagram
would indicate this even more clearly).

AT&T Bell Labs

- direct descent (c.1969-1970)
- strong influence Version 6
(1975)

BSD Version 7
(1977) (1979)

XENIX
(1979 onward)
System III
(1982)

4.2 BSD System V.2

(1984) (1984)

4.3 BSD System V.3

(1985) (1986)

4.4 BSD OSF/1 System V.4

(1993) (c.1992) (1988)

Figure P-1. Unix genealogy (simplified)

For a Unix family tree at the other extreme of detail, see http://perso.
wanadoo.fr/levenez/unix/. Also, the opening chapters of Life with UNIX,
by Don Libes and Sandy Ressler (PTR Prentice Hall), give a very enter-
taining overview of the history of Unix. For a more detailed written his-
tory, see A Quarter Century of UNIX by Peter Salus (Addison-Wesley).

Preface | xiii

This is the Title of the Book, eMatter Edition

Copyright © 2007 O’Reilly & Associates, Inc. All rights reserved.
Exploring the Variety of Random
Documents with Different Content
in the general sense, may be present in the metabolism of some
animals. Certain lower plants, the zoospores of algæ, exhibit
movements identical in character with those of lower animals. At the
base of both kingdoms are organisms, the Peridinians, for instance,
which have much of the structure of the animal (though cellulose is
present in their skeleton), which possess motile organs, but which
also possess a photo-synthetic apparatus, and exhibit the typical
plant mode of nutrition. Further, there are symbiotic partnerships,
that is, associations of plant and animal in one “individual” form (as,
for instance, among the lower worms, Echinoderms, polyzoa,
molluscs, and other groups of animals). In these cases green algal
cells, capable of forming starch from carbon dioxide and water under
the influence of light, become intercalated among the tissues of the
animal. We find, also, that with regard to some fundamental
characters, plant and animal display close similarities: the structure
of the cell, for example, and the highly special mode of conjugation
of the germ-nuclei in sexual reproduction. We must regard all the
distinctive characters of the plant as represented in the animal and
vice versa. Why they have become specialised in different directions
is a question that we discuss later.
The organism, then, in so far as we regard it as a physico-chemical
mechanism, as the theatre of energetic happenings, exhibits the
following general characters:—
(1) It slowly accumulates available energy in the form of
chemical compounds of high potential, work being done upon
it.
(2) It liberates this energy in relatively rapid, controlled,
“explosive reactions,” transforming into movements carried
out by a sensori-motor system of parts, work being done by
it.
 (3) In all these transformations the amount of energy which
is dissipated is relatively small, and tends to vanish.
From the point of view, then, of energetic processes these are the
characters of life, using the term in the general sense indicated
above. 16
Is there an absolute distinction between the organic mechanism and
the inorganic one? Let us note, for the first time, that the actual
physico-chemical transformations themselves, which we study in
inorganic matter, are identical with those which we study in the
organism. Molecules of carbon dioxide, water, nitrate, sodium
chloride, potassium chloride, phosphate, and so on, are just the
same in inert matter as in the organism. Chemical transformations,
such as the hydrolysis of starch, the inversion of cane sugar, or the
splitting of a neutral fat, are certainly just the same processes,
whether we carry them out in the glass vessels of the laboratory, or
observe them to proceed in the living tissues of the animal body.
The same molecular rearrangements, and the same transfers of
energy, occur in both series of events. This, however, is not the
material of a distinction: what we have to find is, whether the
direction of a group of physico-chemical reactions is the same in the
organism and in a series of inorganic processes.
Let us return to the Carnot cycle. This is a series of operations which
occur in an imaginary mechanism in such a manner that the whole
series can be easily reversed. Heat is supplied to the imaginary
engine, which then performs work and yields up its heat to a
refrigerator. Work is then performed on the engine, which thereupon
takes heat from the refrigerator and returns it to the source. The
work done by the engine in the direct cycle is equal to the work
done on it in the indirect cycle. The heat taken from the source and
given to the refrigerator in the direct cycle is equal to the heat taken
from the refrigerator and given to the source in the indirect cycle.
But it is a purely imaginary mechanism, and all experience shows
not only that it has not been realised in practice, but that it cannot
so be realised. If it could be realised, we should show that the
second law of thermo-dynamics is not physically true.
Do the energy processes of life realise such a perfectly reversible
cycle of operations? In order to answer this question we must
consider the fate of the energy which is absorbed in the plant
metabolic cycle, and that which is given out in the animal one. Does
all the energy of solar radiation which is absorbed by the plant pass
into the form of the potential chemical energy of the carbohydrates
and other substances manufactured? Does any of the energy of the
animal which results from the metabolism of its body pass into the
unavailable form—that is, into a form in which it cannot be utilised
by other organisms? That is to say, is energy dissipated by the
organism?
Undoubtedly it is to some extent, but to a far less extent than in the
