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COMPARATIVE
ELECTRO-PHYSIOLOGY
PLANT RESPONSE;
as a
logical Investigations.
Svo. 21s.
1906.
39 Paternoster
Row, London,
Calcutta
.<?t-
COMPARATIVE
ELECTRO-PHYSIOLOGY
A PHYSICO-PHYSIOLOGICAL STUDY
BY
JAGADIS
CHUNDER
BOSE,
M.A., D.Sc.
WITH ILLUSTRATIONS
5^
NEW
All
riRbts
reserved
CO.
PREFACE
This volume concludes the
phenomena
sive
publication of a
in general,
Memoir
which
first
commenced with
the
of
my
publications on the
on respon-
In this
subject
line of investigation
employed
for obtaining
my
at
response-records was
With the
in
object of showing
reaction,
Believing, as
responsive phenomena,
succeeded
in
common
had already
stances, to test
also,
meaning those
The
De
la Gen^ralite
stimulation
la
'
'
COMPARATIVE ELECTRO-PHYSIOLOGY
VI
many
to
quantitative, thus
By
sources of complication.
show
this
was able
As
method
obviating
summary
observations similar
May
7,
and afterwards
my
of
read, with
from the
results
communicated
to
the Royal
accompanying ex-
An
By methods somewhat
tissues.
above,
is
that given
The response
Mimosa, but
1901.
by
inor-
by plant
to mechanical stimulus.
6,
is
is
response
electric
not confined to
universally present.
and leaves
stems,
white
lily,
injury "
is,
of,
among
rhubarb, and
generally
uninjured part.
"
others,
horse-radish.
speaking,
horse-chestnut,
The
from the
negative variation
injured
" is also
vine,
" current
of
the
to
produced.
Very
stimulants,
and of poison.
by poison
response.'
may be well to
when
this
'
I>rEFacE
changes, was
electro-motive
definite
the reading of
my
Paper, on June
Sanderson went so
as
far
regarded
to
1901, Sir
6,
that
state
as
highly-
which followed
the discussion
Indeed, in
controversial.
vii
John Burdon
excitatory
this
an impossibility.
My
which
effects
had shown
to occur
commonly assumed
tinction
my work
between the
to exist
These
results
'
sensitive
'
were published
effects of various
Many
anomalous
effects hitherto
ascribed to
anisotropic
structures,
and
shown that
Among
was
all
other things,
it
in reality derived
'
of
was
from
autonomous response
either to the
It
movements
initiated
differential excitability
to
internal stimulus
specific sensibilities
subsequent ex-
movements, were
its
more
and that
COMPARATIVE ELECTRO-PHYSIOLOGY
Vlll
were thus
With regard
to these points,
my results
have been
in direct
induced by stimulus
is
From
firing-off of a
all
the
it
run-down of energy.
accompanied by an inevitable
and
incessant
ment
as
into
all
it
gigantic
of
environment.
its
the
is
agent that
the
is
the
know,
is
stimuh
multifarious
we
which
embryo
Rather,
analogy of the
plausible
of a combustion-engine,
that
fashions
microscopic
the
And
banyan-tree.
it
clear
is
able
to
by means of
corroborate,
many more
in various directions, so as to
was
my
with
original inten-
of Plants.
the
response, the
electrical
same methods of
mental devices,
in the
inquiry,
pos-
proceeded to apply
one case as
in the other.
have thus
PREFACE
IX
The
animal.
for
the
elucidation
of
problems
biological
Exception
sufficiently obvious.
to the
most complex
the
in
may
in general,
is
must
It
on the
general
in
My mode
determined by the
attempt
it
will
necessary progression
my
conviction
results,
to the
as
And
between them.
existed
of investigation
from
this
which, accord-
were anomalous,
capable of an
and
are, in
fact,
factory explanation.
increasingly simple
must
It
its
extension
also be understood
electrical
the
into
field
satis-
that
my
response of plants,
of Animal
Electro-
tinuity
my
It
disposal,
references
to
These
will
be found,
account given
in
in
the
all
their
detail, in
tissues.
the excellent
COMPARATIVE ELECTRO-PHYSIOLOGV
JC
ment, which
The
have
reactions
forth
set
of different
garded as specifically
in
tissues
different.
As
the
present volume.
a continuity
this,
typically
re-
Thus, nerve
non-motile
re-
its
nerve
is
investigation of
its
method
by the
that
The
electrical.
error,
is
than
It
But
tion.
this
view
is
incorrect.
Experi-
is
So complete,
indeed,
has
that
similarity
this
observation in
its
the
explanation of a
plant, has
Many anomalous
Now
organ,
is
simple plant
I
have been
PREFACE
able
show that
to
XI
excitability
differential
this
widely
is
posed to
known
it
electrical
Few
and that
conclusions in
fishes.
rest
as Pfluger's
Law
law
that this
is
enunciated by
Pfliiger.
that
Finally,
lie
it
But
visual scrutiny.
actually attended
is
beyond
has here
by change
This
single,
is
fact,
lies
In
read
In
'
said
phenomena
the
breach of continuity.
say
"
described
It
is
above
difficult
to
there
draw a
is
line
little
and
or "
is
organisms
"
'
or
That
is
phenomenon of inorganic
lines of
demarcation that
COMPARATIVE ELECTRO-PHYSIOLOGY
xil
of psychical
Such
processes."
can hardly
arbitrary lines
be drawn.
'
We may explain
finding
in the living
common
persistent
this
many
And for
phe-
nomena which
it
may be
or,
at first
said
that
been to attempt to
appear so
the
find,
different.
this
wherever
facts justify
it,
an under-
was
It
that
And
now,
for the
out
set
in
publication,
bringing to
its
close
exf)ress
the
regions of inquiry
may
many
had at one time been regarded as beyond the scope of experimental exploration.
I
efficient
my
C.
BOSE.
CONTENTS
CHAPTER
to stimulus
of solubility
by change of form
negative variation
modifications
Response
PAGE
(d)
CHAPTER
II
Difficulties of investigation
Electrical
response of pulvinus of
electrical response of
lation
and
Division of
Mechanical
and
and transmitted stimuAll forms of stimulus induce excitatory change of galvanometric
into
plants
negativity
'
ordinary
'sensitive'
plants
'
arbitrary
Direct
CHAPTER
.12
III
on period of vibration
Additive
Response
COMPARATIVE ELECTRO-PHYSIOLOGY
xiv
electric
Effect of season
CHAPTER
29
IV
Instantaneous
mechanical
Tabular
..........
