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THE ELDRIDGE REEVES JOHNSON
FOUNDATION FOR MEDICAL PHYSICS
ADVENTURES IN BIOPHYSICS
ADVENTURES IN
Biophysics
By
Α . V. Hill
SC.D., LL.D., M.D., F.R.S.
Foulerton Research Professor of the Royal Society
Philadelphia
U N I V E R S I T Y OF PENNSYLVANIA PRESS
L O N D O N : H U M P H R E Y MILFORD: OXFORD U N I V E R S I T Y PRESS
1 1
9 3
Copyright 1931
UNIVERSITY OF PENNSYLVANIA PRESS
Printed in the United States oj America by
Lancaster Press, Inc., Lancaster, Pa.
PREFACE
If you go to Devonshire in September and wander in the

lanes you saw at the end of March you will find it hard to
believe that they are in fact the same. Reading now in
February the proofs of the lectures given at Philadelphia
in October I am equally astonished at the changes which a
few short months have wrought in our outlook. It is
fortunate to be concerned with a field of research in which
one's colleagues are so active, but it makes it no easier to
describe the present, or any other outlook. The last three
years, indeed, have brought strange and precious growth
in subjects which seemed to be reaching maturity.
I have avoided any detailed alterations in the picture
piesented in the lectures themselves, and have been content
to show, in somewhat numerous footnotes, so far as I am
able, the changes which have occurred in the meantime.
No doubt the picture will be old-fashioned again in six
months: but still, when the primroses return it may help
to explain how they once gave place to blackberries, how—
for example and for a season—lactic acid was replaced by
phosphagen, equilibrium by steady state, bound water by
free.
Α. V. HILL.
UNIVERSITY COLLEGE, LONDON,
February, 1931.
V
CONTENTS
Page
PREFACE Ν
INTRODUCTION ix
LECTURE I. Some Adventures with Vapour Pres-
sure ι
" II. The State of Water in Tissues 29
" III. The Conception of the Steady State. . 55
" IV. The Time-Relations of Events in Mus-
cular Contraction 80
" V. The Mechanics of Muscular Contrac-
tion and Other Matters 116
APPENDIX I. A Discussion of the Several Instru-
mental Factors Involved in the
Attainment of the Greatest Possible
Sharpness of Analysis of the Heat
Production of Muscle 139
A P P E N D I X II. The Effect on the Calibration of Non-
uniformity of Cross-section of Mus-
cle 144
A P P E N D I X III. The Flow of Heat in a Plane Sheet of
Muscle 146
BIBLIOGRAPHY 149
INDEX 159
vii
INTRODUCTION
' T ^ W E N T Y summers ago, on Sunday mornings, certain
·*· young Cambridge physiologists used to go and dig in
Gaskell's garden on the Gog Magog hills, while he regaled
them with histories of the great things he had seen, and
the great days he had spent in Ludwig's laboratory and
elsewhere. These stories supplied, what no paper in a
scientific periodical can supply, a vivid picture of the
human side of scientific research. In our journals we try,
so far as we can, to present a concise and logical account of
our alleged discoveries. The real reasons why we did the
things we did, the delays and imperfections and per-
plexities which beset us, the misery of continual failure,
the joy of occasional success, the faith that with patience
and persistence we should find the unknown something we
were sure was there—all these are unfitting in a scientific
periodical, yet somewhere a hint at least of them should
be recorded. Such lectures as these are not intended to
take the place of reviews or of articles in abstracting
journals: those are much better read than heard. I hope,
therefore, that Professor Bronk will persuade my successors,
as he has persuaded me, to tell you of their scientific
adventures: for so, firstly I shall be in good company, and
secondly you will see better into their minds, you will
realize that for you as for them the pursuit of natural
knowledge may be one of the great adventures of the
human spirit.
I t was Borelli two hundred and fifty years ago who
affirmed that the study of the motion of animals, no less
than Astronomy, is a part of physics, to be enlarged and
adorned (note the word " a d o r n e d " ) by mathematical
demonstrations. The Johnson Foundation for Medical
Physics, in which I have the honour to inaugurate this
annual Series of lectures, is intended to fulfil Borelli's pre-
cept. I wish it good fortune in its enviable task: may it
adorn, as well as extend, the fields of biological and medical
knowledge!
ix
L E C T U R E I
SOME ADVENTURES WITH VAPOUR PRESSURE
N the spring and summer of 1927, while I was at Cornell,

I my colleague and assistant Mr. A. C. Downing was
busy in London with a new type of muscle thermopile,
which he was preparing against my return. This instru-
ment exceeded all our expectations and made it possible
for the first time to measure, with reasonable accuracy, the
rate of resting heat production of a muscle lying on it; it
came into use just three years ago, in October, 1927. I
need not describe it in detail now: it is sufficient to say that
by greatly improved electrical insulation of the wires and
by the use of material of high thermal conductivity for
the framework, it avoided the troublesome zero-errors, and
the differences of temperature within the instrument, which
had dogged the footsteps of previous myothermic observa-
tions. (See fig. ι and Hill 1928a.) A resting muscle
placed upon it and soaked in Ringer's solution for an hour or
two, g a v e — w h e n the solution was replaced by oxygen or
nitrogen—steadyreadings which agreed well with Meyerhof's
recorded observations of the oxygen consumption, or of the
lactic acid formation, under aerobic or anaerobic conditions
respectively. (See Hill 1928 b.) This, with its very con-
stant zero, made measurements of total heat, after stimula-
tion in oxygen or in nitrogen, far more accurate than any
previously possible.
T h e object of the experiments undertaken was to test,
by the myothermic method, a claim advanced from Emb-
den's laboratory in Frankfurt (see Embden, Hirsch-
Kauffmann, Lehnartz and Deuticke (1926); Embden, Leh-
nartz and Hentschel (1927)), that in a series of muscle
twitches a considerable fraction of the lactic acid set free ap-
pears some time after the mechanical response is past. This
1
SILVER FRAME
Side
S I L V E R FRAME
front
FIG. I. Thermopile for frog's sartorius. Scale refers to centre and right only
(a) Vulcanite carrier; (b) brass terminals to electrodes; {c) copper terminals to
thermopile; (d) brass tube for thread to muscle; (e) glass tube carrying electrode
leads; (/) glass tube carrying thermopile leads; (g) rubber stopper; (A) glass cover;
(<) brass carriage; ( J ) silver frame; ( t ) muscle; (/) thermopile; (m) electrodes; (n)
muscle clamp; (o) brass carriage; (p) clamping screw for {q) vulcanite rod; (r) brass
rods and nuts; (s) " E l o " bushes; (/) hot junctions; (a) cold junctions; (o) wire
windings; (to) gas exit-pipe; (*) gas inlet-pipe. (Hill 1928a.)
2
VAPOUR PRESSURE
effect was held by Embden and his colleagues to disprove

what they were pleased to call the " Hill-Meyerhof theory"
of muscular contraction. The facts were contested by
Meyerhof and Lohmann (1926), and by Meyerhof and
Schulz (1927), and, even if true, some of them did not prove
what they were supposed to prove. (See Hill i928d.) The
question, nevertheless, was important, and, assuming lactic
acid formation to be accompanied by heat production,
could be tested easily by the myothermic method. The
ratio of tension to initial heat was measured in nitrogen, in
a single isometric twitch: and the ratio of Σ (tensions) to
total heat, in a series of isometric twitches. If these ratios
were equal we could only conclude that in nitrogen initial
heat is equal to total heat. Within reasonable limits of
error they were equal. Therefore, in the series of anaerobic
twitches all the heat was initial heat: if there was delayed
lactic acid formation it was not accompanied by heat pro-
duction.1
This particular matter will be dealt with further in my
fourth lecture. For the moment I wish to refer to a curious
by-product of these observations which remained unex-
plained, and produced various fantastic hypotheses, for
nearly two years. Finally it led to an admirable method of
measuring the depression of vapour pressure of an aqueous
solution (Hill 1930a) by which, for example, the molecular
weight of the solute in a few drops of solution can be deter-
mined within ι p.c. or less, and by which—of all things—
my colleague, Dr. Margaria (1930), has been able to estab-
lish the existence of a perfectly definite difference of os-
1
Strange to relate, it now appears that there it delayed lactic acid formation,
after all, at least following a tetanic contraction, the heat of which is largely balanced
by the negative energy of phosphagen resynthesis. If there be delayed lactic acid
formation also after a twitch—of which at present there is no direct evidence—then
its energy is exactly balanced by that of phosphagen restoration. See below and
Lecture IV. [ A d d e d March 1931· By improved methods Cattell and Parkinson
have found delayed heat after a series of twitches in nitrogen, amounting to about
10 p.c. of the initial heat.]
3
A D V E N T U R E S IN BIOPHYSICS
motic pressure between the bloods of men and women!

The observation itself was as follows (Hill 1928b).
The galvanometer connected to a thermopile, on the
" h o t " junctions of which a muscle lies, the muscle having
been previously soaked for an hour or two in isotonic
physiological salt solution, shows a steady deflection cor-
responding to the constant rate of heat production of the
muscle. In oxygen the deflection is about twice as great
as in nitrogen: the completion of the oxidative cycle gives
about as much heat as the initial anaerobic phase. If the
muscle be stimulated to a series of twitches the galva-
nometer immediately deflects. In oxygen it remains de-
flected for a long time, owing to the onset of the recovery
heat (Lecture I V , fig. 1 2 ) : in nitrogen, when the stimuli end,
it returns rapidly towards its initial zero. Towards but
not to, that was the phenomenon. However long one
waited the galvanometer never came back to its original
position: it remained deflected, by an amount directly
proportional to the energy liberated by the muscle during
the previous stimulation (fig. 2). There could be only one
conclusion, namely that the rate of anaerobic resting heat
production had been increased by previous activity, to an
extent proportional to that activity. The experiment was
an easy one to make, it was repeated hundreds of times;
yet the result seemed so incredible that, for consolation
and support, I invited various of my friends to witness it—
who agreed at least that the phenomenon was not a product
of my imagination!
All who have thought at all of the problems of general
physiology must have exercised their minds as to the reason
why a living cell, completely at rest, and doing nothing at
all except maintain its continued existence, requires a con-
tinual supply of energy. Usually, in animal cells, this
energy is obtained from the oxidation of foodstuffs: in the
absence of molecular oxygen, either some substance con-
taining combined oxygen is broken down to yield it (as in
4
VAPOUR PRESSURE
200
J-0* .
j-o"
,0- >»
160
/•o •i jrf
Ό
***
0-75"o?s~
120
075 i^f *
0-75
6/772. cm. 4000

Heat 8000 12000 16.000 2QP00
verGm.
FIG. 2. Linear relation between anaerobic resting heat-rate and total heat
liberated by a succession of tetanic stimuli at ίο-minute intervals. T e t a n u s dura-
tion shown in seconds. (Hill 1928^.)
nerve), or some anaerobic process such as the formation of

lactic acid from carbohydrate supplies the energy directly.
When further energy is unobtainable the cell dies. There
are two ways of looking at the phenomenon, the dynamic
and the structural. These are not necessarily different,
but they start from different points of view.
We speak of living matter: there is, as I said at a recent
Discussion on Colloid Science at Cambridge (Hill i93od),
a certain danger in the term, for it appears to imply that
any given identical piece of matter may be, and may
continue to be, "alive." It emphasises the matter rather
than the process. To take a physical analogy, a wave is a
sequence of events in matter—or space—depending on the
properties of that matter or space: we should, however,
only obscure the study of waves were we to attribute their
characteristics to the specific properties of an "oscillatory
matter" or an "oscillatory space." There is nothing
peculiar about the matter—or space—in which waves are
travelling: the peculiarity is in the sequence of events
5
2
making up the waves. So also in biology: " l i f e " in the

physico-chemical sense must be regarded as a self-per-
petuating and generally periodic complex of events, recog-
nisable by its results, not by its outward appearance at a
given moment, occurring indeed in a medium of matter,
depending absolutely on the properties of matter, but as
distinct from matter as music is from the air in which it is
propagated. From the dynamic point of view the energy
required by a resting cell is used in " d r i v i n g " the sequence
of invisible events which make up life. T o break the
continuity of these events for more than the shortest time
is like wetting a short length in the powder train of a fuse:
normal reactions may cease altogether to be propagated,
and " l i f e " may end. There must be some element of
truth at least in this way of looking at things.
T h e other point of view emphasises the known, or the
imagined but invisible, structural elements of the living
cell. There is no doubt, whatever the mechanism of their
maintenance, of the continued existence of surfaces of
separation (or of dynamic processes acting like surfaces
of separation) in the living cell. L i v i n g tissue, kept
aseptically after removal from an animal, gradually breaks
down in a way unknown during life. In muscle, glycogen
becomes lactic acid, " p h o s p h a g e n " yields creatine and
phosphate, various changes take place in the several
phosphorus compounds, and probably as yet unknown
reactions occur (see, e. g., Meyerhof 1930a). Rigor mortis
ultimately sets in, and then the tissue slowly and gradually
breaks up, in the process known as " a u t o l y s i s . " All this
can be prevented, almost indefinitely, by supplying oxygen
and foodstuff's.
In the living cell there are, undoubtedly, many highly
reactive substances, which proceed to react as soon as they
are given the opportunity. T h e opportunity may come,
either (1) when the cell is injured by mechanical or chemical
means, when in fact the surfaces of separation are positively
6
VAPOUR PRESSURE
broken: the rapidity of lactic acid formation in injured

muscle is a notorious example: or (2) when, by oxygen
want or by deprival of energy-liberating substances, the
surfaces of separation can no longer be maintained in their
normal and orderly state. One is led therefore to think of
oxidation in the healthy resting cell as employed in main-
taining a certain dynamic status quo, in which highly
reactive substances are confined to their proper spheres of
action, hindered—so to say—from unsocial conduct, utilized
in an organised scheme of things instead of being per-
mitted to behave at random. The need of an oxidative
maintenance of an orderly and cooperative system may be
due simply to the place of oxidation in a normal sequence
of events—here we return rather to the dynamic point of
view: or, the oxygen may be used to liberate energy by
which surfaces and interfaces are held in their normal,
polarized, impermeable state. We may think of a house,
or perhaps better of a business or a city, in which the
normal architecture, the normal services, and the normal
economic organization can be kept up only by continual
expenditure. So long as these exist, ordinary life is possible:
the inhabitants can go about their lawful business. Re-
move, however, the normal services and restraints, and
chaos rapidly sets in, all kinds of reactions occur which
are usually prevented, and recovery may become slow or
impossible.
These two hypotheses of the function of oxidation in
maintaining the steady state of the normal resting cell had
long been in my mind. Could it be that in the peculiar
increase of the rate of resting heat production of a muscle,
caused by anaerobic stimulation, we were witnessing the
first stages in the breakdown of the living muscle cells?
The greater the extent of activity induced by stimulation
the greater would be the oxygen want, the greater the
reducing power of the substances set free, the greater the
degree to which the surfaces and interfaces of the living
7
cells were p u t out of action. If this were so, the increment