inorganic train of processes. Some of the energy of solar radiation
absorbed by the plant must become transformed, by the friction of
whatever movements occur, into low-temperature heat, and some
quantity of heat, however small, is generated by the metabolism of
the plant. Again, some of the heat of the warm-blooded animal must
be radiated into space, or conducted away from its body; and this
energy becomes dissipated—let us assume, at least, that it is so
dissipated in the physical sense. Probably also some quantity of heat
is generated by the metabolism of the cold-blooded animal, though
this must be a very small proportion of the total energy transformed.
We see, then, that the distinction is one of degree, though the
difference between inorganic and organic energetic processes is very
great in this respect; so great that we must regard it as constituting
a fundamental difference, and as indicative of the limitation of the
second law when extended to the functioning of the organism.
But we have also to consider the effect of the work done by the
organism. We consider the nature and meaning of the evolutionary
process in a later chapter, but in the meantime we may state this
thesis: that the process of evolution leads up to man and his activity.
It leads, if we regard the process as a directed one; but even if we
regard it as a fortuitous process we still find that man, far more than
any other organism, is the result of it. All the facts of biology and
history show that man dominates the organic world, plant or animal;
that the whole trend of his activity is to eliminate whatever
organisms are inimical, and to foster those that are useful. Already,
during the brief period of his rational activity, the wolf has
disappeared from civilised lands while the dog has been produced.
Species after species of hostile or harmful organisms have been, or
are being, destroyed or changed, while numerous other species have
been preserved and altered for his benefit. In the future we see an
organic world subservient to him either entirely or to an enormous
extent.
So also in the inorganic world. Rivers which formerly rushed down
through rapids, dissipating their energy of movement in waste
irrecoverable heat, now pour through turbines and water wheels,
generating electricity and accumulating available energy. Winds
which “naturally” dissipated their mechanical energy in waste heat
now propel ships and windmills. Tides, with their incredibly great
mechanical energy, now simply warm up the crust of the earth by an
infinitesimal fraction of a degree daily, and produce heat which at
once radiates into space. Who doubts that by and by this energy too
will become accumulated for human use? Multitudes of chemical
reactions were potential, so to speak, in the molecules of petroleum,
while the energy which might have produced them ran to waste. But
under human activity this energy became directed and made to
produce chemical reactions formerly existing only in their possibility,
and all the substances of modern organic chemistry came into
existence.
The energy, then, of human activity has been directed towards
averting or retarding the progress towards dissipation, or
irrecoverable waste, of cosmic energy—that of the sun’s radiation,
and of the motions of earth and moon. Human activity has
accumulated available energy. The difference of water-level between
Niagara and the rapids below represents available mechanical
energy. A few years ago an enormous quantity of this energy
became irredeemably lost in waste heat every twenty-four hours:
now it remains available for work; and this quantity of work retained
is enormously greater than is the human energy which was
expended on erecting the water-power installation there.
The processes studied by physics and chemistry are therefore
irreversible ones. We can conceive a perfectly reversible process, as
in the Carnot heat-engine, but this is a purely intellectual conception,
formed as the limit to a series of operations which approximate
closer and closer to an ideal reversibility. It is a conception that has
no physical reality—a guide to reasoning only. On the other hand we
see that all naturally occurring physical processes are irreversible
and in their sum tend to complete degradation of energy.
Mechanistic biology isolates physico-chemical processes in the
functioning of the organism, and sees that they conform to the law
of dissipation, as well as to that of the conservation of energy.
Yet the organism as a whole, that is, life as a whole, on the earth,
does not conform to the law of dissipation. That which is true of the
isolated processes into which physiology decomposes life is not true
of life. In all inorganic happenings energy becomes unavailable for
the performance of work. Solar radiation falling on sea and land
fritters itself away in waste irrecoverable heat, but falling on the
green plant accumulates in the form of available chemical energy.
The total result of life on the earth in the past has been the
accumulation of enormous stores of energy in the shape of coal and
other substances. By its agency degradation has been retarded.
Whenever, says Bergson, energy descends the incline indicated by
Carnot’s law, and where a cause of inverse direction can retard the
descent, there we have life.
 CHAPTER III
THE ACTIVITIES OF THE ORGANISM
The rather lengthy discussion of the last chapter was necessary in
order to show just how far the principles of energetics established by
the physicists applied to the organism. We have seen that the first
law of thermodynamics does so apply with all its exclusiveness. The
more carefully a physiological experiment is made; the more closely