statement of results of
multiple responses
44
CHAPTER V
THE ELECTRICAL INDICATIONS OF POSITIVE AND NEGATIVE
TURGIDITY-VARIATIONS
Motile responses of opposite signs, characteristic of positive and negative
hydrostatic effect of stimulus causes
turgidity-variations - Indirect
Antagonistic elements
in the
electrical
effect
CHAPTER
^^
VI
Hydraulic transmission of energy in plants True meaning of tonic conditionOpposite expressions of internal energy and external stimulus
seen in growth-response Parallelism between responses of growing and
motile
Sudden
retardation of growth
of spiral tendril;
and
electrical response
uncurling
Laws
movement
of mechanical
gp
CONTENTS
CHAPTER
XV
VII
and by intensity of
Sub-tonic, normal and hyper-tonic conditions The
Outward
external stimulus
critical
level
critical
level
is
tissue,
when
exceeded
Three
typical cases
Investigation
CHAPTER
76
VIII
Similar
responsive
effects
seen
in
inorganic
matter
tonic tissue
excitability
stimulus
Volta-chemical
nating fatigue
effect
Rapid
Instances
....
latent
CHAPTER
staircase
and
86
IX
Anomalies in mechanical and electrical response Resultant response determined by differential excitability Responsive current from the more to
Laws of response in anisotropic organ Demonstraby means of mechanical stimulation Vibrational stimulus Stimulation by pressure
Quantitative stimulation by thermal shocks
the less excitable
tion
CHAPTER X
THE NATURAL CURRENT AND
Natural current in anisotropic organ from the
ITS
VARIATIONS
less to the
more excitable
107
COMPARATIVE ELECTRO-PHYSIOLOGY
xvi
tion,
the negative
variation
Action
of hydrochloric acid
Action
of
Re-
CHAPTER
.116
XI
studied
excitability
transmitted stimulation
Effect
by two methods:
of chloroform
direct
(i)
Effect of chloral
negative variation
KHO
CHAPTER Xn
VARIATIONS OF EXCITABILITY DETERMINED BY METHOD OF
INTERFERENCE
Arrangement
for
interference
difference of phase
of excitatory waves
Interference
effects
Effect
of increasing
Diametric
balance
Effect of
.
141
CHAPTER Xni
CURRENT OF INJURY AND NEGATIVE VARIATION
Different theories of current of injury
Reymond
Electrical
Pre-existence
theory of
distribution in a muscle-cylinder
Hermann's Alteration
is
Du
Bois-
Electro-mole-
Theory Experiments
a persistent after-effect
electrical
in
in
'
'
current of injury
149
CONTENTS
XVll
CHAPTER XIV
CURRENT OF DEATH
Anomalous
RESPONSE
BY POSITIVE VARIATION
and dead
Inquiry into
tissue
PAGE
the cause
Dying
living
tissue
Explanation of
.164
CHAPTER XV
EFFECT OF TEMPERATURE ON ELECTRICAL RESPONSE
General observation of
effect of
temperature on plant
Similar
Effect of and
Effect of
temperature in diminishing
rise of
hanced response as
frost
Effect of
diminution of excitability
in
effect
rise
fall
Desmodium
180
CHAPTER XVI
THE ELECTRICAL SPASM OF DEATH
Different post-mortem
death-point
normal
electrical
death-spasm
electrical
Determination
:
determination of
From
spasm
for
(a) in anisotropic
methods Tabulation
of observations
.192
CHAPTER XVII
MULTIPLE AND AUTONOMOUS ELECTRICAL RESPONSE
Transition
COMPARATIVE ELECTRO-PHYSIOLOGY
xviii
PAGE
its
time-
pulsations in
leaflet
............
ing plants
So-called
CHAPTER
207
XVIII
RESPONSE OF LEAVES
and-stalk currents
shown
of petiole on Z)zV?^a
True explanation
of
J/i^j-a
results
Experi-
.."
Parallel experiments on
223
CHAPTER XIX
THE LEAF CONSIDERED AS AN ELECTRIC ORGAN
Electrical organs in fishes
Typical instances,
or pitcher of Nepenthe
Response
dromous
shown to
electrical
to direct
excitation
to
to differential
.241
CHAPTER XX
THE THEORY OF ELECTRICAL ORGANS
Existing theories Their inadequacy
sponse
uni-directioned,
to
Uni-directioned
The
blaze -current
'
so called
Re-
Mimosa
Re Of Eel
Multiple
Multiple responses of Biophyiuin
organ
'
response of pulvinus of
sponse of petiole of
The
Cucurbita
responses of electrical
259
CONTENTS
XIX
CHAPTER XXI
DETERMINATION OF DIFFERENTIAL EXCITABILITY UNDER
ELECTRICAL STIMULATION
PAGE
Advantage of
electrical stimulation,
due
to
fluctuating
Difficulties
sponse of
Musa
to equi-alternating shocks
plagiotropic
and Eel
by electric
Cuctirbita
............
response
272
CHAPTER XXH
RESPONSE OF ANIMAL AND VEGETAL SKINS
General
formula for
all
types of response
of
skin
(2)
skin
of Direct Effect
Response of grape
{c)
chanical
may be
response
tomato
Mimosa ^\s.z.'Aq.
pulvinus of
response of Des7nodium
True
variation
gyrans
in
Autonomous
{b) frog's
variation
Normal
autonomous me-
of skin of neck
of tortoise
Electrical
gecko Explanation
current of rest
of
Method
of frog's skin
of
This
Electrical
response of skin of
Response
of skin
of abnormal response
287
CHAPTER XXni
RESPONSE OF EPITHELIUM AND GLANDS
Epidermal, epithelial, and secreting membranes in plant tissues Natural
resting-current from epidermal to epithelial or secretory surfaces Current
of response from epithelial or secretory to epidermal surface^;
Response
COMPARATIVE ELECTRO-PHYSIOLOGY
X5i
of Dillenia
foot of snail
Natural current
'
due
The
PAGE
to injury
'
positive variation
direction
of responsive
characteristic curves
currents,
diagrammatically represented in
-312
CHAPTER XXIV
RESPONSE OF DIGESTIVE ORGANS
Drosera
Response
Multiple
response in
Re-
329
CHAPTER XXV
ABSORPTION OF FOOD BY PLANT AND ASCENT OF SAP
Parallelism between responsive reactions of root and digestive
organ Alter-
as
in
pitcher
of A<f/^M^
Different
phasic
chemical
reactions,
Response
to
sap Physical
Strasburger's experiments on
Current inference unjustified
sponse of sap-wood
ascent of sap
effect
of poisons on
349
CONTENTS
xxi
CHAPTER XXVI
THE EXCITATORY CHARACTER OF SUCTIONAL RESPONSE
PAGE
root killed
namely (i) the unbalanced, (2) the balanced, (3) the over-balanced
Renewal of suction previously at standstill, by action of stimulus- -Reponsive enhancement of suction by stimulus After-effect of stimulus
Response under
over-balance Response under sub-terminal stimulation Variation of
response under seasonal changes
after-effect of stimulus
365
CHAPTER XXVII
RESPONSE TO STIMULUS OF LIGHT
Heliotropic plant
movements reducible
to
tion or expansion
growing
concomitant to excitatory
not determined by
effects
Electrical
presence or absence of
response
due
excitation
Mechanical
to indirect
effect
chloroplasts
Effect
of
response of leaf of
Mimosa
to light applied
on
induction in
Mimosa
response of leaf of
to light applied
effect
effect,
of light and
positive
after
effect
Plants
by negative response
growth response Examples of
Exemplified by
(a) electrical
and
Periodic
{b)
variation of excitability
Multiple
under
-I-
-f-
392
COMPARATIVE ELECTRO-PHYSIOLOGY
xxii
CHAPTER XXVIII
RESPONSE OF RETINA TO STIMULUS OF LIGHT
Response of retina
rest
Determination
pa(;e
and
retina
The
so-called
positive
variation
of
previous observers
true
excitatory
response in
human
.415
CHAPTER XXIX
GEO-ELECTRIC RESPONSE
Question regarding
expansion Crucial experiment by local application of cold Reasons for
of Statoliths
shoot
Due
to active contraction of
vanometric negativity
upper
Geo-electric
Geo-electric response
side,
of
response of an organ
physically
restrained
434
CHAPTER XXX
DETERMINATION OF VELOCITY OF TRANSMISSION OF
EXCITATION IN PLANT TISSUES
Transmission of excitation in plants not due to hydromechanical disturbance,
but instance of transmission of protoplasmic changes Difficulties in
accurate determination of velocity of transmission
A perfect method
of rise
Effect
of temperature
apd negative
.ft
444
CONTENTS
XXlll
CHAPTER XXXI
ON A NEW METHOD FOR THE QUANTITATIVE STIMULATION
OF NERVE
PAGE
Drawbacks
Gradual
from positive to negative, through intermediate diphasic
ConEffect of depression of tonicity on excitability and conductivity
version of abnormal into normal response by increase of stimulus-intensity
negative after tetanisation
transition
456
CHAPTER XXXn
ELECTRICAL RESPONSE OF ISOLATED VEGETAL NERVE
Specialised conducting
tissues
Isolated
vegetal
{c)
(a) action
nerve
Method
of
ob-
Similarity of responses of
[d) action of
of carbonic
ammonia
and
{e)
ex-
positive
....