in heat rate induced by anaerobic stimulation might be a
sign of the chaotic reactions m a d e possible by the with-
drawal of the normal restraining influence of surfaces, inter-
faces, or dynamic processes m a i n t a i n e d so long as oxygen
was present.
Looked at in perspective now, with the true and simple
explanation of the phenomenon in mind, the hypothesis
m a y appear fantastic. At the time no other seemed
possible, nor (I would plead in extenuation) could any of
my friends suggest a better. I t was not lightly proposed,
and m a n y tests and experiments were made before it was
m a d e public. Firstly one h a d to be sure t h a t it was not
due to some defect of the i n s t r u m e n t s employed. Various
chemical substances, particularly lactic acid, are produced
by stimulation: could the effect be a galvanic one, induced
for example by acid diffusing through the insulating
material and reacting differently with the two metals of the
thermopile? T h e thermopile was most carefully insulated
with bakelite, shellac and paraffin wax: to avoid a n y
possible effect of a diffusing substance a strip of tin-
foil was introduced between the insulating layers: the
phenomenon appeared u n c h a n g e d : it seemed impossible
t h a t any reasonable chemical substance could pass through
such protection. Moreover filling the chamber with C 0 2
would make the muscle as acid as extreme fatigue. I t
caused only a very slight increment in resting heat rate. If
the effect were due to chemical reactions occurring in the
muscle it should have a high t e m p e r a t u r e coefficient: t h e
increment in heat rate increased a b o u t 2.5 times for a rise
of io° C.: a physical error would probably be only slightly
affected by temperature. M o r e o v e r — a n d here was some-
thing certainly significant—the effect might be almost
entirely abolished by allowing the muscle to recover in
oxygen. An example, illustrative only, m a y make this
clearer.
8
VAPOUR PRESSURE
Fresh muscle initially in N 2 : resting heat rate,

galvanometer scale divisions 20
Muscle fatigued in N 2 : final resting heat rate. 200
0 2 introduced: muscle allowed to recover:
final resting heat rate in 0 2 50
N 2 introduced: resting heat rate 25
What could this be other than an effect of oxygen in

restoring the normal status quo in the muscle, in reinstating
those surfaces or agents which maintain an orderly sequence
of events and prevent a biochemical chaos? How other-
wise could we find a smaller heat production in oxygen
than in nitrogen? The same reactions running to com-
pletion in the one would scarcely yield less heat than ending
half way in the other. Clearly oxygen was inhibiting some
unknown reactions. W h a t should those reactions be other
than the ones which, in the absence of oxygen, are known
to lead to the death and disintegration of the cell?
The experiments could not be denied: the hypothesis—
fantastic as even then it seemed—was the only one which I,
or any of my friends, could propose, and an account was
published in the early summer of 1928 (Hill 1928b). This
publication had a most peculiar effect, which some of you,
even in America, may have noted. M y colleague Professor
Donnan had rashly promised to give a public lecture at the
British Association on " T h e Mystery of L i f e " (Donnan
1928). In this, having read my paper, and wishing to
refer to the role of oxygen in maintaining the normal life
and architecture of the cell, he made a few incautious and
far too flattering remarks about my work, with the result
that the daily Press descended like an avalanche on me as
I was taking my summer holiday in Devon. I was charged
with creating life (or alternatively with creating souls 2 )
in the laboratory, I was invited to raise the dead (in
America) by means of oxygen and to write half a dozen
* Souls and cells sound much the same to a newspaper reporter.
9
books. M y friend Professor Meyerhof heard about it

from two old ladies—strangers to him—travelling in a
train in Switzerland, and wondered what had befallen me.
In Frankfurt doubtless they attributed it to the imperfec-
tions of the " Hill-Meyerhof theory." Y o u see why I have
called this lecture " Some adventures with vapour pressure."
B y the autumn of 1928 difficulties began to appear. T h e
existence of an increment in resting heat rate after anaerobic
stimulation could not be confirmed by Meyerhof (Meyerhof,
McCullagh, and Schulz (1930)) by experiments with masses
of muscle in a calorimeter. The increment in resting heat
rate could be abolished, not only by allowing the muscle
to recover in oxygen, but by washing it with Ringer's
solution completely free from oxygen. On the other hand
the phenomenon itself was confirmed in every way, even
with a delicate resistance thermometer to replace the
thermopile: it had nothing to do with the particular
instrument employed. Finally in the summer of 1929 a
most preposterous thing happened—the increment in heat
rate due to anaerobic stimulation was found to be several
times as great in hydrogen 3 as in nitrogen (Hill 1929c).
It was obvious that a muscle could not be using hydrogen
for its metabolism—some physical effect must underlie the
observation. It still took me several days to find the
answer to the riddle—so stupid and slow are physiologists
in solving physical problems, so urgently do we need places
like the Johnson Foundation where we can get sympathetic
help!
Years before I had read a paper by J . S. Haldane
(Haidane, Kellas and K e n n a w a y ( 1 9 1 5 ) ) in which he
showed that the human subject can better stand a given
low partial pressure of oxygen in rarefied air than in a gas
' Hydrogen had been previously avoided, as the thermopiles were hollow and
it was feared that the gas would diffuse through the insulator and blow them out.
This actually happened later when tried. T h e experiments with hydrogen were
made with a special instrument which was not hollow.
10
VAPOUR PRESSURE
mixture at normal atmospheric pressure. Oxygen diffusion

is more rapid in a mixture of lower density. Could some-
thing be diffusing towards my muscle more rapidly in
hydrogen than in nitrogen? On a warm night in July,
suddenly and from nowhere, the answer came: water
vapour. A book of physical constants gave one immediate
assurance that the phenomenon could be explained by the
transfer just of a few milligrammes of water per day,
evaporating from the walls and condensing on the muscle
with an evolution of heat. A couple of experiments next
morning made it certain that at last the riddle was solved.
Not one of the long line of physiologists, from Helm-
holtz downwards, who have applied the myothermic method
had ever paid any attention to the vapour pressure of the
muscles they used. I was in good company. We had all
placed our muscles on a thermopile in a moist chamber,
moistened preferably with physiological salt solution, to
prevent them from drying. We all knew of the depression
of vapour pressure caused by substances dissolved in water.
We all were aware, at least since the days of Ranke (Ranke
1865), that during activity the osmotic pressure of muscles
rises owing to the production of new chemical molecules in
solution, such as lactic acid: and a rise of osmotic pressure
means a fall of vapour pressure. If we had been asked
we should all have admitted the logical necessity that after
stimulation the vapour pressure of the muscle would be
less, with the consequence that water vapour would pass
over from the walls of the chamber and condense on it,
with an evolution of heat (the " l a t e n t heat of evapora-
tion")· But, we should have answered, the effect, though
theoretically necessary, must be so small as to be com-
pletely negligible. For example at 20° C. the depression
of vapour pressure due to dissolving 0.7 g. of N a C l in
100 g. of water (a solution about isotonic with Ringer's
fluid) is only 0.069 m m * H g ; while the change of vapour
pressure due to the production even of 0.32 p.c. of lactic
11
acid ( c o m p l e t e f a t i g u e ) w o u l d be o n l y a b o u t ο . ο ι α mm.
of H g . H o w c o u l d w e r e a s o n a b l y e x p e c t a fall o f v a p o u r
p r e s s u r e o f 12 μ of H g to c a u s e a t r a n s f e r o f w a t e r v a p o u r ,
a c o n d e n s a t i o n on t h e m u s c l e , s u f f i c i e n t to g i v e a m e a s u r a b l e
heat production. Y e t such indeed w a s the case: w i t h the
instruments now in use (Hill 1930a, Hill and Kupalov
1930, M a r g a r i a 1930) 1 m m . on the g a l v a n o m e t e r scale
c o r r e s p o n d s to a c h a n g e o f w a t e r v a p o u r p r e s s u r e o f t h e
o r d e r of 0.1 μ o f H g , a f r a c t i o n o f a w a v e l e n g t h o f l i g h t .
I h o p e I m a y be a c q u i t t e d o f g r o s s s t u p i d i t y in having
b e e n d e c e i v e d for n e a r l y t w o y e a r s b y an e f f e c t so c o m -
pletely unexpected.
I t is well k n o w n t h a t in o r d e r t o o b t a i n r e l i a b l e r e s u l t s
w i t h i s o l a t e d m u s c l e s it is a d v i s a b l e to s o a k t h e m before
u s e in R i n g e r ' s fluid. T h i s has been the recognised practice
for m a n y years, and L u c a s about 1910 designed a special
m u s c l e t r o u g h for class e x p e r i m e n t s in w h i c h t h e prepara-
t i o n is k e p t b a t h e d . T h e p r i m a r y r e a s o n for this use o f
R i n g e r ' s fluid is t o a v o i d d r y i n g : b u t a p a r t f r o m t h i s t h e
beneficial effect of previous soaking has long been em-
pirically realised. A logical basis for the procedure is
a v a i l a b l e in r e c e n t w o r k . Duliere and H o r t o n (1929) h a v e
p r o v e d t h a t w i t h o u t s u c h s o a k i n g a m u s c l e is a p t to s h o w
a s p o n t a n e o u s loss of i r r i t a b i l i t y , w h i c h c a n be r e c o v e r e d
b y subsequent soaking: H o r t o n (1930) has traced the effect
f a i r l y c o n c l u s i v e l y to t h e e s c a p e o f p o t a s s i u m f r o m c e r t a i n
of the fibres: and Fenn (1930) has demonstrated that
d u r i n g t h e s p o n t a n e o u s l y n o n - i r r i t a b l e s t a t e t h e r e m a y be
an a p p r e c i a b l y g r e a t e r o x y g e n c o n s u m p t i o n . These facts
are mentioned, not because they have any immediate
b e a r i n g on t h e q u e s t i o n o f v a p o u r p r e s s u r e , b u t because
the empirical recognition that a muscle behaves better
a f t e r s o a k i n g h a d led m y c o l l e a g u e s a n d m y s e l f for m a n y
y e a r s to s o a k t h e m u s c l e for an h o u r or t w o o n its t h e r m o p i l e
before commencing myothermic observations. This prac-
tice had, moreover, the a d v a n t a g e t h a t b y b u b b l i n g gas
12
VAPOUR PRESSURE
through the fluid in which the muscle lay the attainment

of thermal equilibrium was greatly quickened—which is an
advantage when you are reading to o.ooooi 0 . Thus, it
had become the regular practice to soak a muscle well on
its thermopile before commencing observation of its heat
production.
Now the preliminary soaking had one effect, indeed one
advantage, which was entirely unforeseen: it brought the
muscle, particularly if thin, into fairly exact osmotic
equilibrium with the fluid around it. When the fluid was
withdrawn and replaced by a gas (oxygen or nitrogen) the
muscle still possessed the same identical vapour pressure as
the fluid remaining on the walls and on the instrument,
with the result that no condensation (or evaporation) of
water on (or from) the muscle occurred. Consequently
the deflection of the galvanometer connected to the thermo-
pile was due solely to the heat produced by the muscle
itself, owing to its proper metabolism, and was unaffected
by the heat of condensation of water-vapour—until the
muscle was stimulated. Then its osmotic pressure rose, its
vapour pressure fell, water passed over and condensed on it,
with an evolution of heat which might be several times as
great as that due to its own resting metabolism. So long,
however, as the muscle continued at rest in oxygen, its
osmotic pressure—and therefore its vapour pressure—re-
mained constant and equal to that of the fluid in which it
had been soaked; and its heat production was correctly
measured. In nitrogen, on the other hand, even without
stimulation, its osmotic pressure gradually rose, owing to
the anaerobic formation of lactic acid and other substances,
and in the course of hours its change of vapour pressure
was sufficient to cause appreciable error due to con-
densation of water on it.
One obvious way of testing the vapour pressure hy-
pothesis of the origin of the increment in anaerobic heat
rate caused by stimulation was to prohibit evaporation
13
altogether by filling the muscle c h a m b e r w i t h paraffin oil.