do its results correspond with those which theory demands. It is true
that relatively few experimental investigations can be controlled in
this way, but in those that can be checked by calculation (as, for
instance, in the well-known calorimetric experiments) everything
tends to show that precisely the same quantities of matter and
energy enter the body of an organism in the form of food-stuff, that
leave it as radiated and conducted heat, as work done, and as the
potential chemical energy of the excretions. Even when we are
unable (as in most investigations) to apply the test of
correspondence with theory, we have the conviction that the law of
conservation holds with all its strictness.
Then, whenever it was possible to apply the methods of chemistry
and physics to the study of the organism, it was seen that the
processes at work were chemical and physical. The substance of the
living body was seen to consist of a large (though limited) number of
chemical compounds, differing mainly from those which exist in
inorganic nature in their greater complexity. It was also seen that
physico-chemical reactions occurred in living substance analogous
with, or quite similar to, those which could be studied in non-living
substance. The conclusion, then, was irresistible that the life of the
organism was merely a phase in the evolution of matter and energy,
and differed in no essential respect from the physico-chemical
activities that could be observed in the non-living world.
These conclusions were stated so well by Huxley in his famous
lecture on “The physical basis of life,” over forty years ago, that all
subsequent utterances have been merely reiterations of this thesis in
a less perfect form. The existence of the matter of life, Huxley said,
depended on the pre-existence of certain chemical compounds—
carbonic acid, water, and ammonia. Withdraw any one of them from
the world and vital phenomena come to an end. They are the
antecedents of vegetable protoplasm, just as the latter is the
antecedent of animal protoplasm. They are all lifeless substances,
but when brought together under certain conditions they give rise to
the complex body called protoplasm; and this protoplasm exhibits
the phenomena of life. There is no apparent break in the series of
increasingly complex compounds between water, carbon dioxide, and
ammonia, on the one hand, and protoplasm on the other. We decide
to call different kinds of matter carbon, oxygen, hydrogen, and
nitrogen and to speak of their activities as their physico-chemical
properties. Why, then, should we speak otherwise of the activities of
the substance protoplasm?
“When hydrogen and oxygen are mixed in certain proportions and
an electric spark is passed through them they disappear, and a
quantity of water, equal in weight to the sum of their weights,
appears in their place. There is not the slightest parity between the
passive and active powers of the water and those of the oxygen and
hydrogen that have given rise to it. . . . We call these and many
other phenomena, the properties of water, and we do not hesitate to
believe that in some way they result from the properties of the
component elements of the water. We do not assume that a
something called “aquosity” entered into and took possession of the
oxide of hydrogen as soon as it was formed and guided the aqueous
particles to their places in the facets of the crystal, or among the
leaflets of the hoar frost.”
“Is the case in any way changed when carbonic acid, water, and
ammonia disappear, and in their place, under the influence of pre-
existing protoplasm, an equivalent weight of the matter of life makes
its appearance?”
“It is true that there is no sort of parity between the properties of
the components and the properties of the resultant. But neither was
there in the case of water. It is also true that the influence of pre-
existing protoplasm is something quite unintelligible. But does
anyone quite understand the modus operandi of an electric spark
which traverses a mixture of oxygen and hydrogen? What
justification is there, then, for the assumption of the existence in the
living matter of a something which has no representative or
correlative in the non-living matter which gave rise to it?”
All the investigations of over forty years leave nothing to be added
to this statement of what, in Huxley’s days, was called materialistic
biology. It was a very unpopular statement to make then, but it has
become rather fashionable now. Let the reader compare it with all
that has been spoken and written since 1869, even with the
utterances of the British Association of the year 1912, and he will
find that it expresses the point of view of mechanistic biology far
better than all the subsequent restatements. The only difference he
will find is that the latter have become (as William James has said
about academic philosophies), rather shop-soiled. They have been
reached down and shown so often to the enquiring public, that each
display has taken away something of their freshness.
Now Huxley’s example leads up so well to the consideration of the
differences between the chemical activities of the organism and
those of inorganic matter that we may consider it in some detail.
What, then, is the difference between the explosion of a mixture of
oxygen and hydrogen, and the photo-synthesis of starch by the
green plant?
In the case of the synthesis of water we have an example of an
exothermic chemical reaction. We are to think of the mixture of
oxygen and hydrogen as existing in a condition of “false equilibrium.”
It may be compared with a weight resting on an inclined plane.