Effect of
468
CHAPTER XXXIII
THE CONDUCTIVITY BALANCE
conductivity, and responsivity Necessity
distinguishing
Ad vantages of the Method of Balance Simultaneous comparison of
variations of receptivity, conductivity and responsivity The Conductivity
Receptivity,
for
these
Balance Effect
of
NajCOg on
frog's
nerve Effect
of
CuSO^ Effect
conductivity, velocity
responsivity.
.^
,
479
COMPARATIVE ELECTRO-PHYSIOLOGY
XXIV
CHAPTER XXXIV
EFFECT OF TEMPERATURE AND AFTER-EFFECTS OF STIMULUS
ON CONDUCTIVITY
PAGE
Effect of temperature in inducing variations of conductivity
{a)
by Method
of Mechanical Response
cold
Effect
of rising
497
CHAPTER XXXV
MECHANICAL RESPONSE OF NERVE
Current assumption of non-motility of nerve Shortcomings of galvanometric
modes
electric
of detecting excitation
shocks
Optical
mechanical response
Kunchangraph
Of chloroform
Mechanical
to transmitted stimulation, in
gecko
507
CHAPTER XXXVI
MULTIPLE RESPONSE OF NERVE
cessation of tetanisation
of
nerve by drying
CHAPTER XXXVII
RESPONSE BY VARIATION OF ELECTRICAL RESISTIVITY
Variation of resistance in Dtonaa, by
its
'
modification
'
Characteristic
Excitatory
change,
difficulties of investi-
532
CONTENTS
Morographic
XXV
curves at death-point
record
Similarities
540
CHAPTER XXXVIII
FUNCTIONS OF VEGETAL NERVE
Excitability
and motile
Physiological
sensibility,
-SSI
CHAPTER XXXIX
ELECTROTONUS
Extra-polar effects of electrotonic currents on vegetal
variation of excitability
nerve Electrotonic
increment Investigation into the law of electrotonic variaInvestigation on variation of excitability Conductivity enhanced when excitation travels from places lower to higher
electric potential, and depressed in opposite direction
When feeble,
anode enhances and kathode depresses excitability All electrotonic
phenomena reducible to combined action of these factors Explanation
polarisation
tion of conductivity
of apparent anomalies
..........
5^
CHAPTER XL
INADEQUACY OF PFLUGER'S LAW
Law under high E.M.F. Similar reversals under
Investigation by responsive sensation Experiments on
living
Under moderate E. M. F. intensity of sensation enhanced
kathode, and depressed at anode Under feeble E.M.F., sensation
Reversal of Pfliiger's
feeble
E.M.F.
wounds
at
intensified at
at
578
COMPARATIVE ELECTRO-rHYSIOLOGY
XXVI
CHAPTER XLI
THE MOLECULAR THEORY OF EXCITATION AND
ITS
TRANSMISSION
PAGE
Two
opposite
responsive
and
negative
manifestations,
positive
Such
Arbitrary nature of term excitatory Pro-excitatory and anti-excitatory agents Molecular distortion under magnetisation in magnetic sub'
'
stances
Diff"erent
forms of
response under
magnetic stimulation
Uniform
groupings Ineffec-
magnetic responses
tive
stimulus
Response
made
effective
Effect of
rise of
exhibiting periodic
as in conducting nerve
...
vv^ire,
nerve-and-muscle preparation
587
CHAPTER XLH
MODIFICATION OF RESPONSE UNDER CYCLIC MOLECULAR
VARIATION
Explicable
Continuous
Anomalies of response
molecular changes
tonic conditions
istic
Two methods
of inquiry,
firsl
by means of character-
Abnormal
Abnormal trans-
re-
liarities
descent
All these
of drugs
Response
in-
Elucidation
615
history
CHAPTER XUII
CERTAIN PSYCHO-PHYSIOLOGICAL PHENOMENA
BASIS OF SENSATION
Indications of stimulatory changes in nerve
i,
Electrical
THE
2,
PHYSICAL
Mechanical
CONTENTS
sensory nerves similar
xxvii
Two kinds of nervous impulse, and their character Different manifestations of the same nervous impulse determined
by nature of indicator Electrical, motile, and sensory responses, and
mutual relations The brain as a perceiving apparatus Webertones of sensation
istics
their
Law Elimination
Fechner's
particular relation
644
CHAPTER XLIV
DISSOCIATION OF COMPLEX SENSATION
Conversion of pleasurable into painful sensation, and vice versa, by electrotonus The Sensimeter Mechanical stimulation
Stimulation by ther-
666
CHAPTER XLV
MEMORY
677
CHAPTER XLVI
REVIEW OF RESPONSE OF ISOTROPIC ORGANS
687
700
CHAPTER XLVII
REVIEW OF RESPONSE OF ANISOTROPIC ORGANS
CHAPTER XLVHI
REVIEW OF RESPONSE OF NERVE AND RELATED PSYCHOLOGICAL
PHENOMENA
CLASSIFIED LIST OF EXPERIMENTS
INDEX
7l8
735
747
ILLUSTRATIONS
.....
.........
FIG.
PAGE
1.
2.