A good brand of " m e d i c i n a l " oil is p e r f e c t l y harmless to
l i v i n g tissue, at least within the few hours of such an
experiment as this. T h e muscle w a s first soaked for an
hour or two in R i n g e r ' s solution to render it p e r m a n e n t l y
irritable in the sense of Duliere and H o r t o n (1929). The
solution was then replaced by oil t h r o u g h w h i c h nitrogen
w a s bubbled to stir it and to r e m o v e o x y g e n ( o x y g e n is v e r y
soluble in oil). T h e rate of resting heat p r o d u c t i o n was
read once more. T h e r e was no sign of a n y increment.
When evaporation, therefore, was made impossible the phe-
nomenon disappeared.
T h e converse experiment also w a s n e c e s s a r y : if the rate of
evaporation was altered the effect should change proportionally
and in the appropriate sense. A muscle w a s placed on a
thermopile and brought b y prolonged s o a k i n g into osmotic
equilibrium w i t h Ringer's solution. T h e solution was re-
m o v e d , and in o x y g e n or nitrogen the true resting heat
rate was read. T h e c h a m b e r and the i n s t r u m e n t in it
were then v e r y rapidly washed o u t w i t h a new solution,
w e a k e r or stronger than that originally used, and the gas
r e t u r n e d . I f the new solution were w e a k e r its v a p o u r
pressure would be greater, w a t e r w o u l d pass o v e r from the
walls w e t with it and condense on the muscle, the heat of
condensation would be added to the true physiological
h e a t , and an increased deflection w o u l d result. Conversely
if the new solution were the stronger its v a p o u r pressure
w o u l d be less, water would now e v a p o r a t e from the muscle
(with an absorption of heat) and condense on the walls,
the physiological heat would be diminished b y the l a t e n t
h e a t of evaporation, and a smaller deflection w o u l d result.
T h e experiment w a s an easy one to m a k e , and was q u i t e
decisive. A n y value w h a t e v e r , positive or n e g a t i v e , could
be g i v e n to the g a l v a n o m e t e r deflection recording the r a t e
of h e a t production of the muscle, b y an a p p r o p r i a t e a d j u s t -
m e n t of the strength of the solution used for w a s h i n g o u t
14
VAPOUR PRESSURE
the chamber. Incredible as it seemed the riddle had

really been solved at last!
One last test was necessary—the quantitative one.
Muscles had frequently been left for long periods, showing
exaggerated values of their resting heat rates. If these
were due to the condensation of water on them should
there not be sufficient water deposited to be very obvious?
An extreme value for the increment in resting heat rate in
nitrogen, due to complete exhaustion of the muscle, could
be accounted for by the condensation of only 6 mg. of water
per day on a muscle of 100 mg. A more ordinary value
would correspond to 3 or 4 mg. per day, 1 mg. perhaps
during the course of an experiment. An increase of 1 or
1 p.c. in the weight of one of these small muscles would be
quite inappreciable to the eye: indeed it would be difficult
to measure at all with certainty. Clearly there was no
quantitative difficulty in attributing the effect to the
transfer of vapour in the chamber. The high latent heat
of evaporation of water on the one hand, and the almost
incredible sensitivity of the instrument as a wet-bulb
thermometer (for such indeed it was) on the other, had
combined to make it possible to produce all these complex
effects, these fantastic hypotheses, these disturbances in the
daily Press, with the aid of a few milligrammes of water!
How now could the secondary phenomena be explained?
{A) The effect of temperature on the increment in resting
heat rate: the temperature coefficient of the vapour pressure
of water is just the same as that of the observed increment
in the muscle experiments. (Β) Meyerhof's failure to
detect the effect in calorimetrical experiments: due of
course to the fact that evaporation and condensation were
not possible, or—if possible—occurring in the same cham-
ber, the one exactly balanced the other. (C) The effect of
hydrogen: caused simply by the more rapid diffusion of
water molecules through hydrogen than through nitrogen.
(D) The effect of oxygen: due to oxidative recovery and
15
the removal of lactic acid, to restoration of " p h o s p h a g e n , "

to the fall of the osmotic pressure and the rise of the vapour
pressure to their initial values. Perfectly obvious once the
riddle had been solved.
T h e answers to a number of minor questions now
appeared. W h y , for example, if a muscle had not been
given a sufficient preliminary soaking, was it impossible to
obtain a reasonable or constant value of its resting heat
rate? because the muscle was not in osmotic equilibrium
with the fluid in its chamber. F o r a long time my colleague
Hartree, working at Cambridge with the older type of
vulcanite thermopile chamber (Hartree and Hill 1920a),
had noticed that, even after prolonged soaking, a muscle
suspended in oxygen or nitrogen shows a continual tem-
perature drift in the negative direction (Hartree and Hill
1924, p. 452). One less cautious than Hartree might have
been tempted by the perfectly obvious fact to indulge in hy-
potheses as fantastic as those recorded earlier. T h e living
muscle, lying in a moist chamber at constant temperature,
was unquestionably slightly cooler than its surroundings:
were we witnessing here the direct evasion by a living cell
of the Second L a w of T h e r m o d y n a m i c s ? W h a t sport for
the daily Press at the British Association! Hartree re-
corded the result and went on with his work like a reason-
able man: being an engineer he was unwilling to attack the
Second L a w than which nothing is more certain. T h e an-
swer was really much simpler. T h e vulcanite walls of the
chamber absorb a little water (but not salts) from the drop-
lets of solution lying on them; these become slightly more
concentrated: their vapour pressure falls: water vapour
therefore passes from the muscle to the droplets on the wall:
the surface of the muscle is subjected to continual slow
evaporation, with an absorption of heat, increasing as time
goes on. Hartree has abolished the effect completely,
simply by coating the surface of the vulcanite with a film
of paraffin wax. So ends the possibility of a temporary
16
VAPOUR PRESSURE
discovery of great importance: a living cell at constant

temperature taking heat from its surroundings! It might
be better if other supposed discoveries were treated with
similar caution. Here at least, in the Johnson Foundation
for Medical Physics, you will be able to save your physio-
logical and medical colleagues from mistaking physical for
biological effects. Perhaps indeed some of you expect that
they will all prove to be physical in the end!
I was determined, however, that the matter should not
end here. So utterly unexpected a phenomenon, a device
so sensitive in doing a job for which it was not designed or
intended, must be made to expiate its crimes and to repay
the effort wasted on it by being turned to good account.
Firstly it should be calibrated in some way, and used to
tell us the absolute value of the rise of osmotic pressure in
a stimulated muscle, for comparison with the heat liberated
in activity: and secondly it should be applied in a modified
form (a problem in design and manufacture for Mr.
Downing) to the measurement of the vapour pressure of
small quantities of an aqueous solution. The first task it
satisfactorily performed as I will shortly recount. In the
second task its performance was good beyond expectation,
and an instrument and a method have been devised by
which the depression of vapour pressure can be measured,
in a fraction of ι cc. of an aqueous solution, e.g. in human
blood, with a probable error in a single determination of
the order of 0.2 or 0.3 p.c. B y its means molecular
weights may be measured, " f r e e " water distinguished from
" b o u n d , " activities in the physico-chemical sense de-
termined, osmotic pressures and changes of osmotic pressure
recorded, all on minimal quantities of liquid. I do not
know how important all this is, that is for others to judge,
but it is certainly a strange and amusing development of
the original observation. This second phase will be dis-
cussed in the second and third lectures.
The osmotic pressure Ρ of a solution is related to its
17
vapour pressure (for not too great concentrations) by the

equation
logepo/p = PFjRT,
where p0 is the vapour pressure of the pure solvent, p that
of the solution and V the volume 4 occupied in the liquid
state by one gramme molecule of the vapour of the solvent.
For the case of dilute solutions in water this equation may,
with sufficient accuracy for most purposes, be written
[Oo - p)lpo]r = PU-ζβΤ,
where Ρ is reckoned in atmospheres, and [_{po — ρ)!ρο~\τ is
the relative lowering of vapour pressure at absolute tem-
perature T.
We see therefore that the osmotic pressure may be
simply calculated from the relative depression of vapour
pressure (p0 — p)lpt>. The determining factor in a physio-
logical salt solution is the concentration of N a C l , and the
relative depression of vapour pressure of a solution of
N a C l is easily obtained from its molal 5 concentration m,
remembering that over the whole range of physiological
importance the relative molal depression, (p0 — p)/mp0, is
practically constant at 0.0330, and independent of tem-
perature. It is convenient, therefore, in dealing with
osmotic and vapour pressures, to define them simply in
terms of the corresponding solutions of N a C l . The abso-
lute values can be immediately calculated if desired.
As we shall see in the next lecture the osmotic pressure of
frog's blood is equal approximately to that of a solution
containing 0.725 g. of NaCl in 100 g. of water. Soaked in a
Ringer's fluid practically isotonic with this a thin frog's
muscle comes quickly into equilibrium. Placed on a
thermopile in nitrogen and stimulated, an "increment in
heat r a t e " is measured which records its rise of osmotic
pressure: this rise of osmotic pressure is practically a linear
4
To be precise, V is the increase of volume caused by condensing in a large
quantity of the solution ι g. mol. of the vapour of the solvent.
6
g. mols. of solute to iooo g. of HiO.
18
rate of a muscle in nitrogen, due to a series of maximal tetani, is plotted (as ordinate)
against the total heat set free by stimulation (as abscissa). Units arbitrary.
Durations of successive stimuli shown in seconds along each curve. The muscle
in every case finished appreciably (e.g. A, D) or very (e.g. C, H) fatigued, as shown
by the amount of heat in the last contraction (5 sec.) as compared with the first
(1.5 sec.) [Hill and Kupalov 1930.]
function of the heat produced in the activity induced by

stimulation, as shown in fig. 3. In other words, total heat
produced in anaerobic activity is almost, but not quite,
directly proportional to the total number of molecules
simultaneously set free—right up to the stage of rather
advanced exhaustion. This is in a normal muscle: what
it would be in a muscle poisoned with mono-iodo-acetic acid
(see Lundsgaard 1930a), in which lactic acid formation does
not occur on stimulation, remains to be f o u n d : 6 the experi-
ments of fig. 3 were made before L u n d s g a a r d ' s paper was
published. I t is important, in any case, to know that in
the normal muscle at least the energy is very nearly pro-
portional to the number of molecules set free.
T h e major chemical changes known to occur in normal
stimulated muscle are lactic acid formation and phosphagen
breakdown. According to Meyerhof (1930a, p. 233) the
isometric coefficient of lactic acid (the ratio of the product
of tension developed and length to lactic acid formed in
an isometric twitch) is almost entirely unaffected by tem-
perature, but depends to some degree on the initial tension
and appreciably on the degree of fatigue of the muscle.
B y plotting and differencing M e y e r h o f ' s data (1930a, p.
234) it can be calculated that in successive productions of
0.05 p.c. lactic acid (i.e. from o. to 0.05, 0.05 to 0 . 1 0 , o. 10 to
0 . 1 5 etc.) the isometric coefficient of lactic acid has the
following values, all multiplied by io 6 : 162, 1 3 4 , 1 1 4 , 1 0 0 ,
96, 88, 82. According to my own observations the iso-
metric coefficient for heat ( T l j H ) also is independent of
temperature (Hartree and Hill 1 9 2 1 b ) and is somewhat
affected by initial tension (Hill 1925) but is practically
constant throughout a long series of twitches (Hill 1928c),
up to the beginning of fatigue: in a very fatigued muscle,
however, it diminishes, probably owing to another cause, the
failure of the contractile mechanism. Dividing the one
• T h e same effect has been found in muscles poisoned with iodo-acetic acid. See
footnote 1 2 , p. 25 below and Hill & Parkinson ( 1 9 3 1 ) .
20
VAPOUR PRESSURE
c o e f f i c i e n t b y t h e o t h e r w e o b t a i n t h e r a t i o o f h e a t to l a c t i c
a c i d f o r m e d ; w e see t h a t t h i s d e c r e a s e s c o n s i d e r a b l y b e t w e e n
t h e b e g i n n i n g a n d t h e e n d o f a l o n g series o f t w i t c h e s .
It is very unlikely, indeed impossible, that the same
chemical reaction should yield different quantities of heat
at different s t a g e s of a c t i v i t y : the e x p l a n a t i o n of the fall of
t h e i s o m e t r i c c o e f f i c i e n t for l a c t i c a c i d is a l m o s t certainly
in p a r t t h a t in t h e e a r l i e r s t a g e s o f a c t i v i t y a c o n s i d e r a b l e
f r a c t i o n o f t h e e n e r g y is d e r i v e d f r o m p h o s p h a g e n break-
d o w n , so t h a t t h e l a c t i c a c i d is less a n d t h e c a l o r i c q u o t i e n t
(heat/lactic acid) greater. A b o u t 1/5 o f t h e e n e r g y s e t f r e e
i n s t i m u l a t i n g a m u s c l e t o f a t i g u e (see L e c t u r e V ) m a y be
derived from phosphagen breakdown.7 It was shown by
Eggleton and Eggleton (1927b, 1928) that the ratio of
phosphagen breakdown to lactic acid formation decreases
r a p i d l y as a n a e r o b i c s t i m u l a t i o n is c o n t i n u e d , a n d t h i s h a s
been amply confirmed (see Meyerhof 1930a, p. 101).
D i r e c t d e t e r m i n a t i o n s of the caloric q u o t i e n t for lactic acid
by Meyerhof and his colleagues (see M e y e r h o f 1930a, p.
203) have shown the same general effect. For 0.1 p.c.
lactic acid the quotient was 376 cal. per gramme, for
0.2 p.c., 355 cal., i.e. for the second 0.1 p.c. lactic acid
3 3 4 cal.: w i t h c o n s i d e r a b l e f a t i g u e the v a l u e w a s still lower.
I t is c l e a r t h a t in t h e e a r l i e r s t a g e s o f a n a e r o b i c activity
e n e r g y is l i b e r a t e d t o a n a p p r e c i a b l e e x t e n t a t t h e e x p e n s e
of phosphagen breakdown, in the later stages almost
entirely b y the formation of lactic acid. The myothermic
o b s e r v a t i o n s s h o w , h o w e v e r , t h a t in b o t h s t a g e s , i.e. f r o m
e i t h e r s o u r c e o f e n e r g y , t e n s i o n is d e v e l o p e d (a) w i t h a b o u t
t h e s a m e t o t a l l i b e r a t i o n o f h e a t 8 a n d (b) w i t h a b o u t the
s a m e t o t a l l i b e r a t i o n o f n e w m o l e c u l e s or i o n s .
'Probably more like one-third from sources other than lactic acid. See Hill
and Parkinson (1931).
8 This conclusion is strikingly confirmed by the fact that in a muscle poisoned
with iodo-acetic acid, in which no lactic acid is formed, the ratio Tl/H in a twitch is
exactly the same as in a normal muscle. Fischer (1930a, 1931): Meyerhof, Lunds-
gaard & Blaschko (1930): confirmed by Feng and myself.
21
3
With regard to the latter, so far as the lines of fig. 3 are

not straight they suggest that energy, in the earlier stages
of activity, is liberated with a rather smaller rise of osmotic
pressure than in the later stages. T h i s cannot be explained,
nor indeed can a linear relation, by assuming that in the
earlier stages of activity the energy is derived more from
phosphagen breakdown, in the later stages more from lactic
acid formation. T h e energy liberated, per g. mol., in the
splitting of phosphagen is about 1 1 0 0 0 cal. (Meyerhof
1 9 3 0 a , p. 94). If the breakdown be
" p h o s p h a g e n " molecule —> creatine + phosphate,

then one new gramme molecule is liberated per 11000
calories. If it be
X — "phosphagen" X + creatine phosphate,
where X is some other unknown substance, then two new

gramme moleculcs are liberated per 1 1 0 0 0 calories. In the
formation of lactic acid alone in muscle we have (say)
300 cal. per g. or 27000 cal. per g. mol. For a given amount
of heat therefore the number of new molecules set free is
much less in the case of glycogen than in that of phosphagen
breakdown: this is the wrong w a y round 9 to explain the
curvatures of fig. 3. T h e phosphagen breakdown occurring
particularly at the beginning, i.e. to the left of each diagram,
should give there a greater and not a smaller rate of rise
in the osmotic pressure curve. Clearly there are other
factors at work, hitherto unrecognised: indeed the obser-
vations which I will next discuss furnish a strong hint
that during anaerobic activity, during fatigue, there is an
appreciable excess of new molecules or ions produced, over
• This conclusion is borne out to some degree by the observation (Hill and
Parkinson 1 9 3 1 ) that the liberation of 1 cal. of heat per 1 g. by a muscle poisoned
with iodo-acetic acid (in which no lactic acid is formed) is accompanied by a rise
of osmotic pressure equivalent to that produced by adding 0.40 p.c. N a C l to Ringer's
fluid: whereas 1 cal. per 1 g. in a normal muscle gives a rise of osmotic pressure
equivalent only to 0.33 p.c. N a C l .
22
VAPOUR PRESSURE
and above all those known at present to chemical analysis.