Fig. 8.

Suppose that the plane is a sheet of smoothly polished glass, and

that the weight is a smooth block of glass. By canting the plane
more and more an angle will be found at which the slightest push
starts the weight sliding down. Now in the case of the explosive
mixture of oxygen and hydrogen we have a chemical analogue.
Either the gases do not combine at all at the ordinary temperature
or they combine “infinitely slowly.” But the slightest impulse, an
electric spark requiring an almost infinitesimally small quantity of
energy, starts the combination of the gases, and this continues until
all is changed into water vapour. In this reaction a large quantity of
energy is liberated in the form of heat. This heat becomes
transformed into the kinetic energy of the water particles which
condense from the steam formed in the explosion, and these
particles assume the temperature of their surroundings. The energy
which was potential in the explosive mixture, and which was capable
of doing work, still exists as the kinetic energy of the water formed,
but it has become unavailable for any natural process of work.
We have seen what is the general character of the reaction series in
the course of which carbon dioxide and water become starch; and
then this, becoming first soluble, and becoming associated with the
ammonia or nitrate taken into the plant, becomes protoplasm. It is a
reaction which differs from that just described, in that available
energy becomes absorbed and accumulated, and retains the power
of doing work. It is not a reaction which can be initiated by an
infinitesimal stimulus, but one in which just as much energy is
required in order that it may happen as is represented in the energy
which becomes potential in the living substance generated. The first
reaction is one which may take place by itself; 17 the other is one
which requires a compensatory energy-transformation in order that
it may happen. In the first reaction energy is dissipated; in the
second one it is accumulated.
We are thus led to the consideration of the second principle of
energetics and its limitations, but before entering upon this
discussion we must consider the nature of the activities of the
organism.
By the term “metabolism” we understand the totality of the physico-
chemical changes which occur in the living substance of the
organism. In physiological writings we usually find that two
categories of metabolic changes are described: (1) anabolic
processes, in the course of which simple chemical compounds
possessing relatively little energy are built up into much more
complex substances, containing a relatively large quantity of
available energy, and therefore capable of doing work. The
transformations constituting an anabolic change must be
accompanied by corresponding compensatory energy-
transformations, to account for the energy which becomes potential
in the substances formed. The formation of starch from carbon
dioxide and water, by the green plant, is such an anabolic change,
and the compensatory energy-transformation is the absorption of
radiation from the ether by the cells of the plant. A further anabolic
change in the plant organism is the formation of amido-substances
from the ammonia or nitrate absorbed from the soil, and from the
soluble carbohydrates formed from the starch manufactured in the
green cells.
The typical activities of the chlorophyll-containing organism are of
this nature; they are anabolic. The organism may be a green land-
plant; a marine green, red, or brown alga; a yellow-green diatom, a
yellow, green, red, or brown peridinian or other holophytic
protozoan; an ascidian, mollusc, echinoderm, polyzoan, worm, or
coral containing “symbiotic algæ” (that is the chlorophyll-containing
cells of some plant organism which have become associated with the
animal and incorporated in its tissues). In all these cases the
presence of this chlorophyllian substance confers on the organism
the power of effecting the compensatory energy-transformation, by
the aid of which carbon dioxide and water are built up into starch.
What this transformation is, and what are the steps by which the
carbon dioxide and water become carbohydrate we do not exactly
know. Solar radiation impinging upon an inorganic substance is
partly reflected and partly absorbed. The absorbed fraction may
become transformed in such a way as to render the substance
phosphorescent, or it may transform into chemical energy, as when
light impinges on a photographic plate, but as a general rule it is
transformed into heat. In the green plant, however, the
transformation of radiation into heat does not occur—at least the
heating is very small—and it passes directly or indirectly into the
potential chemical energy of the starch which is synthesised. We
must regard this power of absorbing radiation and utilising it in
compensatory transformations as a general character of protoplasm.
It is true that it is now specialised in the cells containing the
chlorophyll bodies, but there are indications that it may be present in
the tissues of the animal devoid of chlorophyll.
Other anabolic transformations occur in the animal. The food-stuffs
which are absorbed from the intestine are substances which have
undergone dissociations, the nature of which is such as to render
them capable of absorption and of reconstruction. These anabolic
changes in the higher animal are exceptional, and their usefulness
lies in the fact that by their means substances become capable of
being transported by the tissue fluids of the body.
(2) Katabolic changes in the animal body correspond in their
frequency of occurrence to the anabolic changes of the plant
organism. In them complex chemical substances undergo
transformation into relatively simple substances, and the contained
energy at the same time undergoes a parallel transformation,
passing into the form of heat and mechanical energy, while a
fraction becomes dissipated. Food-stuffs taken into the alimentary
canal break down in this way, but to a very limited extent. Proteids
undergo dissociation or decomposition into amido-substances, while
fats are dissociated into fatty acids and glycerine. Doubtless energy
is dissipated in these processes, serving no other purpose but to
heat the contents of the alimentary canal, but this energy-
transformation has not been worked out very completely and it is a
question whether, given a healthy animal and perfect food-stuffs,
any energy would necessarily be lost during the digestive processes.
The reactions involved in the latter do not belong to the category of
chemical changes proceeding from the complex to the simple, with a
liberation of energy; but appear to involve rather a rearrangement of
the constituents of a complex molecule, a process in which the
contained energy need not undergo change in quantity. These
processes involve the action of enzymes.
Enzymes play a great part in modern physiological theory and we
must consider them in detail. Let us attach a concrete meaning to
the general notion of enzyme-activity by considering the phenomena
known as catalysis. The metal platinum can be brought into a very
fine stage of division when it is known as platinum black. In this
condition it brings about reactions in chemical mixtures or
substances which would not otherwise occur: a mixture of oxygen
and hydrogen explodes when brought in contact with platinum
black, and a mixture of coal gas and air inflames, a reaction which is
made use of in the little gas-lighting apparatus which most people
have seen. If, again, a powerful electric current be passed between
platinum wires which are a little distance apart, and are immersed in
water, the metal becomes torn away from the points of the wire in
the form of an impalpable powder, colloidal platinum. The liquid
containing this colloid then has the power of setting up chemical
changes in other substances, changes which would not otherwise
occur, or, at least, would occur very slowly.
In general such catalysts, platinum black or colloidal platinum for
instance, have the following characters: (1) a small quantity is
sufficient to cause change in a large (theoretically an infinite)
quantity of the substance acted upon; (2) the nature and quantity of
the catalyst remain at the end the same, as at the beginning of the
reaction; (3) a catalyst does not start a reaction in any other
substance or substances, it can only influence the rate at which this
reaction may occur: apparently it does, in some cases, start a