Response of Indiarubber
Response of Selenium to the Stimulus of Light
Negative Response of Galena to Hertzian Radiation
Positive Response of Ag' to Electric Radiation
Electric Response in Metals
Uniform Electric Response in Tin
Fatigue in the Electric Response of Metals
Stimulating Action of NaaCO^ on Electric Response of Platinum
Abolition of Response in Metal by Oxalic Acid
Response by Method of Relative Depression
Arrangement for observing Simultaneous Mechanical and Electrical
Responses
17
13.
19
14.
Photographic Record
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
....
....
...
.......
......
.
"^
.....
.
Negativity
of
of
Pulvinus
of
17.
18.
....
.
21.
22.
23.
24.
Additive Effect
25.
28.
Response Recorder
Photographic Record of Uniform Responses (Radish)
Stimulation by Thermal Shocks
Photographic Record of Uniform Response in Petiole of Fern to
29.
27.
of Block
transmitted excitation
26
31
32
33
.........
1:2:3:4
25
26
-27
.30
in leafstalk of turnip
24
26.
20
16.
20.
Response by Galvanometric
Mimosa, when leaf is physically
19.
Electrical
34
34
36
38
39
39
XXX
COMPARATIVE ELECTRO-PHYSIOLOGY
FIG.
............
30.
31.
Responses
32.
33.
34.
Fusion of
35.
stalk)
Stalk of Cauliflower
of Plants (Style of
36.
37.
38.
39.
Efi"ect
41.
42.
Datura
44.
45.
46.
48.
49.
50.
51.
52.
alba)
54.
41
'43
....
...
45
47
47
49
52
'55
............
......
43
-Si
moderate stimulation
Response Curve from Rheotomic Observation in Stem of Amaranth
under strong stimulation
Artificial Hydraulic Response of il/zw^^a
Experimental Arrangement for obtaining Records on Smoked Drum
of Responses given to Direct and Indirect Stimulation, by Leaf of
56
57
58
60
The Abnormal
53.
43
47.
40
Specimens of
Mimosa
43.
40.
Two
PAGE
Cauliflower
....
79
82
56.
84
91
Bryophyllum
...
.
57.
58.
59.
60.
Fatigue in Celery
61.
62.
is
91
92
92
not
......
73
74
64
66
55.
61
62
93
93
93
94
ILLUSTRATIONS
XXXI
PAGE
FIG.
63.
64.
Rapid Fatigue under Continuous Stimulation in (a) Muscle (d) Leafstalk of Celery (Electrical Response)
Photographic Records of Normal Mechanical Response of Mimosa to
Single Stimulus (upper figure), and to Continuous Stimulation
65.
94
;
..........
...........
96
(lower figure)
66.
67.
68.
Alternate Fatigue
Radiation
flower;
Multiple
in
{b)
Biophyttan
Biophytum
(c)
and
97
98
Electrical
in
(a)
Responses
Electric
Peduncle
of
of
in Multiple
{d) in
69.
70.
Periodic
71
72.
73.
74.
Preliminary Staircase,
75.
76.
77.
Differential Contractile
78.
oi
Desmodium gyrans
Fatigue
Pulsation
in
of
(Pembrey
and
.
tion in Contractile
in
Responses of
the
103
103
Temporary Stop
Response of
....
Artificial Strip
79.
80.
81.
The Thermal
Variator
83.
84.
Effect of Variation of
Petiole of J/wi^a
82.
104
109
to
87.
88.
90.
Convex Side
92.
115
118
'Pulvinus of
Mimosa
..127
136
-131
Responses of Leaf-stalk of Cauliflower ,
Photographic Record showing Action of Formalin (Radish)
132
Abolition of Response at both A and b Ends by th^ Action of
.
91.
iii
89.
-^13
86.
108
.110
85.
loi
102
followed by Fatigue,
Muscle (Brodie)
99
.100
98
-99
Heart
Frog's
Phillips)
96
NaOH
134
COMPARATIVE ELECTRO-PHYSIOLOGY
XXXll
PAGE
FIG.
93.
94.
95.
Sugar
Photographic Record showing Continuous Action of 2 per cent.
96.
97.
98.
Response by strong
Na^COg Solution
Abolition of
KOH
135
HCl Acid
99.
per cent.
KHO
KHO
.
138
138
phase-diflFerence
103.
104.
105.
106.
107.
Experimental Arrangement
108.
109.
no.
Electrical
111.
Record of Responses
112.
100.
1
01.
102.
Isolated
.......
.
Diametric Method
for
150
151
152
154
155
with
Plant-cylinder
in
.
Lamina
Ends
Opposite
.
.169
114.
115.
116.
117.
118.
Nymphcea
172
originally isotropic
175
175
171
one
alba^
Specimen
1 59
165
156
by Method
of Negative Variation
119.
147
154
Distribution
Sectioned
terminal
146
End
3.
143
144
in
1 1
139
147
Section
136
136
more
.
176
.177
ILLUSTRATIONS
XXXIU
PAGE
FIG.
20.
121.
122.
123.
1
24.
181
182
183
184
185
oi
Amaranth)
125.
(Stem
.
186
126.
127.
190
128.
195
198
Lowering of Temperature
129.
Filament of Passijlora
1
30.
...
188
131.
132.
133.
^//mr^M
203
137.
138.
139.
Photographic Record
140.
141.
Initiation of Multiple
142.
Photographic
143.
144.
145.
134.
135.
136.
...........
Biophytufti
Different
Organs
Thermal Shock
of 35
Pulsation of
Pulsation in
Z>(?j-///^<^?M/;/
Leaflet of
Restraint of Leaflet
210
Leaflet of
Desmodium
212
Record of Autonomous
Leaflet
209
?.i\
in Lateral
originally at Standstill
Desmodium
209
Single
Response
208
.210
Electrical Response to
Stomach
in P'rog's
High Temperature
of Multiple
204
Leaflet
Mechanical
..
Pulsation
214
,218
Physical
after
^
.213
in
220
COMPARATIVE ELEdTRO-PHYSIOLOGV
xxxiv
'
fAGE
FIG.
146.
147.
148.
149.
50.
Crocus
......
...
'
.,..,.,.
154.
155.
153.
22
229'
Burdon Sanderson's Fundamental Experiment on Diomea Leaf
Experiment in Sheathing Petiole of Musa
229
Positive Response of certain Leaves of Dionaa
230
Diphasic Response of Leaf of Dioncea in its prime. Positive followed
by negative
230
Positive Response of same Leaf when 'modified' by previous
stimulation
230
Parallel
'
152.
224
151.
...
233
,of
Petiole
of
Musa depending on
Transmitted Stimulus
the
Effective Intensity
233
of
238
Response of Lamina of Nymphcea alba due to Transmitted
Excitation from Petiole
.
246
Diagrammatic Representation by Du Bois-Reymond for Explanation.
of Electrical Response in Organ of 7b?;^^(/(p
.