We are not yet at the end of the biochemistry of muscle.
The rise of osmotic pressure shown by the increment in
heat rate can be measured in absolute units, and compared
with the heat set free during preceding activity. By
washing out the chamber with Ringer's fluids of different
concentrations the apparent rate of heat production can be
altered: and it is clear that the speed of condensation of
water vapour on the muscle, owing to a given rise in its
osmotic pressure, must be the same as that due to an equal
fall in the osmotic pressure of the fluid in the moist chamber.
For example, if a deflection of 200 mm. be given when a
Ringer's fluid isotonic with 0.75 p.c. of NaCl is replaced
on the walls of the chamber by one isotonic with 0.45 p.c.
NaCl, then it is obvious that a deflection of 200 mm.
obtained by stimulation is to be attributed to a rise of
osmotic pressure equal to that caused by adding 0.3 p.c. of
N a C l to the fluids of the muscle. Kupalov and I (1930a)
have made a number of such observations and found that,
on the average, for 1 calorie per gramme of heat set free in
fatiguing a muscle, there is a rise of osmotic pressure
equivalent to the addition of 0.335 8· NaCl to 100 g.
of its fluids.10
In complete fatigue there may be 0.3 to 0.35 p.c. of lactic
acid set free, 1.05 to 1.225 calories of heat liberated per
gramme. Taking 0.3 p.c. of lactic acid as a standard, this
degree of fatigue will lead, according to our results, to a
rise of osmotic pressure equivalent to the addition of
0.35 p.c. of NaCl to the fluids of the muscle. It is inter-
esting to compare this value with that calculated from all
the substances known to be produced in fatigue. We will
assume that the muscle is 77 p.c. " f r e e " water and that the
lactate ion exerts a normal osmotic pressure; it is easy to cal-
culate that the increase of osmotic pressure on stimulation to
10
Confirmed by Hill and Parkinson (1931) who found a mean value of 0.326 in
similar experiments.
23
fatigue is 2.8 times as great as the osmotic pressure of the

lactate ions liberated.
T o explain the discrepancy it might be argued that the
lactate ion is necessarily accompanied by an alkali ion,
e.g. K , and that before combination with lactate the Κ ion
was not free in solution to affect the osmotic pressure.
There is, however, grave difficulty in assuming that the
Κ ions associated after stimulation with the lactate ions
were not free in solution before. T h e existence of elec-
trically undissociated compounds of Κ in muscle is unlikely,
and even if the anions were unable to diffuse (e.g. if they
were part of the protein structure of the muscle) the cations
would be free to change partners and so to move about
and presumably to affect the osmotic pressure. Moreover,
as I shall show in my next lecture, the osmotic pressure o f
resting muscle can be calculated fairly exactly from the
known soluble constituents dissolved in the water of the
muscle. I f we were to argue that the cations before
stimulation were not free to affect the osmotic pressure we
should be left with a serious deficit in our calculation.
Finally, direct experiments in which an acid ( C 0 2 ) was
added artificially to resting muscle caused an increase of
osmotic pressure rather less than that of its anions alone,
certainly not twice as great: an effect which has been
confirmed by Margaria (1931b) by experiments in which the
rise of osmotic pressure due to the combination of C 0 2 w i t h
blood was found to be appreciably less than that calcu-
lated for the bicarbonate ions alone, certainly again not
twice as great. I t is clear therefore that the cations play
no part in the phenomenon described.
Lactic acid, however, is not the only substance liberated
in fatigue. T h e breakdown of phosphagen into creatine
and phosphate, if we suppose it to occur according to the
second scheme referred to earlier, 11 liberates two new
11
There is evidence from recent work of Eggleton (1930) that " p h o s p h a g e n " in
living muscle is not a simple uncombined molecule: phospho-creatine can diffuse
24
VAPOUR P R E S S U R E
molecules for every atom of Ρ set free. According to

Parnas and Mozolowski (1927) the ammonia content of
muscle increases in severe fatigue. According to Lohmann
(1928), in exhausted muscle a certain amount, say, one
third, of the pyrophosphate initially present in it is hydro-
lysed into orthophosphate; if this pyrophosphate had
previously been combined with adenylic acid (Lohmann
1929) each molecule split would become three. Taking
account, therefore, of every known possibility, and making
a maximal allowance in each case, the increase in total
molal concentration as the result of fatigue can be calcu-
lated as follows:
L a c t a t e (0.3 p . c . ) 0.043 Μ
Creatine 0.023 Μ
Phosphate 0.023 Μ
Ammonia 0.002 Μ
Pyrophosphate-adenylic acid 0.004 Μ
Total °.°95 Μ
Now 0.095 Μ ' s sum m

° l a l concentrations of the
ions of a solution of 0.28 g. of N a C l in 100 g. of water.
This is only 80 p.c. of 0.35, the observed change. We have
taken the most favourable view possible of all the reactions
known, or supposed, to be involved, in order to provide as
many new molecules as possible. In spite of it there is
still a deficit. 12 The existence of this deficit has more
through a parchment membrane: phosphate and creatine can diffuse through the
living muscle: phosphagen can not. Presumably the phosphagen breaks down
from combination in a larger complex,
X — phosphagen —1• X + phosphate + creatine.

u In muscles poisoned with iodo-acetic acid, in which lactic acid formation is
impossible, the chemical changes known to occur are (Lundsgaard 1930b):
(a) phosphagen becomes phosphate and creatine;
( i ) adenyl-pyrophosphoric acid becomes orthophosphate, inosinic acid and
ammonia;
(r) the phosphate formed is esterified with hexose derived from glycogen, about
70 p.c. as the di- and 30 p.c. as the mono-ester;
(d) some glycogen is broken down to hexose. These changes, at the best, ac-
25
recently been confirmed by Meyerhof (1930b) in experi-

ments in which the extra depression of freezing point,
caused by stimulating muscles to fatigue, was compared
with the various substances produced. Presumably, there-
fore, some other changes, at present unrecognized, are in-
volved. It is indeed unlikely that, after the striking prog-
ress of the last few years, we should at this particular
moment have discovered all the reactions involved in
muscular activity. M a y not carnosine, for example, be
found to supply part of the missing osmotic pressure?
There is no evidence indeed as yet that it will, but for
many years creatine had also apparently no particular role
to play. Carnosine is present in considerable quantities,
specifically in skeletal muscle. There is said to be 0.25 p.c.
in frog's muscle, considerably more in the muscles of some
mammals. Little attention has been paid to it, and
methods of determining it are at present very uncertain.
Its molecular weight is 226, so that if 0.25 p.c. of carnosine
were set free in muscular activity it would form a 0.012 Μ
solution in the water of the muscle—which is about half
the quantity we require to complete the story.
L e t me show you, in closing, how consistent nature is.
We have seen a very considerable rise of osmotic pressure
in a frog's muscle stimulated anaerobically. Cannot the
same phenomenon be demonstrated in man? This, by the
w a y , is always a good question to ask: it often leads you
into work more interesting even than the original discovery.
It is easy to make a man take exercise of the required na-
ture: any very severe effort, lasting even for a minute,
leaves behind it a considerable " o x y g e n debt." We should
expect the muscles of a man exhausted by violent exercise
count for 50 to 70 p.c. of the increase of osmotic pressure observed (Hill & Parkinson
1 9 3 1 ) , unless we assume that the phosphagen and perhaps the adenyl-pyrophosphoric
acid were combined in some more complex form in the resting muscle, and so were
unable to exert an osmotic pressure of their own. Otherwise it is necessary, as in
the normal muscle, to suppose that some reaction, hitherto unknown, occurs during
activity.
26
VAPOUR P R E S S U R E
to have a very high osmotic pressure. This should rapidly

be communicated to the blood, partly by diffusion outwards
of the products of metabolism, but more particularly by
the diffusion of water in. The experiments of Margaria
(1930) to be referred to again in a later lecture have shown
a surprisingly constant value in the osmotic pressure of the
blood of men at rest: the mean value is equal to that of a
solution containing 0.945 g. of NaCl in 100 g. of water: the
probable deviation of an individual value from the mean is
only ± 0.005. The of a runner who had competed
in the Amateur Athletic Championships in London, with-
drawn 75 sec. after the end of the two miles steeplechase,
had a value of 1.048, more than 1 1 p.c. higher than the
mean. Margaria himself, after 1 minute of standing run-
ning, showed a value of 0.989: he had to go slow, however,
in order to be in a fit state at the end to withdraw his own
blood: our colleague, Hukuda, who had no such obligation,
after minutes of standing running, gave the following
values:
J
Blood withdrawn f min. afterwards -°33
Blood withdrawn i j min. afterwards 0.996
Blood withdrawn 4} min. afterwards 0.982
M y own blood, withdrawn 60 seconds after 1 minute only

of standing running, gave a value of 1.028.
Standing running at maximum speed is extremely ex-
hausting exercise; correspondingly it raises the osmotic
pressure of the blood, in a very short time, to astonishing
values: after which the osmotic pressure slowly returns to
normal as recovery proceeds. The increment recorded in
these experiments is quite outside the range of normal
resting values: it exceeds the probable deviation twentyfold
or more. I have little doubt that the blood of a first class
runner, taken shortly after a quarter or a half mile race,
would show considerably greater increments than these:
in fact I am inclined to put a value of 1 . 1 5 as well within
the range of possible accomplishment, for exercise of ex-
27
treme severity lasting for a minute or two in a first rate

subject. It may be interesting to some of you, in this land
of record-breaking, to try. One can never hope to obtain
a value as high as that of the muscles themselves: diffusion
is not rapid enough between muscle and blood, and recovery
sets in too fast. But at least one m a y hope, in the blood
of the human athlete, to show an increment of osmotic
pressure which is 60 p.c. of that found in the isolated muscle
driven to exhaustion. M a y not such a change of osmotic
pressure, leading temporarily to a large upset in the water
distribution of the organism, be the cause of some of the
phenomena of fatigue? Might not, for example, an osmotic
suction of one atmosphere, suddenly applied, considerably
upset the central nervous system ?
28
L E C T U R E II
THE STATE OF W A T E R IN TISSUES
O R the calculations given in the last lecture we assumed

F that the substances formed during muscular activity
are simply and normally dissolved in the water of the tissue,
which was taken as being 77 p.c. by weight. Actually
there is 80 to 81 p.c. of total water in frog's muscle: the
value 77 p.c. was chosen for reasons which will be referred
to later. I f , however, the chemical bodies had been dis-
solved in water equal in weight to 61 j p.c. of the muscle
and not to 77 p.c. they would have given the full value of
the observed increase in osmotic pressure: there would have
been no cause to argue that some chemical body still un-
known is liberated in muscular activity. Alternatively,
we might attribute the observed rise of osmotic pressure
not entirely to an increase in the quantity of the dissolved
substances themselves but partly to a decrease—caused
by activity—in the volume of the solvent. F o r example,
if the free water of the muscle, initially 77 p.c., were reduced
b y some colloidal process occurring in fatigue to 72 p.c.
the dissolved constituents initially isotonic with 0.725 p.c.
N a C l would be confined to a smaller volume, and their
osmotic pressure would rise to a value equivalent to that
of 0.725 X 77/72 = 0.776 p.c. N a C l . T h e chemical sub-
stances formed in a c t i v i t y , moreover, instead of being
dissolved in 0.77 g. of w a t e r per g. of muscle, would be
dissolved in 0.72 g. of water, and their osmotic pressure
would be that, not of a 0.28 p.c. N a C l solution, but of a
0.28 X 77/72 = 0.297 p.c. solution. T h u s the final os-
motic pressure would be that of a solution of (0.776 + 0.297)
= 1.073 P- c · N a C l . T h i s is about 0.35 p.c. in excess of the
initial value 0.725 p.c., so that the whole of the deficit in
the calculated osmotic pressure below the observed could
29
be explained by the simple hypothesis that the " f r e e "

water of the muscle diminishes from 0.77 g. to 0.72 g. per
g. of muscle as the result of fatigue.
Clearly this possibility could not be neglected: it required
careful examination. Indeed if it could be shown to be a
fact it might provide a factor of extraordinary potency in
altering or adjusting the osmotic pressure of living tissues,
or their fluids. It is easy to imagine colloidal changes to
occur by which some of the water is removed from its
normal state as a solvent, and combined in some other state
with the colloid: the consequence would be that the sub-
stances dissolved would be confined to a smaller volume and
their osmotic pressure would rise, without the liberation
of any molecules at all.
T h e possibility had indeed been claimed as a fact as long
ago as 1910. Jensen and Fischer ( 1 9 1 0 ) and Jensen (1912)
determined the " bound " water of muscle by making cooling
curves and calculating from their areas the heat absorbed.
A comparison of the results with those obtained with solu-
tions of N a C l was believed to allow a calculation of the
" b o u n d " water. In fresh muscle they found 4 p.c. of the
total water to be " b o u n d , " in muscle killed by freezing
and thawing 14 to 17 p.c., in muscle heated to ioo° C.
22 p.c. Changes much smaller than these, occurring in
fatigue, would be sufficient to explain completely the ob-
served excess of osmotic pressure.
Since the publication of Overton's experiments (1902)
it has been commonly supposed that a large proportion of
the water of muscle exists in some " b o u n d " form, in-
capable of taking part in the osmotic changes which occur
when the tissue is immersed in hypo- or hyper-tonic solu-
tions. From the fact that a muscle swells to much less
than double its initial weight when immersed in a solution
of half the initial osmotic pressure Overton concluded that
" o n l y part of the water found in muscle can be present as
a solvent." Overton's experiments can easily be confirmed.
30
S T A T E OF WATER IN TISSUES
Their explanation, however, as I have shown in a recent