reaction, but in such cases we suppose that the latter proceeds so
slowly as to be imperceptible; (4) the final state of the reaction is
not affected by the catalyst; it depends only on the nature of the
interacting substance or substances; (5) the final state is not
affected either by the nature or quantity of the catalyst: it is the
same if we employ different catalysts, or a large or small quantity of
the same catalyst. Finally, it appears that the phenomena of catalysis
are universal: “There is probably no kind of chemical reaction,” says
Ostwald, “which cannot be influenced catalytically, and there is no
substance, element, or compound which cannot act as a catalyser.” 18
Enzymes, then, are agents which are produced by the organism, and
which act by influencing (accelerating or retarding) chemical
reactions. An enzyme, as such, need not exist in a tissue; it is there
as a zymogen, a substance which may become an enzyme when
required. An enzyme need not be active: it may be necessary that it
should be “activated” by a kinase, another substance produced at
the same time. Associated with many enzymes are anti-enzymes,
substances which undo what their corresponding enzymes have
done. Finally some, perhaps most, enzymes are reversible, that is, if
they produce a change in a certain substance they can also produce
the opposite kind of change: the meaning of this will become clearer
a little later on. We have spoken of enzymes as “agents” or
“substances,” but it is not at all certain that they are definite
chemical compounds. In the preparation of an enzyme what the bio-
chemist obtains is a liquid, a glycerine or other extract which
possesses catalytic properties. An actual catalytic substance, like
platinum black, cannot be obtained from this liquid. A white powder
may be obtained, but this usually proves to be proteid in
composition; it is not the actual enzyme itself but is the impurity
associated with the latter. Now the very great number of enzymes
“isolated” by the physiologists has rather destroyed the original
simplicity of the idea of enzyme activity and suggests a parallel
statement to that made by Ostwald about catalysts: any tissue
substance may influence the reactions that may possibly occur in
other tissue substances. But while pure chemistry has to deal with
definitely known chemical compounds in the phenomena of catalysis,
this cannot be said to be the case with physiology in dealing with
enzymes. Reasoning by analogy, we may say that it is probable that
enzymes are definite proteids, or chemical substances allied to
these, but this has not been clearly demonstrated, and it is possible
that the phenomena of enzyme activity may belong to some other
category of energy-transformations.
However this may be, the conception is a useful one in describing
the reactions of the organism, and it may be illustrated by
considering the digestion and absorption of fat in the mammalian
intestine, a process which appears to be better known than that of
proteid digestion. A neutral fat consists of an acid radicle, oleic,
palmitic or stearic acids, for instance, united with glycerine. The
action of the pancreatic or intestinal enzymes is to dissociate this
fatty salt. Let us write the formula of the latter as G F, G being the
glycerine base, and F the fatty acid; then
GF G+F
which means that the enzyme can cause the neutral fat to dissociate
into glycerine and fatty acid. This action will go on until a state of
equilibrium is attained, in which there is a certain quantity of each of
the radicles, and a certain quantity of unchanged neutral fat, the
ratio of all these to each other depending on various things. When
this state of equilibrium is attained the enzyme does indeed go on
splitting up more neutral fat, but it is a reversible enzyme, and it also
causes the glycerine and fatty acid already split up to recombine,
forming neutral fat. A condition is, therefore, reached in which the
composition of the mixture remains constant.
Now there is dissociated fat in the intestine after a meal, but there is
only neutral fat in the wall of the intestine. The fat itself cannot pass
through the cells forming the intestinal wall, but the glycerine and
fatty acid into which it is dissociated can so pass, since they are
soluble in the liquids of the intestine. We suppose that the cells of
the wall of the intestine also contain the fat-splitting ferment; this
ferment in the cells acts on the glycerine and fatty acid immediately
they enter and recombines these radicles again into neutral fat, the
above equation now reading from right to left. But after a time this
reaction in the cells will also begin to reverse, for the enzyme will
begin to split up the synthesised neutral fat when the state of
chemical equilibrium in the new conditions is attained. Fatty acid and
glycerine will then diffuse out from the cells into the adjacent lymph
stream or blood stream—perhaps neutral fat will also pass from the
cells into these liquids, we are not sure. At all events the lymph and
blood after a meal containing much fat are crowded with minute fat
globules. But why are there no fatty acids or glycerine in the blood,
for the latter also contains lipase (the fat-splitting enzyme)? The
explanation is, apparently, that either an anti-enzyme is produced, or
that the enzyme passes into a zymoid condition. Why also does fat
accumulate in the tissues? Here, again, the activity of the enzyme,
which from other considerations we may regard as being universally
present almost everywhere in the body, must be supposed to be
arrested by some means.
The conception of a catalytic agent, such as we can study in pure
chemistry, thus carries us a long way in our description of the
processes of digestion, absorption, and assimilation. We have
applied it to the case of fat-digestion, but very much the same
general scheme might also apply to many other processes in the
body. Obviously it enables us to describe these processes in terms of
physico-chemical reactions, but we cannot fail to see that ultimately
we are compelled to assume the existence of reactions which were
not included in the original conception—the activation of the enzyme
at the proper moment by the kinase, the operation of the anti-
enzyme, and the passage of the enzyme into the zymoid. Just why
these things happen as they do we do not know, yet the whole
problem becomes shifted on to these reactions.
In the same way we apply the purely physical processes of the
osmosis and diffusion of liquids to the circulation of substances in
the animal body. The nature of these processes will probably be
familiar to the reader, nevertheless it may be useful to remind him
that by diffusion we understand the passage of a liquid, containing
some substance in solution, through a membrane; and by osmosis
the passage of a solvent (but not of the substance dissolved in it)
through a “semi-permeable membrane.” The molecules of the
solvent (water, for instance) pass through the membrane (the wall of
a capillary, or lymphatic vessel), but the molecules of the substance
(salt, for instance) dissolved in the solvent do not pass. Let us
suppose that a strong solution of common salt in water is injected
into the blood stream: what happens is that osmosis takes place, the
water in the surrounding lymph spaces passing into the blood
stream because the concentration of salt there is greater than it is in
the lymph. While this is happening, the capillary walls are acting as
semi-permeable membranes, allowing the molecules of water to
pass through but not the molecules of salt. Very soon, however, the
process of osmosis becomes succeeded by one of diffusion, and the
salt molecules pass through the capillary wall into the lymph and are
excreted.
Undoubtedly the purely physical processes of diffusion and osmosis
occur all over the animal body and are the means whereby food-
materials, secretory, and excretory substances are transported from
blood to lymph, or vice versa, from lymph to cell substance or to
glandular cavities, and so on. But it is also the case that in very
many processes the activity of the cells themselves plays an
important part. It may even be the case that a particular process,
after all physical agencies are taken into account, reduces down to
this action of the cells. To understand this we must consider the
mode of working of some well-known organ, and the best possible
example of such an organ, considered as a mechanism, is that of the
sub-maxillary salivary gland of the mammal.