247
Photographic Records of Responses given by Leaf of Coleus aromaticus,
when both Surfaces are Excited Simultaneously by Thermal
Shock
248
Experimental Arrangement for Rheotomic Observations
252
Records of Two Successive Responses in Leaf of Bryopkyllum
calycinum under Equi-alternating Electrical Shocks
.253
Response- curve from Rheotomic Observation on L.to.i oi Nymphaa
.
Electrical
156.
157.
58.
159.
160.
161.
254
by Leaf of Plerospermum suberifolium to
Stimulus of Equi-alternating Electrical Shocks
255
Photographic Record of Responses of Carpel of Z)z7/^wm zW/<ra
256
Photographic Record of Normal Responses given by Pitcher of
Nepenthe, under Equi-alternating Electric Shocks
256
Responsive Currents in Lead Wire
264
Flat Strip of Lead, of which lower Surface is Brominated
265
Photographic Records of After-effect of Homodromous f and Heterodromous | Induction-shocks in prepared Strip of Lead
266
Photographic Record of Responses to Equi-alternating Electric
Shocks in Prepared Lead Strip
267
Response of Pulvinus of Mimosa io Equi-alternating Electric
268
Shocks
Experimental Arrangement for Determination of Excitatory Aftereffect of Equi-alternating Electrical Shocks
276
280
Method of Direct Effect of Excitation by Equi-alternating Shocks
281
Excitation by Equi-alternating Shocks
Photographic Record of Response of Petiole of Musa \o Equi-aliernating Electric Shocks, before and after Application of Chloroform
284
alba
163.
164.
.......
.
165.
166.
167.
168.
169.
170.
171.
173.
62.
172.
....
....
XXXV
ILLUSTRATIONS
PACE
FIG.
174.
175.
176.
177.
178.
to Equi-alternating Electric
Shocks
285
285
291
Shocks
296
Electrical
179.
1
80.
181.
301
301
183.
184.
185.
...
P.M. to 6 a.m.
305
187.
Isolated Responses of
188.
189.
190.
191.
192.
193.
194.
Photographic Rec^" d
Electrical
Shocks
Tomato
of Skin of
Drum
._.
Armpit
of- Electrical
.
...
A and
198.
B,
201
315
319
320
321-
Surfaces
-.
324-
Response in
-.
Pulvinus of yf/Zwoja due to Fatigue
.Photographic Record showing Reversal of Response in Carpel of
-;
Z)z7/^>a zVz^zVa, Under Sub-minimal Stimulation
Pitcher of iVd?/>^;zM^, with lid removed
.
Glandular Surface of a Portion of the Living Membrane 0^the Pitcher
...... ..
oi Nepenthe
^
.
.-
-.^
200.
313
Characteristic Curves
199.,
311
Two
310
Human
197.
309
;'
Responses of Intact
.
196.
'
i^^
Photographic Record of Electrical Response of Intact Human Lip
195.
308
Responses in Skin of
.
307
Shock
to Equi-alternating
...
306
Gecko
Single Response
302
303
186.
Equi-alternatin
302
299
300
182.
'
'
-.
326
328
335'
336^
COMPARATIVE ELECTRO-PHYSIOLOGY
xxxvi
PACE
FIG.
202.
203.
Photographic Record of Series of Normal Negative Responses of Glandular Surface of Nepenthe in Fresh Condition to Equi-alternating
204.
205.
206.
207.
208.
209.
210.
Photographic Record of
211.
212.
213.
214.
215.
216.
217.
Electric
Shocks
338
......
343
341
342
Mechanical Stimulation
of Colocasia
340
in
Positive Phase
to
340
341
Stomach
336
345
345
346
347
in
Wood
353
354
362
to
..........
The Shoshungraph
219.
220.
221.
Effect of Strong
KNO3
222.
Effect of Strong
NaCl
223.
224.
225.
Sub-terminal
226.
227.
228.
229.
230.
ture
............
.....
Mode
Solution
Solution
of Application of Stimulus
.....
.....
.....
Suction
Stimulus
372
375
376
377
effect of
363
369
378
380
380
383
383
384
385
386
ILLUSTRATIONS
XXXVU
FIG.
231.
232.
233.
Experimental
lation
388
Arrangement
Detection of
for
Change
Electrical
234.
235.
236.
237.
.........
.
238.
PACE
386
402
239.
240.
406
241.
407
242.
408
243.
244.
245.
Diagrammatic Representation of
246.
-405
in
Type
Effect in
After
flower
.........
I.
Type
Phasic
After-effect
.
412
............
Type
III.,
247.
248.
Experimental Arrangement for Determination of Differential Excitability of Optic Nerve and Cornea
.
.
249.
251.
252.
253.
III.
to Equi-alternating Electric
Experimental Arrangement
responses
positivity
.
417
417
for
.......
ment of maximum
413
413
Shocks
...
Demonstration of Differential Excitability as between Retina and Optic Nerve
Series of Photographic Records of Excitatory Responses in Frog's
Retina to Equi-alternating Electric Shocks
Photographic Record of Multiple Response of Retina of Frog under
Continuous Action of Light
Response of petiole of Bryophyllum. Light was cut off on attain-
Eye
250.
411
and After
Alternations,
III
Effects in
409
second of
the
in
.
419
426
multiple
the
.
418
427
XXXVlll
COMPARATIVE ELECTRO-PHYSIOLOGY
PAGE
254.
255.
256.
first
257.
258.
259.
260.
261.
262.
-.
427
oscillation
....
263.
0//^?t7<r^j>/^a/<j fish
...
.
427
428
428
429
...
430
432
'
264.
265.
...
267.
266.
268.
269.
432
435
436
437
437
438
440
270..
27.1.
272.
Diagrammatic Representation of .Electrical Connections for Determination of Velocities of Centrifugal and Centripetal Trans-
273.
Experimental Arrangement
274.
275.
276.
...
463
277.
464
278.
Abnormal
Lily
missions
-.
after
Thermal Tetanisation
447
for
441
442
being Injured
Geotropic
to
...
....
457
Thermal
After-effect of
...
454
462
Positive
279.
v..
..
,.
-i
466
469
XxxiX
iLLCSTRATiONS
PAGE
Fro.
280.
281.
Plant-nerve
Photographic Record of Effect of
282.
283.
of Alcohol
Photographic Record of Effect of
CO2 on
Electrical
472
Response of
Plant-nerve
......
Petiole of
284.
473
Walnut
473
Ammonia on Ordinary
474
'
on Plant-nerve
Tissue of
Ammonia
474
..
285.
Nerve
Positive, in
475
286.
Photographic Record of Effect of Tetanisation in Inducing Enhancement of Normal Negative Response in Nerve of Fern
287.
288.
Normal Negative,
Nerve of Fern
Abnormal Positive Response
Diphasic into Normal Negative by the
after Tetanisation,. T, in
how
289.
........
290.
291.
292.
296.
Frog's Nerve
Record showing Enhancement of Responsivity by
Application of CaCla
Photographic Record showing Depression of Responsive Excitability
by Application of KCl
Photographic Record exhibiting Comparative Effects of NaCl and
297.
NaBr on Responsivity
.