paper (Hill 1930b), is not a " b i n d i n g " of the water, but
(a) progressive changes, owing to survival and immersion,
{b) the slowness of diffusion, and (c) the loss of semi-
permeability in a considerable fraction of the fibres. Over-
ton's experiments, in fact, provide no support for the theory
of " b o u n d " water in muscle. They must be referred to,
however, for they seem to have started the tradition.
A very obvious objection was raised by Rubner (1922) to
the experiments of Jensen and Fischer, viz., that thermal
conduction may be quite different in muscle and in solutions
of N a C l . Rubner proposed an alternative method based
on the same general idea, viz. that " b o u n d " water may be
defined as that which cannot be frozen out by cooling the
tissue to such temperatures as — 20° C. The material to
be investigated was cooled to a low temperature for two
hours and then dropped into a water calorimeter, the heat
required to melt it being measured. He found that 1
gramme of dry substance was associated with the following
amounts of " b o u n d " water:
Egg-white 0.33 g.
Blood corpuscles 0.63
Elastic tissue 0.44
Blood vessels 0.45
Beef muscle (dead) 0.76
Beef heart-muscle (dead) 0.64
Frogs' muscle (alive) 0.90
The frog's muscle was cooled to — i8° C. and 100 g. of

muscle contained 61.7 g. of " f r e e " water (mean of 11
observations); similar muscle contained 79.8 p.c. of total
water. 1
Rubner's method also is not free from possible objection:
(i) it assumes that the " b o u n d " water is the same at
1 In dilatometric experiments carried out at —2o° C . Moran and Smith (i930)
found " t h a t the amount of water remaining unfrozen cannot possibly be greater
than 6 p.c. of the total water in the muscle." I t is "quite possible," they point out,
" t h a t muscle contains no bound water," even at — 20° C .
31
— 20° C . as at the ordinary t e m p e r a t u r e s in which we are

interested; the " b i n d i n g " of w a t e r b y a hydrophilic colloid
is likely to be an exothermic reaction, in which case it
m i g h t proceed appreciably further at a low temperature
t h a n at a high.
(ii) it assumes t h a t no reactions other than the melting
of ice occur when the temperature rises.
(iii) the specific heats of the solid constituents of muscle
m a y not be v e r y accurately k n o w n , nor the heats of their
solution negligible. It is possible, moreover, that water
p r e v e n t e d from freezing by association with hydrophilic
colloids m a y nevertheless be c a p a b l e of dissolving sub-
stances present in the tissue, and so be in t h a t sense free.
R u b n e r ' s method was e m p l o y e d b y T h o e n e s (1925); in-
deed it is continually a t t r i b u t e d b y A m e r i c a n writers to
T h o e n e s . T h e latter found in a gelatin jelly about 2 g. of
w a t e r " b o u n d " by each 1 g. of d r y m a t e r i a l ; in agar j e l l y
a b o u t 4 g. In the muscles of y o u n g animals he found about
2 g. of " b o u n d " water per 1 g. of d r y substance, in those of
old animals about 1 g. T h e muscles were frozen and the
h e a t of t h a w i n g measured. " I n this w a y , " he claimed,
" i t can be shown w i t h considerable c e r t a i n t y t h a t there is
a change during rigor in the a m o u n t of water b o u n d . "
Robinson (1927, 1928) applied the same method to inves-
t i g a t i n g the hardiness of insects exposed to low temperatures
during winter. In some insects as m u c h as one half of the
w a t e r they contain m a y be " b o u n d , " in the sense that it is
not frozen by cooling to — 20° C . In h a r d y insects (Pro-
mothea) exposed to low t e m p e r a t u r e s the proportion of
w a t e r " b o u n d " m a y increase from a b o u t 8 p.c. at the s t a r t
to o v e r 40 p.c. after t w o or three weeks' exposure. The
m e t h o d e m p l o y e d by T h o e n e s and b y Robinson is discussed
in detail by G o r t n e r (1929). It is open to the same possible
objections as t h a t of R u b n e r .
A n o t h e r definition of, and another m e t h o d of determin-
ing, the " f r e e " w a t e r were suggested by N e w t o n and
32
S T A T E OF W A T E R IN TISSUES
Gortner (1922), who added a known amount of cane sugar

to expressed plant juice and measured the resulting depres-
sion of freezing point. 2 T h e greater the amount of water
" b o u n d " the less will be the volume of water free to dis-
solve the added cane sugar, and the greater will be the
depression of freezing point. B y comparing the depression
of freezing point observed with that caused by adding the
same amount of cane sugar to an amount of water equal
to the total quantity contained in the juice, they showed
that an appreciable fraction of the water was " b o u n d , " in
the sense that it took no part in the solution of the cane
sugar. In one case in which the solids made up 0.178 g.
per ι g. of juice, of the total water (0.822 g.) 0.130 g.
was found to be " bound." Newton (referred to by Gortner
(1929)) employed the same method of studying the state
of water in the sap of winter wheat, in drought-resistant
crops, and in the press juice of grasses.
In a recent paper Gortner (1930) has emphasised his
sense of the importance of the conception of " b o u n d "
water in biology. T h e questions raised as to the status of
water in various animals and plants are clearly of great
interest and merit further research and thought. I
have had no personal experience of the matter except in
so far as it has been presented by the problem which I
discussed in my first lecture: the observed changes in the
osmotic pressure of muscle obviously demanded a knowl-
edge of the amount (if any) of " b o u n d " water in muscle
and of its changes (if such occurred) during and as the result
of activity. In investigating the problem I have been led
1 This method has recently been criticised by Grollman (1931) and by Moran and
Smith (1930), particularly on the ground of the anomalies occurring in salt solutions
to which high concentrations of sucrose are added. Indeed, according to Grollman,
the method employing high concentrations of sucrose may often be shown to lead
to impossible results. He suggests that the addition of small quantities of N a C l or
K C l and the use of a delicate means of measuring the depression of vapour pressure
(see below) provide the most appropriate method of approaching this problem.
With such a method he found that the amount of water bound in gelatin solutions
at p H 7.0 is relatively small, and in gum acacia solutions negligible.
33
incidentally to study blood, the body fluids of inverte-

brates, casein solutions and eggs. The opinion, therefore,
which I shall express, that in muscle, and in such fluids, the
" b o u n d " water is of little, if any, importance, and that the
" f r e e " water is nearly equal to the total water as measured
by drying, must be understood at the moment to apply
solely to such fluids. The methods which we will discuss
could be applied to a variety of other problems, but the
results of such applications must not be forestalled. 3
As a preliminary to the description of the experimental
study of the problem, it is interesting to see how far the ob-
served osmotic pressure of blood and muscle can be calcu-
lated from the known concentrations of their soluble
constituents as determined by analysis, supposing these to
be dissolved in the total water as measured by drying. Inci-
dentally mammalian blood is an excellent physico-chemical
" m o d e l " of muscle; it contains about the same relative
amount of water, and its chief protein, haemoglobin, is an
efficient buffer, being (like the proteins of muscle) the
ionised alkali salt of a weak acid.
Mammalian blood. The mean osmotic pressure of human
blood, taken from men at rest, is very accurately known
from the investigations of Margaria (1930), which will be
referred to in detail in the next lecture. It is equal to that
of a N a C l solution containing 0.945 g. in 100 g. of water.
Abderhalden's (1898) analyses of the blood of two cattle,
two sheep, two horses, two dogs, one goat, one pig, one
rabbit and one cat, allow the following mean values to be
calculated. It is realised that by taking the mean of his
12 values for each constituent we obtain a result which is
true in detail for no particular animal; since, however, our
only object is to find the sum of the molal concentrations
for mammalian blood, no error is introduced by taking the
mean, and the result is more accurate.
• S e e , however, Grollman ( 1 9 3 1 ) .
34
STATE OF WATER IN TISSUES
TABLE I.—Mean Values Calculated from Abderhaldens Data for Mammalian Blood
g. to g. to Molal
Substance 100 g. 100 g. concen- Remarks
blood HjO tration
HiO 80.2 — —
Hb 12.5 15.6 Ο.ΟΟ23 Molecular weight assumed 67000

(Adair (1925), (1928))
Sugar 0.0790 0.0980 0.0054 —
Na 0.2460 0.3060 O.I33J —
Κ 0.0790 0.0980 0.0252 —
Ca 0.0044 0.0055 O.OO IO Assumed 70 p.c. as free Ca ions

Mg 0.0031 0.0039 O.OOL6 —
CI 0.2890 0.3604 O.IOI5 —
Total Ρ 0.0314 0.0392 0.0088 See note
Sum. . Ο.2789
Note.—The molal concentration for total Ρ assumes that the number of dissolved
molecules containing phosphorus is 70 p.c. of the number of phosphorus atoms.
See Hill 1930b, p. 482.
A b d e r h a l d e n ' s list, h o w e v e r , a l t h o u g h it m a k e s u p nearly