Fig. 9.
What, then, is this mechanism and how does it act? The gland is a
compound tubular one, its internal cavity being prolonged into the
duct which opens into the mouth. The saliva prepared in the gland
issues from this duct. Blood is carried to the gland by twigs of the
facial artery, and, after circulating through it, is carried away by
factors of the jugular vein. Two nerves supply the gland: one is the
chorda tympani, a branch of a cranial nerve, and the other is a
sympathetic nerve. Lymph also leaves the gland by a little vessel.
Now suppose we have laid bare all this mechanism in a living animal
and make experiments upon it. If we stimulate the chorda tympani
there is a copious flow of thin watery saliva, but if we stimulate the
sympathetic there is a less copious flow of thick viscid saliva. Why is
this? We find on closer analysis that the chorda contains fibres which
dilate the small arteries so that there is an increased flow of blood
through the gland; but that, on the other hand, the sympathetic
contains fibres which constrict the arteries, thus leading to a reduced
flow of blood. This accounts for the fact that “chorda-saliva” is
abundant and thin, while “sympathetic-saliva” is scarce and thick. It
was thought at one time that the chorda contained fibres which
stimulated the gland to produce watery saliva, while the sympathetic
contained fibres which stimulated it to produce mucid saliva. This,
however, is not the case. Both nerves contain the same kind of
secretory fibres: their other fibres differ mainly in that they act
differently on the arteries.
It might be the case—indeed it was at one time thought that it was
the case—that secretion of saliva was simply a matter of blood-flow:
an abundant arterial circulation gave rise to abundant saliva, a
sparse flow to a sparse saliva. Undoubtedly the secretion depends
on blood supply, but not solely. If it did, then the whole process
might be conceived to be a very simple mechanical one—filtration or
diffusion of the saliva from the blood stream through the thin walls
of the blood vessels, and the walls of the tubules into the cavity of
the gland. If this were the case, then the liquid in the gland would
be the same in composition and concentration as the liquid part of
the blood—the plasma. But it is really different in composition and it
is not so concentrated. Now osmotic pressure—on the action of
which so much is based—cannot help us, for the liquid in the gland
is less concentrated than that in the blood vessels, so that water
ought to pass from gland to blood instead of from blood into gland.
Again, if we tie the duct, so that the saliva cannot escape, secretion
still goes on, though the hydrostatic pressure of saliva in the cavity
of the gland may be considerably greater than that of the liquid in
the blood vessels. Yet again, if we stop the blood flow by tying the
artery, secretion of saliva may still go on for a time.
Therefore the only physical agencies we can think of do not explain
the secretion. The latter is actually the work of the individual cells,
stimulated by the nerves. If the volume of the gland be measured
just while it is being stimulated to secrete, it will be found that the
organ becomes smaller, yet while it is being stimulated the blood-
vessels are being dilated so that the volume of the whole structure
ought to become greater. Obviously part of the substance of the
gland is being emptied out through its duct as the secretion.
If we examine the cells of the gland in various states we see clearly
that granules of some material, different in nature from the
substance of the protoplasm itself, are being formed within them.
Evidently these granules swell up during secretion and discharge
their contents into the ducts. Further changes in the characters of
the cell-substance, and in the nucleus, can be observed, and all
these indicate that the protoplasm of the cells, as the result of
stimulation, elaborates certain substances; that these substances are
then washed out, so to speak, into the duct by the withdrawal of
water from the cell; and that thereafter the cell absorbs fresh
nutritive material from the lymph which exudes from the blood
vessels, along with water. The distinctive part of the whole train of
processes is, then, this elaboration of material by the cells
themselves; while the concomitant changes in the calibre of the
blood vessels and in the flow of blood and lymph are subsidiary
ones. In the process of secretion of saliva energy is absorbed from
the chemical substances of the blood to bring about the passage of
water from a region of high to a region of low osmotic pressure;
oxygen and nitrogen, with other elements of course, are withdrawn
from the arterial blood stream for the purpose of the secretion, and
carbon dioxide and other substances are given off to the venous
blood and lymph.
The problem thus is pushed back from the mechanical events
occurring in the nervous and circulatory processes, to the physico-
chemical ones occurring in the cells of the gland tubules; and it thus
becomes much more obscure. It is true that we can formulate a
hypothesis which describes, in a kind of way, these intra-cellular
metabolic changes, in terms of physico-chemical reactions, and,
without doubt, reactions of this kind must occur within the cell. But
if we could test any such hypothesis as easily as the mechanical
ones suggested, should we find it any more self-sufficient? 19
Irritability and contractility are general properties of the organism.
These properties are illustrated by the irritability of an Amœba or
Paramœcium to stimuli of many kinds; by the movements of the
pseudopodia of the former animal, or of the cilia of the latter; by the
nervous irritability of the higher animal, and the contraction of its
muscles when they are stimulated. They are among the fundamental
properties or functions of living protoplasm, and their study is of
paramount interest, and carries us to the very centre of the problem
of the activities of the organism. Naturally physiologists have never
ceased to attempt to describe irritability and contractility in terms of
physics, but though we may be quite certain that the things that do
occur in these phenomena are controlled physico-chemical reactions,
it must be remembered that what we positively know about their
precise nature is exceedingly little.
What is the nature of a nervous impulse? When a receptor organ is
stimulated, as, for instance, when light impinges on the cone cells of
the retina, or when the nerve-endings in a “heat-spot” in the skin
are warmed, or when the wires conveying an electric current are laid
on a naked nerve, an impulse is set up in the nerve proceeding from
the place stimulated, and we must suppose that approximately the
same amount of energy moves along the nerve as was
communicated to the receptor or the nerve itself by a stimulus of
minimal strength. How does it so move? Several facts of capital
importance result from the experimental work. (1) The impulse
travels with a velocity variable within certain limits, say from 8 to 30
metres per second; (2) it travels faster if the temperature is raised
(up to a certain limit); (3) it is difficult to demonstrate that the
passage of this impulse is accompanied by definite chemical changes
in the nerve substance: it is stated that carbon dioxide is produced,
but this is not certainly proved; (4) an electric current is produced in
the nerve as the result of stimulation; (5) no heat is produced, or at
least the rise of temperature, if it occurs, is less than 0.0002° C.