Photographic Record of Effect of Dilute
298.
Effect of
293.
300.
301.
302.
303.
304.
305.
.........
.........
295.
299.
CUSO4 on
Photographic
294.
NagCOg
477
the
converted through
Increasing Effective Intensity of Stimulus, due to Lessening the
.
Distance between the Responding and Stimulated Points
is
476
477
482
482
484
485
4^5
4^^
487
4^7
(-5
488
(2 per cent.
Solution on Conductivity
of
....
489
491
492
492
493
494
494
307.
Plant-nerve
...
495
495
499
COMPARATIVE ELECTRO-PHYSIOLOGY
xl
F^G.
PAGE
308.
309.
310.
Experimental Arrangement
.......
504
311.
504
312.
313.
314.
315.
316.
317.
of Frog's Nerve
516
Photographic Record showing Abolition of Mechanical Response of
Frog's Nerve by Action of Solution of Morphia
.516
Arm
for
of the Balance
for
for
De.
505
.510
in
511
Obtaining Trans-
512
318.
50x3
Electric Tetanisation
319.
517
Photographic Record showing Effect of Chloroform on Mechanical
Response of Frog's Nerve
.518
320.
Photographic
321.
.........
322.
323.
324.
325.
326.
327.
328.
Resistivity Variation
version at 6o-8
Pistil of
Hibiscus.
524
528
529
530
539
544
546
329.
330.
Mechanical Response
inversion at 59*6
522
inversion at 59 '6
521
546
in
Filament of Passi/lora,
Critical point of
546
ILLUSTRATIONS
xli
548
333.
554
334.
335.
336.
337.
338.
339.
341.
Diagram
Diagram
Diagram
342.
Diagram
343.
Experiment with Petiole of Fern demonstrating Variation of Conductivity by Polarising Current, Excitation travelling electrically
Downhill
Experiment with Petiole of Fern demonstrating Variation of Conductivity by Polarising Current, Excitation travelling electrically
331.
332.
..........
Effect of
550
Papaya
from
340.
-6 to
-4
557
Stem of
.......
.......
560
Volts
561
Hermann's
563
^63
565
Uphill
565
Experiment, when
345.
346.
347.
348.
to Anodic Region
Experimental Arrangement to Exhibit the Enhancement of Excitability at Anode, when the Acting E.M.F. is feeble
last
Intensity of Polarising
566
E.M.F
..........
.
349.
350.
351.
352.
Experimental
354.
567
568
569
at
569
570
of
.
572
353.
562
562
under
Polarisation-increment
344.
558
560
COMPARATIVE ELECTRO-PHYSIOLOGY
xlii
FIG.
355.
Experimental
Arrangements
for
Showing
so-called
Polarisation-
is
.....
Reversed
356.
357.
359.
360.
361.
358.
in
iiidica
......
.
364.
365.
366.
Molecular Model
367.
Method
368.
Repetition
370.
....
.
duction Current
372.
373.
374.
375.
376.
377.
378.
379.
381.
383.
384.
'
604
607
608
6il
......
'
602
'603
609
610
382.
595
380.
595
.601
Photographic Record
371.
594
594
594
in Sluggish
.
369.
to
589
598
600
of Resistivity Variation
Powder
579
582
363.
575
575
Human
Hand
362.
574
612
620
621
622
623
625
627
'
628
ILLUSTRATIONS
xliii
FIG.
I'
385.
386.
388.
389.
390.
391.
392.
Platinum
387.
.......
............
..........
..........
enhanced
after Tetanisation
Normal
ation
393.
Series showing
how
after Tetanisation in
629
629
630
630
632
after Tetanis-
Tetanisation
Normal Response
in the A phase
AGE
in
Abnormal
632
633
into
Tetanisation enhances
Normal Response
633
in the
B Phase
634
394.
395.
ment of Response
after
635
.......
638
397.
398.
642
399.
Response of Contraction
Record of Response in Nerve of Gecko showing the Effect of Arith-
649
400.
401.
402.
403.
404.
Photographic
405.
The Sensimeter
660
670
406.
683
396.
Series
...........
..........
........
sponses
.......
........
...........
to Stimuli I, 2, 3,
I, 2, 3, 4, 5, 6,
Stimulus
Record
of Magnetic
Responses
.
in Steel
.
642
657
12, increasing
in Arithmetical Progression
Stimuli
639
658
659
659
to Arith.
COMPARATIVE EI.ECTRO-PHYSIOLOGY
CHAPTER
negative
Method of
variation Sign
modifications
Response
resistivity
Permeability
variation
(a)
of response changed
variation
Variation
positive
variation
under
molecular
different
of
of
(d)
electrical
re-
sistance
in
tissues
organic responses
Method
called.
their
tissues,
we
found
is
find
in
one of
the
fact
On
stimulus.
tissue returns to
original condition.
its
to that state.
is
The
excitatory condition
by change of form,
is
sometimes shown
as in the case
__
^HHIIIIRII^^^H
muscle
(fig.
This might
i).
be
^HI|IH|ylBHl^^^H
^^^^^^^^^^^^^*
Fig.
I.
Series of Contractile
Responses
in
Muscle
COMPARATIVE ELECTRO-PHYSIOLOGY
changes, and
example,
may
induce a mole-
may
turn
induce,
in the
in
its
a variation
say,
Bichromated
becomes less permeable under the action of light.
stance to liquid.
gelatine
Fig.
2.
Response of India-rubber
at
The
solubility of a
may
soluble
carbon disulphide,
in
substance
is
example,
for
rendered
usually
insoluble
under
with accuracy,
we should be
stimulus
The two
effects
which have
of accurate rneasurement.
But
in the detection of
molecular
changes by
electrical
may
be used
that of
Resistivity
The impact
now
induces in
it
molecular
return
to
its
original
conductivity.
Thus
is
in
the case of
increased, or the
to light, given
In
on a
fig.
3
3 is
series of
Fig.
for
Response of Selenium
3.
(Resistivity variation
method)
of decreased resistance,
we may
arbitrarily distinguish as
negative.
by a mass of metallic
when acted upon by
also given
particles
electric radiation.
In
4 are
fig.
negative
There
J.
on
the
other
substances
which
are,
hand, some
-.'
ii
i_
is
to
increased,
is
that
Fig.
4.
(Resistivity variation
method)
COMPARATIVE ELECTRO-PHYSIOLOGY
of response does
not depend
negativity of the
substance
highly electro-positive
on the electro-positivity or
seen in the fact that while
is
potassium gives
positive
response,
molecular conditions,
the
will give
different
Again, while
positive,
tive,
By
^''''^'
E^t^ciSF^iior^^^''''
employment of the
method of resistivity
the
^^'"^
variation,
also.
The
electric
response,
however, employed
to
obtain
This electric
reaction
contradistinction
to
has been
physical.
regarded as
But
have
vitalistic
shown
in
that
inorganic.
If,
then,
we
we take an
Living.
cf.
and
the
Non-
wire,
it
if its
obvious that
is
uniform.