87 p.c. of the sum of the m o l a l c o n c e n t r a t i o n s , m u s t be
s u p p l e m e n t e d b y the following d a t a , o b t a i n e d from v a r i o u s
sources.
TABLE I I . — C o n s t i t u e n t s of Mammalian Blood
Mola]
Substance Concentration assumed concen- Remarks
tration
Protein other than One third of the value 0.0008 A rough estimate, but the
Hb for Hb value is practically negli-
gible in any case
Bicarbonate 50 c.c. combined COi per 0.0280
100 g. blood
Lactate 16 mg. per 100 g. blood. 0.0022
SO. 0.0003
Urea 0.030 p.c. in blood 0.0062
Amino acids 0.006 p.c. Ν in blood. . . 0.0053 One Ν atom assumed to
each molecule
Creatine 0.007 P-c· ' n blood 0.0006
Creatinine 0.001 p.c. in blood O.OOOI
Uric acid 0.002 p.c. in blood
0.0002
Sum
0.0437
35
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décida que la durée du carême était de quarante et non de
cinquante jours[241]. Il rendit obligatoire la fête des Rogations,
récemment instituée à Vienne en Dauphiné par saint Mamert, et qui
de là s'était répandue rapidement dans le reste de l'Église. Il voulut
que les trois jours qu'elle durait fussent des jours de jeûne et
d'abstinence; il décida que les esclaves des deux sexes seraient
dispensés de tout travail afin de pouvoir assister aux processions, et
il donna pouvoir à l'évêque de punir le prêtre qui refuserait d'y
participer[242]. Ainsi continuait au sein de l'Église la floraison
liturgique; chaque génération en s'écoulant ajoutait un joyau au
diadème de ses fêtes, et le cercle enchanté de ses prières se nouait
en guirlandes parfumées autour de toute l'année chrétienne.
[240] Canon, 26, Sirmond, I, p. 182; Maassen, p. 8.
La discipline ecclésiastique était peut-être, de tous les sujets, celui
que l'Église soignait avec le plus de sollicitude; aussi ne s'étonnera-
t-on pas d'y voir consacrer un grand nombre de canons. Il faut parler
d'abord des attributions réservées aux évêques en leur qualité de
chefs de diocèse. Le diocèse était dans l'Église primitive, et avant le
mouvement de concentration qui s'est fait autour de la chaire
romaine, l'organisme par excellence de la vie religieuse, et l'évêque
était le centre et la source de toute autorité et de toute discipline. Le
lien qui rattachait les fidèles à leur évêque était le lien le plus fort qui
les rattachât à l'Église elle-même: il fallait veiller, s'il y avait lieu de
l'élargir, à ce qu'il ne pût jamais être défait ou rompu. Voilà pourquoi
l'on faisait aux fidèles dispersés dans les paroisses rurales
l'obligation d'affirmer par intervalles l'unité diocésaine, en venant
assister aux offices de la cathédrale aux fêtes de Noël, de Pâques et
de Pentecôte. Le concile d'Orléans renouvela cette prescription[243].
Il rappela aussi aux fidèles que toutes les églises qui se
construisaient dans le diocèse, que ce fût dans le domaine d'un
particulier ou ailleurs, restaient sous la juridiction de l'évêque[244]:
mesure d'une importance capitale, qui sauvegardait l'unité
religieuse, et constituait la barrière la plus solide que la féodalité
envahissante ait rencontrée sur son chemin. Le concile consacra
l'autorité de l'évêque sur toutes les personnes comme sur tous les
biens de son église; il lui en subordonna les religieux comme les
laïques; il ne permit ni à ses prêtres ni à ses moines d'aller trouver le
roi pour lui demander un bénéfice sans la permission de l'évêque
diocésain; celui qui contreviendrait à cette défense devait être privé
de son rang et de la communion jusqu'à ce qu'il eût satisfait[245].
Mais en même temps qu'il veillait à conserver intacte l'autorité
épiscopale, le concile voulut que l'évêque se souvînt aussi de ses
devoirs: il exigea que tous les dimanches, sauf empêchement, il
assistât aux offices de l'église la plus voisine[246]; il ne lui permit pas
de manier l'arme de l'excommunication contre un laïque qui
revendiquerait les biens d'une église ou d'un évêque[247]. Il est
intéressant de constater ces restrictions que les évêques eux-
mêmes apportent à leur pouvoir: rien ne montre mieux l'action
modératrice des conciles.
[243] Canon 25, Sirmond, I, p. 182; Maassen, p. 8.
[244] Canon 17, Sirmond, p. 181; Maassen, p. 6.
Plusieurs autres dispositions des conciles antérieurs furent
renouvelées en ce qui concernait la vie du clergé. Telle fut en
premier lieu celle qui défendait aux clercs de tout rang, tant aux
évêques qu'aux prêtres et aux diacres, d'avoir dans leur maison
d'autres femmes que leurs parentes les plus proches[248]. Il fut
interdit aux veuves de clercs de se remarier; celles qui avaient
contracté mariage furent contraintes de rompre leur union, sous
peine d'excommunication tant pour elles que pour leurs
complices[249]. Enfin il fut décidé que le prêtre ou diacre coupable
d'un crime capital serait privé de son office et exclu de la communion
des fidèles[250].
Tout cet ensemble de mesures était relatif au clergé séculier; il faut y
ajouter celles qui concernaient le clergé régulier. Quatre importants
canons furent consacrés à la vie monastique, et il faut remarquer
qu'ils ont pour caractère général le renforcement de l'autorité
épiscopale sur le clergé régulier. Les abbés des monastères, se
souvenant, dit le concile, de l'humilité dont leur profession leur faisait
un devoir, eurent à reconnaître l'autorité de l'évêque, et celui-ci
garda sur eux un droit de correction. Tous les ans ils devaient se
réunir à l'endroit où il leur avait donné rendez-vous. Eux-mêmes, de
leur côté, voyaient confirmer leur autorité sur leurs moines. Le
religieux qui, contrevenant à sa règle, possédait quelque chose en
propre, devait en être dépouillé par l'abbé; celui qui s'évadait de son
monastère devait y être ramené et mis sous bonne garde, avec
l'aide de l'évêque. L'abbé lui-même était déclaré coupable s'il n'usait
pas de son droit de correction, ou s'il accueillait un moine fugitif[251].
Il fut défendu aux moines de quitter leur monastère pour se bâtir des
cellules à part, à moins qu'ils n'eussent l'aveu de leur évêque et de
leur abbé; les Pères du concile voyaient dans cette tendance à
s'isoler une preuve de vanité et d'outrecuidance[252]. Ils fermèrent
l'accès de tout grade dans l'ordre ecclésiastique à quiconque, après
avoir professé la vie religieuse en prenant le manteau de moine,
l'avait ensuite quittée pour contracter les liens du mariage[253]. Enfin,
descendant jusque dans le détail, ils réglèrent de menues questions
de costume monastique[254].
Les simples fidèles s'entendirent rappeler une des défenses les plus
impérieuses de cette époque: celle du mariage entre beaux-frères et
belles-sœurs, et il faut remarquer que par belle-sœur on devait
entendre, au sens du concile, aussi bien la femme du frère que la
sœur de la femme[255]. Deux canons, le onzième et le douzième,
furent consacrés aux pénitents, classe de fidèles toujours
nombreuse, et qui comprenait plusieurs catégories. Il y avait ceux
que l'Église avait condamnés à la pénitence pour expier leurs fautes;
il y avait aussi ceux qui se l'étaient imposée spontanément et par
ferveur de contrition. Ceux-ci étaient tenus de respecter leur vœu et
ne pouvaient retourner à la vie du siècle, sinon ils étaient exclus de
la communion, et nul fidèle ne pouvait les admettre à sa table sans
s'exposer à partager leur sort. Toutefois, si un prêtre ou un diacre
avaient, par pénitence, abandonné le service de l'autel, il leur fut
permis, par égard pour le salut des âmes, d'administrer le sacrement
de baptême en cas de nécessité[256].
Dans les mesures qu'il prit par rapport aux biens ecclésiastiques, le
concile, comme dans tout l'ensemble de ses dispositions, ne fit
qu'étendre, confirmer ou interpréter des canons antérieurs. Tous les
biens immeubles de l'église, ainsi que les esclaves et le bétail,
devaient être à la disposition de l'évêque, qui en faisait l'usage
prescrit par les canons. Si, dans une vue d'humanité, il abandonnait
pour un temps déterminé à des prêtres ou à des moines
l'exploitation de champs ou de vignes, aucune prescription ne
pouvait jamais éteindre son droit de propriété, et les dispositions de
la loi civile ne pouvaient pas être invoquées contre lui[257]. Quant
aux offrandes en nature que les fidèles faisaient sur l'autel, si c'était
dans la cathédrale, elles devaient se partager par moitié entre
l'évêque et le clergé de cette église[258]. Dans les églises rurales,
l'évêque avait droit à un tiers seulement, les deux autres tiers
appartenaient au clergé local[259]. Une question toute neuve, c'était
celle de la répartition des biens que l'Église devait à la libéralité de
Clovis, ou qu'elle en attendait encore. Fallait-il les soumettre aux
règles ordinaires, ou l'évêque pouvait-il en disposer à son gré? Le
concile répondit en rappelant les principes canoniques sur l'emploi
des revenus de l'Église: un tiers revenait au clergé pour sa
subsistance, un tiers aux pauvres et au rachat des captifs, un dernier
tiers à l'entretien des églises et du culte. Cette clause semblait dure
à certains prélats, qui, paraît-il, auraient voulu regarder les libéralités
royales comme des faveurs personnelles. Mais le concile s'éleva
avec force contre cette prétention; il menaça l'évêque récalcitrant
d'une réprimande publique de la part de ses comprovinciaux; s'il ne
se soumettait, il devait être exclu de la communion de ses frères
dans l'épiscopat[260]. Loin de pactiser ainsi avec l'égoïsme et
l'avidité de ses propres membres, l'épiscopat franc leur rappela dans
un canon spécial toute l'étendue de leur devoir de charité: L'évêque,
dit le seizième canon, doit, dans la mesure du possible, fournir les
aliments et les vêtements aux pauvres et aux infirmes que leur santé
empêche de travailler de leurs mains[261]. On sait quelle riche
variété d'œuvres charitables couvre l'ampleur magnifique de cette
formule, qui mettait dans la clientèle de l'Église toutes les misères et
toutes les souffrances d'ici-bas.
[261] Sirmond, p. 181; Maassen, p. 6.
Avant de se séparer, les évêques, Cyprien de Bordeaux et les autres
métropolitains en tête, signèrent les actes et en adressèrent une
copie au roi, avec une lettre ainsi conçue:
«A leur seigneur, fils de la sainte Église catholique, le très glorieux
roi Clovis, tous les évêques à qui vous avez ordonné de venir au
concile. Puisque un si grand souci de notre glorieuse foi vous excite
au service de la religion, que dans le zèle d'une âme vraiment
sacerdotale vous avez réuni les évêques pour délibérer en commun
sur les besoins de l'Église, nous, en conformité de cette volonté et
en suivant le questionnaire que vous nous avez donné, nous avons
répondu par les sentences qui nous ont paru justes. Si ce que nous
avons décidé est approuvé par vous, le consentement d'un si grand
roi augmentera l'autorité des résolutions prises en commun par une
si nombreuse assemblée de prélats[262].»
[262] Sirmond, p. 177; Maassen, p. 2.
Cette lettre était un acte de déférence de l'épiscopat envers la
majesté royale, ou, pour employer l'expression du concile lui-même,
c'était sa réponse au questionnaire de Clovis. On se tromperait si,
de la formule respectueuse de la fin, on tirait la conclusion que les
canons d'Orléans avaient besoin de la confirmation royale. L'Église,
chez les Francs mérovingiens, légiférait avec une souveraineté
absolue dans son domaine; ses canons étaient obligatoires en
conscience pour tous les fidèles, y compris le roi lui-même, et nul
n'aurait pu, sans se charger d'un péché grave, y contrevenir en
quelque matière que ce fût. Elle n'avait donc pas à demander à
Clovis une confirmation dont elle pouvait se passer; ce qu'elle
désirait, c'est qu'en se montrant disposé à y obéir lui-même, il
augmentât le prestige et l'autorité des résolutions conciliaires. D'en
faire passer la substance dans le droit civil, cela ne vint à l'esprit de
personne: c'est plus tard seulement, et dans une mesure d'abord
très restreinte, que les dispositions du droit ecclésiastique
commencèrent à y pénétrer. En attendant, les résolutions du concile
d'Orléans avaient force de loi pour l'Église franque, même celles qui
auraient été en contradiction avec le code[263].
[263] Voir Lœning, Geschichte des deutschen Kirchenrechts, t. II, pp. 150 et suiv.
Nous ne quitterons pas la mémorable assemblée de 511 sans faire
un rapprochement qui se sera sans doute présenté à l'esprit du
lecteur. C'est une œuvre législative qui a ouvert les annales des
Francs, et c'est une œuvre législative qui ferme le règne de Clovis.
Mais depuis les séances des quatre prud'hommes qui délibèrent
sous les chênes de Salaheim jusqu'à celles des trente-deux pontifes
qui siègent sous les voûtes du sanctuaire d'Orléans, quel chemin
parcouru! La loi salique est le code d'un petit peuple païen; les
canons de 511 sont la charte d'une grande nation chrétienne. Là, on
arrêtait le bilan de la barbarie; ici, on continue l'œuvre de la
civilisation. Là, un certain nombre de dispositions purement pénales
résument l'activité négative du passé; ici, les prescriptions positives
d'une loi morale supérieure font pénétrer dans le droit public les
influences fécondantes de l'avenir. L'histoire de la fondation de la
monarchie franque est comprise entre ces deux dates, et toute la
philosophie de cette histoire tient dans ce simple rapprochement.
VII
CLOVIS ET L'ÉGLISE
Il serait d'un haut intérêt, après avoir envisagé les sommets de
l'histoire de Clovis, de jeter un coup d'œil dans ses replis, et de
l'étudier dans la menue activité de la vie quotidienne. Combien elle
s'éclairerait pour nous, si nous pouvions joindre, à l'histoire de ses
exploits militaires, au moins quelques aperçus de son administration
et de son gouvernement! La pénurie de nos documents nous réduit
à ne presque rien connaître de ces sujets, qui prennent une place
capitale dans l'histoire de tant de souverains. C'est là ce qui rend la
vie de Clovis si difficile à écrire: elle finit chaque fois au retour d'une
campagne, c'est-à-dire là où les exigences de l'esprit moderne
voudraient la voir commencer.
Nous essayerons du moins, dans les pages qui vont suivre, de
grouper tous les renseignements qu'il a été possible de recueillir. Ce
sera la faute des matériaux et non celle de l'auteur, si le tableau
produit l'effet d'une mosaïque formée d'une multitude de fragments
rapportés.
De l'administration civile de Clovis, nous ne savons absolument rien.
Deux anecdotes, d'ailleurs fort légendaires, nous le montrent
conférant le duché de Melun à un de ses fidèles nommé
Aurélien[264], et le comté de Reims à un autre du nom d'Arnoul[265].
On n'a d'ailleurs pas besoin de ces indications pour admettre que
l'institution des ducs et des comtes de l'époque mérovingienne est
aussi ancienne que la dynastie elle-même.
[264] Eo tempore dilatavit Chlodovechus amplificans regnum suum usque
Sequanam. Sequenti tempore usque Ligere fluvio occupavit, accepitque Aurilianus
castrum Malidunensem omnemque ducatum regionis illius. Liber historiæ, c. 14.
Je ne garantis pas tout ce passage, que la présence du fabuleux Aurélien rend
justement suspect; mais j'admets, contre Junghans, p. 30, et Krusch, note de son
édition du Liber historiæ, p. 260, que l'auteur aura eu souvenance d'un comte de
Melun nommé Aurélien, et qu'il l'aura identifié avec le personnage de la légende.
Cet Aurélien historique était-il un contemporain de Clovis? On n'en peut rien
savoir.
[265] Ex Vita sancti Arnulfi martyris (dom Bouquet, III, p. 383).
Législateur, Clovis occupe dans les traditions de son peuple une