Thus it is quite certain that physical changes accompany the
propagation of the nerve-impulse, for the latter has a certain
velocity, which depends on the temperature, and an electric change
also occurs in the substance of the nerve. Is this electric change the
actual nerve impulse? It is hardly likely, since the velocity of the
impulse is very much less than that of the propagation of an electric
change through a conductor; besides, the passage of the impulse is
not accompanied by a measurable heat evolution, although the flow
of electricity along a poor conductor must generate heat and
dissipate energy. Is it a chemical change? Then we should be able to
observe metabolism in the nerve substance—that is if the energy-
change is a thermodynamic one—while it is not at all certain that
metabolic changes do occur. Nevertheless it seems probable that a
physico-chemical change is actually propagated when we consider
the chemical specialisation of the substance of the axis-cylinder of
the nerve. Now the velocity of propagation of the nervous impulse is
of the same order of magnitude as that of an explosive change in
chemical substances (using the term “explosion” to connote
chemical disintegrations rather than combustions). If we imagine a
long rod of dynamite, or picric acid, or a long strand of loosely-
packed gun-cotton to be exploded by percussion at one end, then a
transmission of the chemical disintegration of any of these
substances will pass along the rod, etc., with a velocity which will
certainly vary with the physical condition of the material. It would be
a high velocity in a rod of dynamite, or fused picric acid, but a lower
velocity in a loosely aggregated strand of gun-cotton, or a trail of
picric acid powder. Is this what happens in the nerve when an
impulse travels along it? Obviously not, since the substance of the
nerve is not altered appreciably, while that of the explosive
substance passes into other chemical phases. We might imagine,
then, such a change in the nerve fibrils as that of a reversible
transformation of some chemical constituent:—
(2) (1)
:a + b:a + b:a + b:a + b:a + b:
: : : : : :
:c + d:c + d:c + d:c + d:c + d:
Let us imagine the substance of the fibril to be composed of, or at
least to contain, the substances a + b which dissociate reversibly
into the substances c + d. At any moment, and in any particular
physical state, as much of a and b pass into c and d as c and d pass
into a and b. There will be equilibrium. But now let a stimulus alter
the physical conditions: prior to the stimulus the phase was am + bn
= cp + dr—the suffixes m, n, p, r, denoting the concentrations of a,
b, c, and d—but after the stimulus the phase may be am1 + bn1 =
cp1 + dr1. Now the element of the nerve substance (1) forms a
system with the element (2). The condition in (2) is am + bn =
cp + dr, and that of (1) am1 + bn1 = cp1 + dr1, but these two
together now fall into a new state of equilibrium and this is
transmitted along the whole nerve-fibril with a velocity which
belongs to the order of magnitude of that of chemical changes. If
the stimulus remains constant (a constant electric current for
instance), the new condition of equilibrium will be established
throughout the whole length of the fibril and the nervous impulse
will be a momentary one (as it is in this case). But if the stimulus is
an intermittent one (an interrupted electric current, light-vibration,
sound-vibrations), then in the intervals the former condition of
equilibrium will become re-established and the nervous impulse will
be intermittent (as it is). There would be no work done on the whole
in the changes, except that done by the transmission of the changed
state of equilibrium to the substance of the effector organ in which
the nerve-fibril terminates—the substance of a muscle fibre, or the
cell of a secretory gland, for instances. There would, probably, be a
certain dissipation of energy as in the case of the propagation of an
electric impulse through a poor conductor, but all our knowledge of
the chemistry of the nerve fibre points to this amount of dissipation
as tending to vanish.
Something analogous to this may be expected to take place in a
muscle fibre when it contracts; except that, of course, energy is
transformed in this case. What precisely does happen we do not
know and at the present time no physico-chemical hypothesis of the
nature of muscular contraction exactly describes all that can be
observed to take place. Certain positive results have, of course, been
obtained by chemical and physical investigation of the contracting
muscle: carbon dioxide is given off to the lymph and blood stream,
and the amount of this is increased when an increased amount of
work is done by the muscle; heat is produced and this too increases
with the work performed; glycogen is used up, and lactic acid is
produced; finally oxygen is required, and more oxygen is required by
an actively contracting muscle than by a quiescent one. Now the
obvious hypothesis correlating all these facts is that the muscle
substance is oxidised, and that the heat so produced is transformed
into mechanical energy. “We must assume,” says a recent book on
physiology, “that there is some mechanism in the muscle by means
of which the energy liberated during the mechanical change is
utilised in causing movement, somewhat in the same way as the
heat energy developed in a gas-engine is converted by a mechanism
into mechanical movement.”
Now, must we assume anything of the kind? To begin with, life goes
on, and mechanical energy is produced in many organisms living in a
medium which contains no oxygen. Anaerobic organisms are fairly
well known, and we cannot suppose that in them energy is
generated by the combustion of tissue substance in the inspired
oxygen. A muscle removed from a cold-blooded animal will continue
to contract in an atmosphere containing no oxygen, and it will
continue to produce carbon dioxide. It is true that the contractions
soon cease, even after continued stimulation under conditions
excluding the fatigue of the muscle, but do the contractions cease
because the oxygen supply is cut off, or because the muscle dies in
these conditions? We know that some complex chemical substance is
disintegrated during contraction and that mechanical energy and
heat are produced and that carbon dioxide is also produced. We
know that the carbon contained in the latter gas corresponds
roughly with the carbon contained in the muscle substance which
undergoes disintegration, but does all this justify us in saying that
this substance is oxidised in order that its potential chemical energy
may be transformed into mechanical energy? Obviously not, since
we might equally well suppose that the complex metabolic substance
of the muscle splits down into simpler substances and that in this
transformation energy is generated. Suppose that these simpler
substances are poisonous and that they must be removed as rapidly
as formed. The rôle of the oxygen may be to oxidise them, thus
transforming them into carbon dioxide, an innocuous substance
which can be carried away quickly in the blood stream. This line of
thought, according to which the rôle of oxygen is an anti-poisonous
one, is held at the present day by some physiologists, and many
considerations appear to support it; the existence of “oxidases,” for
instance, enzymes which produce oxidations which would not
otherwise occur in their absence. Such enzymes exist in very many
tissues, and they may, apparently, be present in an inactive form,