Hence
its
physical
properties will
condition
electrical
its
likewise
also
will
be
be the
There
will,
therefore,
made
different
from that of
be an electrical difference,
Fig.
{a)
Method
of block
Balancing
6.
{d)
;
effect, R,
Electric
Response in Metals
same
I
wire.
shall
now
describe
the
method by which
may
may
the
be employed
method of
For
first,
two
this purpose,
the
method of
electrical
be obtained from
different
methods
relative depression.
the
(fig.
6).
being
B,
made with a
by means of two
We may
now produce
COMPARATIVE ELECTRO-PHYSIOLOGY
excitation of the
A end
Using this
method of experiment, I have obtained with different substances two different types of response namely, positive
and negative. In the positive, the responsive current flows
through the wire from the unexcited to the excited, or
towards
the
excited
that
is
to
the
say,
excited
point
to uniform stimulus.
response,
The
intensity of the
is
platinum
^Ixic Response'i^
Ti^'
is
dying of the
The
note.
more excited
We
the
is
The
from the
direction of
less excited
A.
when a substance
is
ordinary
silver
negative response.
said,
positive,
stimulation, has
continued
Thus, as already
gives
its
response re-converted, as
In the
it
we
normal
of, say, tin, or platinum, becoming converted by molecular modification into negative, to be again reconverted under continuous stimulation to the normal positive.
There are, again, certain other substances, of which the
normal response is negative. Thus a wire of brominated
lead, for instance, when suitably prepared, is found to give an
electro-motive response in which the current flows from the
also,
find the
positive response
away from
the
in the case
of animal tissues.
say, the
becomes
negative.
excited
that
is
point
galvanometrically
Other
tissues,
The
under
molecular
lation.
The
electrical responses
Fig.
8.
Response of Metals
to an increase or decrease
which is paralleled by the same phenomenon in the response
of animal tissues under similar circumstances.
That is to
COMPARATIVE ELECTRO-PHYSIOLOGY
8
say, fatigue
is
as of the organic
As
(fig. 8).
in
depressors and
still
a third class
such
By
(fig.
as oxalic acid
(fig.
method of
If
as
10).
at a second
others as
9),
we
facts,
that
is
arrive
to say, the
UUi
Before
Fig.
Stimulating Action of
9.
Records to the
left
After
NagCOg on
is
stimulation
the
there will
be no resultant galvanometric
be opposed, and
effect.
This was
altogether
the
abolishing
the
counteracting
excitability of that
If
now
excitatory
A being
point,
left
effect
as,
in its
at
say,
by
by the
B
normal con-
now found
is
to flow in a
to say,
that
it
of
variation
by the
(fig.
it
The method
of
so-called
variation,
which
is
ii).
response
negative
generally
employed
in
sponsive
phenomena
animal
is
causes a negative
studying rein
tissues, is in reality,
Before
^^^^^
Abolition of Response in
Fig. io.
metal by Oxalic Acid
in the
course of this
change
sion
may
re-
responsive
find expres-
different
ways.
expression
may,
in
This
The
Fig.
1 1.
variation
vanometric
Or
the
of gal-
f represents current of
| represents
negativity.
opposite
change, expressed
mechanically as
rest
must be borne
in
mind
positivity.
ex-
But
COMPARATIVE ELECTRO-PHYSIOLOGY
lO
as to
electrical.
of stimulus.
Again, those various responsive phenomena and their modifications which are the subject of our inquiry, are such as are
induced by different external agencies. Under the influence
of certain conditions, the responses of living matter undergo
an abolition
we
In
death.
And
transformation.
formation
some molecular
when
it
'
In
my own
into
'
physico-chemical processes.
physiological,' therefore,
it
will
tissues,
but as
in
no
'
We
shall,
the
in
following chapters,
study excitatory
phenomena
will
vegetable tissues,
and
it
will
in the case
is
of
no
animal tissues, which has not its exact correspondence in the vegetable. Those anomalies, further, which
have been observed in the response of the animal, will be seen
types of
II
the
simplest
form
of
CHAPTER
II
Difficulties
of
investigation
Electrical
response
of
pulvinus
of
'
'
'
into
two
distinct
plants, as regards
classes
*
:
ordinary
their
'
and
such plants
Hence the
'
sensitive.'
excitatory electro-motive
phenomena
in
plants.
They
ii.
pp. I-31.
cf.
On
leaf
stimulation
of the
phenomena
these electrical
to be consequent
on hydrostatic
By means
of this
'
The
phenomena
in
vegetable organs.
of water.
The
greater
one
absorptiveness of
ever,
different points
process,
is
We
which
shall
this
'
(Chapter X.).
induced
by
as, say,
study later
current of
rest,*
so
called,
It
has
is
actually
on
depends
already been
stated
that
to see,
first
negative swing
observed might be due to the disturbance of the diffusionprocess through alterations of the protoplasm.
COMPARATIVE ELECTRO-PHYSIOLOGY
14
Munk
effects
which he observed
in
electrical
maximum
in
changes being
one
initiated in
the other.
it
'
the other.
become
positive
'
The
excitation
the
mode
The
direction
such as
become
to
indicate
that
in
excitation
excited
of
is
cells
become negative
to unexcited.'
of leaves
'
by Burdon Sanderson
know which
difficult to
the results of
what
and which
as
cause.
It will
theories
some other
advanced by different observers in this field are someThis must have been due
to the difficulties
met with
in
be found
was
difficulties
of
apparently
But,
rest.
opposite
are
negative,
For an
the
(i)
fact
it.
that
Chief
among
and
positive
these
negative
variations
current
combination with
in
as
matter
responsive
not always
of
fact,
these
positive
variations,
two
and
added algebraically
to
There was
was due, as
shall
movement of
Under
by mechanical
it may be
one or the other of these which becomes prominent.
hope to show that each of them is definite and dis-
water.
different conditions,
either the
I
shall
With regard
it
may
strated
in
responsive
sensitive
the
course
phenomena
plants
of the
as
may be
all
plant organs.
work
observed
to be
is
demon-
that
in ordinary plants,
I
shall
such
in the case
first fact
present
of
occur
and are
be able to show,
of these phenomena,
much
COMPARATIVE ELECTRO-PHYSIOLOGY
than
simpler
available.
theory of
the
electro-motive
molecules,
is
It will
which
shown
for
the
not only
same, but
it
electrical
the
is
is
all
those conditions
And,
no breach of continuity
responses in plant and animal,
state
between
response,
electrical
by
to be modified
physiological
the
affect
lastly, it will
as
opposite.
fact
again, will be
is
response
of
sign
of the tissue.
both
in
cases
the
in
is
of the
that
plant also.
In
characteristic of excitation,
of the
reaction.
first
electrical
select for
Mimosa^ because
v^^e
the
is
response
experiment a
leaf,
we have
It is
by
stimulus,
traction.
it
This
responds
in
'
is
phenomenon
its
negativity.
electrical
it
and
its
is
'
to this
in living tissues,
we should henceforth
it
is
distinguish
of
excitation,
a contraction
of,
Under
of the
leaf.