place qui n'est pas indigne du fondateur de l'État. La loi salique
n'existait jusqu'à lui que dans le texte germanique, arrêté par les
quatre prud'hommes de la vieille patrie. Selon le prologue de ce
célèbre document[266], il en fit faire, après son baptême, une
recension nouvelle, qu'il aura dépouillée de tout caractère païen.
Cette rédaction écrite en latin, sans doute à l'usage des habitants de
la Gaule romaine, a fait entièrement oublier l'ancienne version
germanique, et est seule arrivée jusqu'à nous, avec son escorte de
textes dérivés ou remaniés au cours des âges. Chose curieuse, pour
la Lex salica de Clovis, la terre franque, c'est le pays situé entre la
Loire et la forêt Charbonnière, c'est-à-dire la Gaule chrétienne et
civilisée qui était sa récente conquête. La France primitive, le pays
des vrais Francs germaniques, la terre de Clodion, de Mérovée et de
Childéric, ne compte plus, et l'on dirait quelle n'existe pas. Faut-il
donc croire que le roi des Francs soit devenu à tel point un étranger
pour sa propre race, qu'il n'ait plus même pris la peine de légiférer
pour elle? Non certes, et s'il n'est fait aucune mention de la mère
patrie dans le texte latin de la loi, c'est apparemment qu'elle restait
en possession de l'ancien texte germanique arrêté par les quatre
prud'hommes.
[266] V. la note suivante.
Le code élaboré par Clovis marque une nouvelle étape dans la voie
du progrès social chez les Francs. Il n'est pas la reproduction pure et
simple du texte germanique; il ne se contente pas non plus d'en
biffer les dispositions qui sentent trop l'idolâtrie, il le tient au courant,
si je puis ainsi parler, du développement total de la nation, devenue
un peuple civilisé depuis son introduction dans la Gaule romaine et
son baptême. «Ce qui était obscur dans le pacte, Clovis l'éclaira; ce
qui y manquait, il y pourvut[267].» Cette formule sommaire mais
expressive de la Loi salique nous laisse deviner une activité
législative qui a dû être considérable, mais que nous devons nous
résigner à ne connaître jamais.
[267] At ubi Deo favente rex Francorum Chlodeveus torrens et pulcher et primus
recepit catholicam baptismi, et quod minus in pactum habebatur idoneo per
proconsolis regis Chlodovechi et Hildeberti et Chlotarii fuit lucidius emendatum.
Prologue de la Loi salique. Pardessus, Loi salique, p. 345; Hessels et Kern, Lex
salica, p. 422.
Elle indique aussi ce que les monuments contemporains nous
montrent, à savoir, un prodigieux accroissement de la puissance
royale chez les Francs. Est-ce l'influence naturelle de ses conquêtes
et de ses victoires, est-ce la proximité de l'influence romaine, est-ce
le caractère sacré donné au pouvoir royal par la doctrine chrétienne,
ou bien plutôt ne sont-ce pas toutes ces raisons à la fois qui ont
placé le roi si haut au-dessus de son peuple? Il n'est plus le prince
tel que l'a connu la vieille Germanie; il est un maître dont le pouvoir
n'a pas de limites dans le droit, il est armé du ban, qui est la sanction
redoutable donnée par des pénalités spéciales à chacune de ses
volontés, il remanie et complète la législation avec une autorité
souveraine, et son præceptum suffit pour lui garantir l'obéissance.
Voilà la place conquise par le roi dans la vie du peuple franc. Celle
qu'il prend dans l'Église a un caractère spécial; il y exerce une
influence qui n'est égalée par nulle autre. Sans doute il n'est pas,
comme l'empereur, placé au-dessus d'elle pour la dominer, ni,
comme les rois ariens, en dehors d'elle pour la combattre. Il en fait
partie à la fois comme simple fidèle et comme souverain; fidèle, il
obéit à ses lois, il croit à sa doctrine; roi, et roi catholique, il écoute
les conseils de ses prélats, il la protège selon ses forces, il a sur sa
vie une action et une autorité qu'elle ne lui dispute pas.
Nous l'avons vu investi du droit de convoquer les conciles; mais ce
n'est pas tout. La première de ces assemblées qui se soit tenue
depuis sa conversion a subordonné à la volonté royale l'entrée des
hommes libres dans le clergé. En matière d'élections épiscopales,
sans jouir d'aucun droit canonique d'intervention, il dispose en fait
d'une influence considérable. Sans violer ni contester le libre
recrutement du sacerdoce, il y intervient avec une autorité à laquelle
tout le monde défère. Quand le roi catholique a dit quel homme il
veut voir mettre sur un siège épiscopal, il ne se trouve personne
pour être d'un autre avis, et de fait ce sera lui qui nommera l'évêque.
Le roi n'est-il pas lui-même membre de l'Église, et, si l'on peut ainsi
parler, son pouvoir électoral ne doit-il pas être en proportion des
intérêts qu'il représente? Nous le voyons, lors de la vacance des
sièges épiscopaux de Verdun et d'Auxerre, jeter les yeux sur des
hommes qu'il respecte, et leur offrir ces hautes charges, et c'est leur
refus seul qui empêche que sa volonté se fasse, mais en combien
d'autres occurrences elle aura eu force de loi! Ce qui semble pouvoir
être affirmé, c'est que, dans aucun cas, un siège épiscopal n'aurait
pu être donné contrairement à sa volonté. Au dire du biographe de
saint Sacerdos, ce prélat fut élevé au siège épiscopal de Limoges
par l'élection du clergé, aux acclamations du peuple, avec le
consentement du roi Clovis[268]. Voilà bien, désormais, les trois
éléments distincts qui constituent l'élection d'un évêque.
[268] Ex vita sancti Sacerdotis (dom Bouquet, III). Cette formule semble
empruntée au canon 10 du Ve concile d'Orléans en 549: cum voluntate regis, juxta
electionem cleri aut plebis (Maassen, Concilia p. 103). Mais il est manifeste que le
concile d'Orléans ne put que consacrer un état de choses antérieur, et il est
impossible de supposer que cet état de choses ne remonte pas au règne de
Clovis.
Un épisode bien authentique va nous montrer de fort près cette
situation de la royauté en face de l'Église, et la nature de l'influence
qui lui est reconnue. Sur la recommandation de Clovis, saint Remi
de Reims avait conféré les ordres sacrés à un certain Claudius. Cet
individu était probablement déjà suspect; après la mort du roi, il
donna un grand scandale. On voit qu'entre autres il avait
frauduleusement dépouillé de ses biens un nommé Celsus, et saint
Remi convient lui-même qu'il était coupable de sacrilège.
Néanmoins il intervint en sa faveur et demanda qu'il fût admis à la
pénitence, alors qu'aux termes du concile d'Orléans il devait être
excommunié. Cette indulgence lui valut d'amers reproches de la part
de trois évêques, Léon de Sens, Héraclius de Paris et Théodore
d'Auxerre. Autant qu'il est possible d'entrevoir leur attitude, ils
rendirent l'évêque de Reims responsable des fautes de son protégé;
ils lui firent notamment un devoir de rechercher et d'indemniser lui-
même les créanciers de Claudius; enfin, ils lui rappelèrent que si ce
malheureux avait pu jeter le discrédit sur sa robe, on le devait à la
pusillanimité de Remi, qui l'avait ordonné à la prière du roi et
contrairement aux canons. Dans sa réponse, qui nous a été
conservée, le saint se défend assez mollement sur la question du
fond; il convient d'ailleurs d'avoir déféré au désir de Clovis et
continue sur un ton énergique:
«Oui, j'ai donné la prêtrise à Claudius, non à prix d'or, mais sur le
témoignage du très excellent roi, qui était non seulement le
prédicateur, mais encore le défenseur de la foi. Vous m'écrivez que
sa demande n'était pas conforme aux canons. C'est le maître du
pays, c'est le gardien de la patrie, c'est le triomphateur des nations
qui me l'avait enjoint[269].»
[269] M. G. H., Epistolæ merovingici et karolini ævi, p. 114.
On ne prendra pas au pied de la lettre cette dernière expression,
inspirée au saint vieillard par le sentiment d'une détresse morale qu'il
ne parvient à cacher que d'une manière imparfaite à ses
contradicteurs. L'âpreté même de leurs reproches et la faiblesse de
ses excuses nous permettent de nous rendre un compte exact de la
situation qui est l'objet de cette correspondance. Clovis avait obtenu
de saint Remi un acte contraire à la législation canonique. On peut
mettre une bonne partie de la condescendance de l'évêque de
Reims sur le compte de ses relations spéciales avec Clovis. Le
pontife avait pour son royal filleul la tendresse d'un père, avec le
respect presque religieux qui lui faisait voir en Clovis l'instrument
manifeste de la Providence. C'était sa conquête à lui, c'était sa
gloire, c'était le fruit de ses sueurs. Toute sa pensée gravitait autour
de l'homme providentiel: qu'aurait-il refusé à son fils, à son
néophyte, à son roi? Il y a quelque chose de touchant à le voir, après
cinquante-trois ans de pontificat, obligé de défendre sa conduite
auprès de collègues plus jeunes que lui, et qui, comme il le leur
rappelle, lui devaient leur ordination. Mais ces confrères avaient pour
eux la lettre des canons, et ce débat entre évêques au sujet de
l'intervention du roi marque bien la distance qu'il y avait entre le droit
strict qui ne lui accordait rien, et la déférence qui lui cédait tout[270].
[270] Sur les élections épiscopales sous les Mérovingiens, il faut lire le bon
mémoire de M. Vacandard dans la Revue des Questions Historiques, t. LXIII
(1898), où est citée, p. 321, n. 1 et 2, la bibliographie antérieure.
Souvent même, c'est l'Église qui allait au-devant du roi, et qui le
sollicitait de trancher des questions, le prenant pour arbitre et
l'honorant de sa confiance. Lorsque saint Fridolin fut élu abbé de
Saint-Hilaire, à Poitiers, il hésita longtemps, nous dit son biographe,
à accepter cette dignité, malgré les instances de l'évêque saint
Adelfius; finalement, vaincu à demi par les prières de l'évêque, il lui
propose d'aller ensemble trouver le roi, pour qu'une affaire de telle
conséquence ne fût pas entreprise sans son concours. Et les voilà
qui partent tous les deux pour le palais royal, l'évêque à cheval,
comme l'exigeait son rang, l'abbé à pied, comme il faisait
d'habitude[271]. Ne voit-on pas comme un tableau en raccourci de
toutes les relations entre l'Église et l'État dans cet évêque et cet
abbé qui vont amicalement trouver le roi, pour le prier de les mettre
d'accord sur une question qui n'est pas de son ressort, mais qu'ils lui
soumettent par déférence et par respect?
[271] Ex vita sancti Fridolini (dom Bouquet, III, p. 388).
Un pareil degré de condescendance de la part de l'Église ne
s'expliquerait guère, si l'on ne savait qu'il était réciproque de la part
du roi. C'est la confiance qui formait la base des relations mutuelles.
Au lieu de délimiter anxieusement leurs frontières, les deux pouvoirs
semblaient s'inviter mutuellement à les franchir. Clovis convoquait
des conciles et intervenait dans les élections épiscopales; mais lui-
même, jusqu'à quel point ne se laissait-il pas inspirer, guider,
conseiller par les évêques? Toute sa politique intérieure, toute son
attitude vis-à-vis des indigènes, c'est l'épiscopat qui l'a dictée, et l'on
a vu plus haut que ce sont des évêques qui ont suggéré la
convocation du concile national. En un mot, son action sur l'Église a
pour contrepoids une action non moins énergique de l'Église sur
l'État. Les évêques composaient son conseil: saint Remi resta
jusqu'à la fin en grand crédit auprès de lui, et on nous dit que saint
Mélaine, évêque de Rennes, compta également parmi ses
conseillers les plus écoutés.
Toute l'hagiographie du temps est remplie des marques de respect
qu'il donna aux évêques. Les récits qui nous en ont gardé le
souvenir n'ont pas tous le degré d'authenticité nécessaire pour
s'imposer à la croyance du lecteur; mais dans l'impuissance où nous
sommes d'y faire le partage exact du vrai et du faux, quoi de plus
légitime que de les reproduire dans leur simplicité, comme des
documents qui ont droit tout au moins à l'attention de l'histoire? C'est
pour cette raison que nous avons cru devoir réserver une place, sur
ces pages, aux épisodes suivants.
Étant en Aquitaine, Clovis entendit parler des vertus de saint
Germier, évêque de Toulouse. Il le fit venir auprès de lui, l'invita à sa
table, et prit grand plaisir à sa conversation. Le saint distribua des
eulogies au roi et à ses grands; eux lui confessèrent leurs péchés et
écoutèrent ses exhortations à la pénitence. Le roi, voyant la sainteté
du prélat, le supplia de prier pour lui, et lui dit:
«Demandez-moi ce que vous voudrez de mes biens, et mes
serviteurs vous accompagneront pour vous le donner.
—Donnez-moi seulement, reprit le saint, dans le territoire de
Toulouse, autant de terre que mon manteau pourra en recouvrir
auprès de Saint-Saturnin, pour que je puisse dormir en paix sous la
protection de ce patron céleste.»
Mais le roi ne voulut pas se laisser vaincre en générosité: il donna
au saint la terre d'Ox avec six milles à la ronde, et, pour son
tombeau, il lui accorda tout le territoire que sept paires de bœufs
pourraient labourer en un jour. Toutes ces libéralités furent
consignées dans des chirographes que le roi et ses grands
scellèrent de leurs sceaux. Le roi y ajouta cinq cents sicles d'or et
d'argent, des croix d'or, des calices d'argent avec leurs patènes, trois
crosses épiscopales en or et en argent, trois couronnes dorées, et
autant de voiles d'autel en byssus. C'est ainsi qu'après être resté
avec le roi pendant une vingtaine de jours, le saint partit chargé de
trésors: le roi l'embrassa en lui faisant ses adieux, et se
recommanda à lui comme un fils[272].
[272] Ex Vita sancti Germerii (dom Bouquet, III, p. 386). Voir l'appendice.
Auch, la vieille cité métropolitaine de la Novempopulanie, a
enveloppé dans un récit aux couleurs bibliques le souvenir qu'elle a
gardé du héros franc. Lorsqu'il approcha de cette ville, dit une
tradition, l'archevêque saint Perpet alla à sa rencontre, et lui
présenta le pain et le vin, comme autrefois Melchisédech à
Abraham. Le roi récompensa magnifiquement le vieux pontife: il lui
donna toute la ville d'Auch avec ses faubourgs, et plusieurs églises;
il offrit également à l'église Sainte-Marie sa tunique et son manteau
de guerre; il lui offrit encore une aiguière d'or, et cent sous d'or pour
faire des couronnes de lumière; il lui assigna de plus un revenu de
cent douze sous d'or à toucher sur le fisc royal; il lui donna enfin
l'église royale de Saint-Pierre-de-Vic. Reconnaissante de tant de
libéralités, l'Église d'Auch célébrait tous les ans, au 3 juin, l'office
double de sainte Clotilde[273].
[273] La plus ancienne attestation de ce récit se trouve dans un acte de 1292,
consigné au registre des enquêtes du parlement de Paris et reproduit par R.
Choppin, De jure monachorum, p. 307; il figure aussi dans un extrait du cartulaire
du chapitre d'Auch, nº 132, reproduit en appendice, nº 7, dans de Brugèles,
Chronique ecclésiastique du diocèse d'Auch, Toulouse, 1746. Voir aussi Baiole,
Histoire sacrée d'Aquitaine, Cahors, 1644, p. 332; Loubens, Histoire de l'ancienne
province de Gascogne, Paris, 1839, pp. 90-91; Monlezun, Histoire de la
Gascogne, Auch, 1846, t. I, p. 189; Lafforgue, Histoire de la ville d'Auch, Auch,
1851. Selon Monlezun, l. c., une des couronnes faites avec l'or offert par le roi a
subsisté jusqu'en 1793; on l'appelait la couronne de Clovis.
Tournai racontait un épisode non moins intéressant. Attiré par la
réputation de l'évêque, saint Éleuthère, Clovis serait venu revoir la
vieille capitale de ses ancêtres, et assister à la prédication du prélat.
Mais une inspiration divine révéla au saint le tourment secret du roi:
il avait péché après son baptême, et il n'osait confesser sa faute.
Profondément ému, le roi essaya vainement de contester la vérité de
cette révélation que l'évêque lui communiqua; il versa des larmes, et
le supplia de prier pour lui. Et voilà que le lendemain, pendant que
l'évêque célébrait le divin sacrifice aux intentions de Clovis, un ange
du Seigneur lui apparut au milieu d'une lumière éblouissante, et lui
annonça que ses prières étaient exaucées. En même temps il lui
remettait un écrit contenant la faute secrète du roi. Clovis rendit des
actions de grâces à Dieu et à saint Éleuthère, et ne quitta Tournai
qu'après avoir comblé l'évêque de ses pieuses largesses[274].
[274] Vita sancti Eleutherii auctior dans les Acta Sanctorum des Bollandistes, 20
février, t. III, pp. 183-190, et Ghesquière, Acta Sanctorum Belgii, t. I, pp. 475-500.
Cette attitude vis-à-vis de l'épiscopat s'expliquerait déjà à suffisance
par des raisons d'ordre politique supérieur. C'étaient les évêques qui
avaient aidé le roi des Francs à établir son pouvoir; c'est par eux et
avec eux qu'il gouvernait. Il le savait, et sa déférence pour eux était
antérieure à sa conversion. Mais, après le baptême, des motifs de
piété s'ajoutèrent aux considérations de la politique pour augmenter
son respect envers les évêques. Il vit en eux des hommes qui
avaient reçu l'Esprit-Saint, et qui étaient les dispensateurs des
faveurs célestes. Leur science, leur sagesse, leur piété, leurs vertus,
la majesté de cette vie sacerdotale qui les élevait au-dessus de la
terre et qui faisait d'eux des hommes surnaturels, tout cela agissait
puissamment sur son âme, religieuse et impressionnable comme
toute âme de barbare. Il se sentait plus rapproché du Dieu qu'il
adorait dans leur société, et il comptait sur leurs prières comme sur
le moyen le plus efficace d'arriver au ciel. L'épiscopat, qui était le
point d'appui de sa politique, était aussi la sûre direction de sa
conscience de chrétien. Comme sa vie publique, sa vie privée
semblait la vérification de cette parole qu'il prononça un jour: «Où
serait l'espoir de vaincre, si nous offensions saint Martin?» Entendez
ici, par saint Martin, l'épiscopat de la Gaule.
Les mêmes sentiments, au dire de la légende, dictaient la conduite
du roi vis-à-vis de toutes les personnes qui, sans occuper un rang
dans la hiérarchie ecclésiastique, se distinguaient par l'éminence de
leurs vertus. Il croyait, avec tous ses contemporains, à l'efficacité de
leurs prières; il était convaincu qu'elles avaient le don d'opérer des
miracles. Lui-même, au dire d'un hagiographe, fut favorisé d'une
guérison miraculeuse obtenue par l'intercession d'un vénérable
solitaire. C'était la vingt-cinquième année de son règne, celle qui
allait être rendue mémorable par la conquête de l'Aquitaine. Il y avait
deux ans qu'il était en proie à la maladie, et ni les prières de son
clergé ni les soins de ses médecins ne parvenaient à le soulager.
Enfin, l'un de ces derniers, nommé Tranquilinus, conseilla au roi de
faire venir Séverin, abbé de Saint-Maurice en Valais, homme doué
de l'esprit de Dieu, et dont les prières obtenaient une multitude de
guérisons miraculeuses. Aussitôt le roi fit partir son chambellan
Transoarius pour Agaune, et le saint, déférant à ses prières, apparut
au chevet du royal malade comme plus tard saint François de Paule
auprès du lit de Louis XI. Après avoir adressé au ciel de ferventes
prières, il ôta son manteau, en revêtit le roi, et à l'instant la fièvre
abandonna le malade. Clovis, plein de reconnaissance, tomba aux
pieds du saint, et le pria de prendre dans son trésor toutes les
sommes qu'il voulait pour les distribuer aux pauvres; il lui offrit aussi
de faire relâcher tous les coupables qui se trouvaient enfermés dans
les prisons[275]. On veut que l'église Saint-Séverin de Paris, qui est
sous le patronage de l'abbé d'Agaune, ait été élevée en souvenir de
cet heureux événement.
[275] Ex Vita sancti Severini Abbatis Agaunensis (dom Bouquet, III, p. 392.) Ce
récit est loin d'être garanti, bien qu'il en soit souvent fait état même par des
historiens peu tendres à l'endroit des légendes, notamment par Junghans, p. 77,
n. 1, par W. Schultze, Das Merovingische Frankenreich, p. 72, et en dernier lieu
par Arnold, Cæsarius von Arelate, p. 242. Voir l'Appendice.
D'autres saints personnages, au dire de la légende, ont été en
rapports intimes avec Clovis. Saint Fridolin de Poitiers, admis à sa
table, a réparé miraculeusement une belle coupe de verre, qui s'était
cassée en tombant des mains du roi au moment où il la présentait
au saint[276]. Un saint ermite du nom de Léonard, qui demeurait
dans la forêt de Panvain, près de Limoges, fit la connaissance du roi
dans des circonstances fort dramatiques. Clotilde, qui était venue
résider avec son époux dans le château de cette forêt, était
menacée de périr dans les douleurs de l'enfantement, et Clovis au
désespoir implora le pieux solitaire de venir à son aide. Léonard se
mit en prières, et la reine fut sauvée par miracle[277].
[276] Ex Vita sancti Fridolini (dom Bouquet, III, p. 388).
[277] Arbellot, Vie de saint Léonard, solitaire en Limousin, Paris, 1863. Les pages
277-289 contiennent le texte d'une vie inédite de saint Léonard, d'après plusieurs
manuscrits dont un du onzième siècle.
Sans doute, la plupart de ces récits ont été embellis par la pieuse
imagination des hagiographes, et il n'est pas interdit de croire que
les épisodes qui sont à la fois les plus extraordinaires et les moins
prouvés appartiennent au domaine de la fiction pure.
Ce qui se dégage des plus authentiques, c'est l'intimité des rapports
entre le roi et les saints, c'est la justesse de l'instinct qui poussait la
royauté à se rapprocher de ceux qui représentaient le mieux les
aspirations chrétiennes de leur peuple. Avec un admirable sentiment
des vrais intérêts de sa couronne, Clovis se mêlait familièrement,
sans crainte de compromettre son prestige, aux hommes humbles et
pauvres revêtus d'une majesté supérieure par le respect public, et le
nimbe de leur sainteté jetait une partie de son éclat sur le front du
souverain. Rien n'a plus contribué à sa popularité que l'amitié des
saints. Les actes de clémence qu'ils lui inspiraient affermissaient son
pouvoir en lui ouvrant les cœurs. Bien des fois, saint Remi et sainte
Geneviève arrachèrent au rude justicier la grâce des malheureux qui
remplissaient les prisons publiques. Parmi ceux que menaçait sa
vengeance, il y avait un grand seigneur du nom d'Euloge, qui se
réfugia dans l'église Notre-Dame de Reims: à l'intercession de Remi,
le roi lui laissa la vie et la possession de ses biens[278]. Au dire d'un
hagiographe, l'évêque aurait même obtenu du roi que chaque fois
qu'il passerait par la ville de Reims ou par son territoire, tous les
prisonniers seraient aussitôt mis en liberté, et, ajoute-t-il, cet usage
se conserve encore aujourd'hui[279].
[278] Hincmar Vita sancti Remigii, dans les Acta Sanctorum des Bollandistes, 1er
octobre, t. I, p. 153 A.
[279] Vie de saint Léonard, éditée par le chanoine Arbellot, c. 3.
Les vastes ressources de la couronne permirent au roi de témoigner
de la manière la plus efficace sa bienveillance à l'Église en la
comblant de ses dons, en venant à son aide dans ses œuvres de
charité et dans ses créations de tout genre. Il faut se souvenir que la
générosité était la première vertu d'un roi germanique. Sa main
devait toujours être ouverte, excepté quand elle brandissait l'épée. Il
passait sa vie à faire des cadeaux, à distribuer à ses amis l'or
travaillé sous forme de bracelets à tours multiples, dont il détachait
les morceaux, et, pour la poésie barbare, il était avant tout le briseur
d'anneaux. Lorsqu'avec les pièces de métal précieux entassées
dans ses trésors, il put disposer aussi des domaines sans nombre
que la conquête avait fait tomber entre ses mains, alors il eut de
nouveaux moyens d'être généreux, et la série des donations de
terres commença. L'Église fut au premier rang des amis qui
participèrent à ces libéralités. On peut dire, sans crainte de se
tromper, que tous les diocèses eurent leur part[280]. Après la
conquête de la Gaule romaine, après celle de la Gaule visigothique,
il s'ouvrit comme deux phases d'abondance qui furent employées à
prodiguer les richesses aux églises. Les actes du concile d'Orléans
parlent expressément des libéralités royales faites ou promises à
tous les diocèses[281]. L'hagiographie ne nous mentionne pas une
seule fois les relations du roi avec quelque saint sans nous faire
connaître les cadeaux dont il le combla. Nous l'avons vu prodiguer
ses dons aux églises Saint-Martin de Tours et Saint-Hilaire de
Poitiers; nous l'avons vu enrichir aussi généreusement saint Germier
de Toulouse et saint Perpet d'Auch; nous savons avec quelle
libéralité il aida saint Eptade à racheter les captifs. Il ne fut pas
moins prodigue envers saint Mélaine de Rennes, qui put faire une
multitude de bonnes œuvres avec les ressources que le roi mettait à
sa disposition[282]. L'église de Vannes se glorifiait de devoir à ses
pieuses largesses le trésor de reliques qu'elle conservait depuis les
jours de saint Paterne, son premier évêque[283]. L'église de Nantes
montrait avec orgueil, dès le douzième siècle, la charte contenant
les faveurs dont l'avait comblée le premier roi de France[284]. Ce
serait une tâche fastidieuse que de relever, dans les biographes et
les chroniqueurs, les récits souvent légendaires qui nous ont
conservé la trace de toutes ces générosités, et il suffit de dire d'une
manière générale que Clovis partagea largement avec l'Église les
richesses considérables qui affluaient de toutes parts dans son
trésor et dans son domaine.
[280] Illi (il s'agit surtout de Clovis) monasteria et ecclesias ditaverunt; isti (il s'agit
de ses petits-fils) eas diruunt ac subvertunt. Grégoire de Tours, iv, 48.
[281] De oblationibus vel agris quos domnus noster rex ecclesiis suo munere
conferre dignatus est, vel adhuc non habentibus Deo inspirante contulerit, ipsorum
agrorum vel clericorum immunitate concessa, id esse justissimum definimus ut...
Sirmond, Concilia Galliæ, I, p. 179; Maassen, Concilia ævi merov., I, p. 4.
[282] Vita sancti Melanii, dans les Acta Sanctorum des Bollandistes. Sur les divers
textes de cette vie, voir l'Appendice.
[283] Un sermon prêché dans la cathédrale de Nancy au douzième siècle et
conservé dans le manuscrit 9093 latin de la bibliothèque nationale à Paris contient
le passage suivant: Circa initia etiam hujus nascentis ecclesiæ, divinæ
misericordiæ dulcor in hoc se aperuit quod Clodovæus rex Francorum
illustrissimus per beatum Paternum patronum nostrum transmisit huic ecclesiæ
desiderabilem thesaurum videlicet etc. Suit une énumération de reliques. V. A. de
la Borderie, Histoire de Bretagne, t. I. p. 204. note 2 et p. 331.
[284] V. dans Dom Morice, Mémoires pour servir de preuves à l'histoire de
Bretagne, t. I, p. 547, le texte de la charte de Louis-le-Gros, datée de 1123, dans
laquelle sont rappelées les libéralités de Clovis; on y lit: «Quoniam vir venerabilis
Bricius Namneticæ sedis episcopus præsentiam nostram non absque magno
labore itineris humiliter adiit et præcepta antiquorum et venerabilium Francorum
regum Karoli, Clodovæi et filii ipsius Clotarii attulit et ostendit, etc. L'authenticité de
cette charte, contestée par Travers, Histoire de la ville et du comté de Nantes, I, p.
244 et par M. C. Port, Dictionnaire de Maine-et-Loire, t. II, s. v. Loué, est défendue
par M. L. Maître, Étude critique sur la charte du roi Louis VI, Rennes 1887, et par
M. A. Luchaire, Louis VI le Gros, Annales de sa vie et de son règne, pp. 153, 323
et suivantes. Au surplus, les défenseurs de l'authenticité ne sont pas d'accord sur
la personne de ce Clovis, père de Clotaire, car cette désignation convient aussi
bien à Clovis II qu'à Clovis Ier; bien plus, si l'on admet qu'ici Clodovæus équivaut à
la forme Hludovicus usitée au onzième siècle, on peut penser à l'une des séries
royales Charlemagne, Louis le Débonnaire et Lothaire, ou encore Charles le
Simple, Louis d'Outre-Mer, Lothaire. M. Luchaire penche pour une de ces
dernières hypothèses, M. Maître, o. c. et M. Orieux (Bulletin de la Société
archéologique de Nantes, t. 39, 1898, p. 59), pensent à Clovis Ier. Selon moi, le
rédacteur de l'acte, authentique ou non, n'a pu penser qu'à un Clovis, et je suis
porté à croire que c'est Clovis Ier.
De tous les prélats sur lesquels il fit pleuvoir ainsi les preuves de sa
munificence, le plus favorisé fut naturellement saint Remi de Reims.
Dès le neuvième siècle, nous entendons la tradition énumérer les
dons qu'il tenait de son généreux filleul. Ils consistaient surtout en
domaines territoriaux, répartis dans plusieurs provinces de la
France. Le saint ne voulut en garder que quelques-uns, situés dans
la partie orientale du royaume, et distribua le reste aux autres
églises, pour qu'on ne pût pas lui reprocher de faire de l'amitié du roi
une source de profits[285]. Toutefois l'église de Reims gardait dans
son trésor un encensoir et un calice émaillé provenant, selon la
tradition, d'un grand vase en argent que Clovis avait donné à saint
Remi, pour en faire ce qu'il voulait[286].
[285] Hincmar, Vita sancti Remigii, 66, dans les Bollandistes, p. 149 C.