requiring the agency of a “kinase” before they are able to act.
The usual view among physiologists is that the muscle fibre is a
thermodynamic apparatus transforming the heat generated during
metabolism into mechanical energy. How is this transformation
effected? It cannot be said that we have any one hypothesis more
convincing than another. It has been suggested that alterations of
surface tension play a part, or that the heat produced by oxidation
causes the fibre to imbibe water and shorten. Engelmann has
devised an artificial muscle consisting of a catgut string and an
electrical current passing through a coil of wire, and by means of
this he has reproduced the phenomena of simple contraction and
tetanus. But it remains for future investigation to verify any one of
these hypotheses.
When Huxley published his Physical Basis of Life, probably few
physiologists had any doubt that protoplasm was a definite chemical
substance, differing from other organic substances only by its much
greater complexity. But in 1880 Reinke and Rodewald published the
results of an analysis of the substance of a plant protoplasm and
these appear to have demonstrated that the substance was really a
mixture of a number of true chemical compounds and was not a
single definite one. Now all of these substances might exist apart
from protoplasm, and in the lifeless form, and a simple mixture of
them could hardly bring forth vital reactions. These results were
followed by the morphological study of the cell—the discovery of the
architecture of the nucleus, and so on, and so opinion began to turn
to the hypothesis that the vital manifestations of protoplasm were
the result of its structure. Microscopical examination of the cell
appeared to disclose a definite arrangement, the “foam” or “froth” of
Butschli, for instance. But, again, it was easily shown that the foam,
or alveolar structure of protoplasm was merely the expression of
physical differences in the substances composing the cell-stuff—they
reduced to phenomena of surface tension and the like. Artificial
protoplasm and artificial Amœbæ were made—at least mixtures of
olive oil and various other substances were made which simulated
many of the phenomena of protoplasm in much the same way as
crystalline products may be made which simulate the growth of a
plant stem with its branches. For instance, one has only to shake up
a little soapy water in a flask to see what resembles surprisingly the
arrangement of certain kinds of connective tissues in the organism.
Obviously these artificial phenomena have nothing to do with living
substance.
Yet if we grind up a living muscle with some sand in a mortar we do
destroy something. The muscle could be made to contract, but after
disintegration this power is lost. We have certainly destroyed a
structure, or mechanism, of some kind. But, again, the paste of
muscle substance and sand still possesses some kind of vital activity,
for with certain precautions it can be made to exhibit many of the
phenomena of enzyme activity displayed by the intact muscle fibres,
or even the entire organism. Mechanical disintegration, therefore,
abolishes some of the activities of the organism, but not all of them.
If, however, we heat the muscle paste above a certain temperature,
the residue of vital phenomena exhibited by it are irreversibly
removed, so that heating destroys the mechanism. This we can
hardly imagine to be the case (within ordinary limits of temperature
at least) with a physical mechanism, but again a mechanism which is
partly chemical might be so destroyed. We see, then, that
protoplasm possesses a mechanical structure, but that all of its vital
activities do not necessarily depend on this structure. The full
manifestation of these activities depends on the protoplasmic
substance possessing a certain volume or mass, and also on a
certain chemical structure.
If living protoplasm has a structure, and is not simply a mixture of
chemical compounds, what is it then? Two or three physico-chemical
concepts are at the present time very much in evidence in this
connection. When the substances known as colloids were fully
investigated by the chemists, much attention was paid to them by
the physiologists, so that life was called “the chemistry of the
colloids,” just as after the investigation of the enzymes it was called
the “chemistry of the enzymes,” and when the discovery of the
relative abundance of phosphorus in cell-nuclei and in the brain was
discovered, it was called the “chemistry of phosphorus.” Colloids
(e.g. glue) are substances that do not readily diffuse through certain
membranes, in opposition to crystalloids (e.g. solution of common
salt) which do readily so diffuse. They form solutions which easily
gelatinise reversibly, that is, can become liquid again (glue); or
coagulate irreversibly, that is, cannot become liquid again
(albumen); which have no definite saturation point; which have a
low osmotic pressure (and derived properties), etc.; and the
molecules of which are compound ones consisting of combinations
of the molecules of the substance with the molecules of the solvent,
or with each other, that is, they are molecular aggregates.
Colloids pass insensibly into crystalloids on the one hand and into
coarse suspensions (water shaken up with fine mud, for instance) on
the other. We may replace the concept of a colloid by those of
“suspensoids” and “emulsoids.” A suspensoid is a liquid containing
particles in a fine state of division—if the division is that into the
separate molecules we have a solution, if into large aggregates of
molecules we have a suspension. If the substance in the liquid is
itself liquid, the whole is called an emulsoid. On the one hand this
approaches to a mixture of oil in soap and water—an emulsion—and
on the other hand to such a mixture as chloroform shaken up with
water, when the drops of chloroform readily join together so that
two layers of liquid (chloroform and water) form. What we see, then,
in protoplasm is a viscid substance possessing a structure of some
kind, and containing specialised protoplasmic bodies in its mass
(nuclei, nucleoli, granules of various kinds, chlorophyll, and other
plastids, etc.). It may contain or exhibit suspensoid or emulsoid parts
or substances, or it may contain truly crystalloid solutions. These
phases of its constituents are not fixed, but pass into each other
during its activity. Nothing that we know about it justifies us in
speaking about a “living chemical substance.” On analysis we find
that it is a mixture of true chemical substances rather than a
substance. It is no use saying that in order to analyse it we must kill
it, for what we can observe in it without destroying its structure or
activities indicates that it is chemically heterogeneous.
This is not a textbook of general physiology, and the examples of
physico-chemical reactions in the organism which we have selected
have been quoted in order to show to what extent the chemical and
physical methods applied by the physiologists have succeeded in
resolving the activities of the organism. The question for our
consideration is this: do these results of physico-chemical analysis
fully describe organic functioning? Dogmatic mechanism says “yes”
without equivocation.
Now it is clear, from even the few typical examples that we have
quoted, that physiological analysis shows, indeed, a resolution of the
activities of the organism into chemical and physical reactions. How
could it do otherwise? How could chemical and physical methods of