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Adventures in Biophysics A. V.

Foulerton Research Professor of the Royal Society

If you go to Devonshire in September and wander in the

SOME ADVENTURES WITH VAPOUR PRESSURE

N the spring and summer of 1927, while I was at Cornell,

effect was held by Embden and his colleagues to disprove

motic pressure between the bloods of men and women!

6/772. cm. 4000

nerve), or some anaerobic process such as the formation of

making up the waves. So also in biology: " l i f e " in the

broken: the rapidity of lactic acid formation in injured

cells were p u t out of action. If this were so, the increment

Fresh muscle initially in N 2 : resting heat rate,

What could this be other than an effect of oxygen in

books. M y friend Professor Meyerhof heard about it

mixture at normal atmospheric pressure. Oxygen diffusion

through the fluid in which the muscle lay the attainment

altogether by filling the muscle c h a m b e r w i t h paraffin oil.

the chamber. Incredible as it seemed the riddle had

the removal of lactic acid, to restoration of " p h o s p h a g e n , "

discovery of great importance: a living cell at constant

vapour pressure (for not too great concentrations) by the

function of the heat produced in the activity induced by

With regard to the latter, so far as the lines of fig. 3 are

" p h o s p h a g e n " molecule —> creatine + phosphate,

where X is some other unknown substance, then two new

and above all those known at present to chemical analysis.

fatigue is 2.8 times as great as the osmotic pressure of the

molecules for every atom of Ρ set free. According to

Now 0.095 Μ ' s sum m

X — phosphagen —1• X + phosphate + creatine.

recently been confirmed by Meyerhof (1930b) in experi-

to have a very high osmotic pressure. This should rapidly

M y own blood, withdrawn 60 seconds after 1 minute only

treme severity lasting for a minute or two in a first rate

THE STATE OF W A T E R IN TISSUES

O R the calculations given in the last lecture we assumed

be explained by the simple hypothesis that the " f r e e "

Their explanation, however, as I have shown in a recent

The frog's muscle was cooled to — i8° C. and 100 g. of

— 20° C . as at the ordinary t e m p e r a t u r e s in which we are

Gortner (1922), who added a known amount of cane sugar

incidentally to study blood, the body fluids of inverte-

Hb 12.5 15.6 Ο.ΟΟ23 Molecular weight assumed 67000

Na 0.2460 0.3060 O.I33J —

Κ 0.0790 0.0980 0.0252 —

Ca 0.0044 0.0055 O.OO IO Assumed 70 p.c. as free Ca ions

CI 0.2890 0.3604 O.IOI5 —

Total Ρ 0.0314 0.0392 0.0088 See note

A b d e r h a l d e n ' s list, h o w e v e r , a l t h o u g h it m a k e s u p nearly

Législateur, Clovis occupe dans les traditions de son peuple une

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