It is
excitable.
This, however, is an error, for
both upper and lower halves are excitable, and contract
under stimulation. If localised stimulus be applied on the
of the pulvinus
upper
is
is
erected.
yet
the
of
excitability
due to the
lower
causes
resultant
two
fall
is
sides of the
of a contraction
FiG. 12.
generally
is,
therefore
Hence the
The
organ.
the
stimulation
17
Arrangement
for
Electrical Responses
the
and
fall
of
leaf.
we make
suitable
is
made with
electrical
connections
a galvanometer.
One
is
way we can
In
of
to be
by
distal point.
is
spot of light,
thrown on the
C
COMPARATIVE ELECTRO-PHYSIOLOGY
recording drum.
by
sudden
its
This spot of
shows,
change.
To show
trical
exact
moment
arrangement
is
an Optical Lever.
from
of
this
mirror moves
galvanometer
the
in
horizontally or
two
by means of
laterally.
necessary to
is
it
spot of light
that
For the
have the
one direction.
in
The
reflection
of
is
inclined.
Stimulation
may
be effected
the
in
neighbourhood of
two responses
that the
mechanical
and
being
practically
As
variation.
excited point
that
is
with
is
take
of the leaf
induced electrical
the
fall
excited vegetable,
tissues.
coincident
electrical
is
the
same
Biophytum^ whose
(fig. 13),
obtained from
It
The
by up-curves.
tivity
metric positivity
will,
erectile
A few words may be said here as regards the synchronism between the two forms of response. The excitatory
molecular change takes place instantaneously, and the
electro-motive variation
concomitant with
it.
is,
This
shown
is
in
by the
Chapter IV.
rheotomic
It will
there
Fig. 13.
by the
tissue
(e)
Responses in Biophylum
same moment.
is
practically
trical variation
no latent period.
But
if
the
same
elecwill
be a lag in the response, owing to the inertia of the galvanometer needle. Similarly, in the mechanical response,
is
is
when the
tissue
is
But
this
may
in a state of
be
still
depressed
COMPARATIVE ELECTRO-PHYSIOLOGY
20
excitability.
mental
It will
excitatory
when
instantaneous,
is
may
in
the funda-
outward
its
nevertheless
consequence of the
it
inertia
should further
the
restrained,
response
electrical
takes
unimpeded.
I
shall
here relate an experiment
place
leaf
of
Mimosa
re-
somewhat
taking from
Photographic record of ElecResponse by Galvanometric
Negativity of Pulvinus of Mimosa,
when leaf is physically restrained from
falling.
The first series in response to
given uniform stimuli ; the second
Fig.
prolonged,
eighteen
five to
minutes, according
to
the
14.
trical
In order to obtain
season.
within
recoveries,
reasonable time,
find that
it
is
necessary to apply
moderate stimulus. There is then a moderate fall, without
complete collapse, and recovery under such circumstances is
found to take place within a minute or so. In order to show
prevented from
petiole in a clamp,
seen in
in
fig.
14.
when the
The
first
leaf
held
is
the
were taken
and the
intensity,
We may
One
by increased
increased
is
here see
intensity
21
is
how
of stimulus.
the trend of
of the pulvinus.
And we
same way
found to be
is
is
accompanied
by galvanometric
negativity.
fallacious in
possible only
contractile,
extreme.
the
and there
is
is
Thus, when
tive' Mimosa would appear to be insensitive.
we place a cut branch of Mimosa in water, the pulvini of the
leaves,
On now
fall
is
found to
take place
is
will often
dark room.
is
this
of the
It will
then be
Mimosa
cumstances, which
may
And
in
the plant
fail
to
movement under
these cir-
COMPARATIVE ELECTRO-PHYSIOLOGY
22
much
varies so
that
its
by the unequal
is
it
excitabilities
lower halves of the pulvinus that that differential contraction is induced which brings about the lateral response of
Mimosa
the
leaf.
Now
it
is
clear
from
this
that
if
the
be reduced or abolished, by
any means whatsoever, there will then be a corresponding
We
and
Mimosa, from
in
induced
We
may
is
in
consequence of
whatever
state of turgor,
in the relative
pulvinus.
And
this, to
certainly,
will
Mimosa
day.
Thus,
Thus
movements of the belauded sensitive plants,
Mimosa, on which depended the arbitrary assumption
the mechanical
such as
itself
extremely unreliable.
identical plant
Mimosa
is
we have
a criterion by
which to
any motile
On
indication.
applying
independently of
this test,
have found
and
from
organs of
all
this
in spite
was
all
23
all
plants
And
mechanical contraction.
The common
error
of regarding
on
all sides.
Hence those
movements, dependent on
lateral
which are seen so conspicuously disMimosa, cannot take place here. But that the
organ as a whole undergoes a responsive contraction has
been demonstrated by recording the consequent induced
Such longitudinal contraction is
shortening of its length.
considerable
for instance, in the filamentous
sometimes very
corona of Passiflora it may sometimes be as much as 20 per
differential contraction,
played
in
that
all
In
of
shall
electrical
excitation, since
applied at a
reach
it.
It
COMrARATIVE ELECTRO-PHYSIOLOGY
24
we take
making
or stem, and
petiole
suitable electrical
connections
(fig,
15),
response
is
same
the
again,
it is
'
like that of
animal
plants,
'
ordinary
and that
'
is
of
in both,
tissues.
induce an
identical
may
excitatory
response of galvanometric
Any
negativity.
Fig. 15.
Method
of Transmitted Stimulation
of temperature
or compression
prick or cut
acid
any variation of
first,
any
electrical currents
stimulatory action of
all
these
The
demonstrated in my work on Plant Response,' the excitation induced being there shown to find expression in
*
shall deal
more
The
1906.
will
be taken
in special chapters
here
and of gravity,
currents, of light
electrical
up
25
shall
tatory effects
of stimulus enumerated.
We
responsive
from
heat,
The
various
forms of
mechanical stimulation
may now
be
effects
of
subjected
and
demonstration,
to
we have
first
to observe the
from
sudden tension. The specimen is
clamped securely in the middle
effect of the stimulus resulting
so that
(fig. 16),
is
given
the
to
half alone
when a vertical
is
upper
Fig. 16.
that
When
Plant securely clamped.
suddenly pulled, tension induces galvanometric negativity of A.
Tension
half,
subjected to a sud-
unaffected.
entirely
left
Excitation by Sudden
pull
Under
A becoming galvanometrically
to mechanical compression,
is
an
negative.
and
electrical response,
for this
is
next subjected
electrical
The
halves of a cork.
is
A may now
This gives
rise to
the
same
electrical
response as before.
This
fact, that
may
electric
men
connection at
(fig.
down,
this
subjected
17).
When
A with
by
first
making an
the tissue at
is
to
COMPARATIVE ELECTRO-PHYSIOLOGY
26
The
sion.
result
negative
similar
is
elec-
trical response.
Excitation
may,
by
means of a sudden blow
again, be produced
This blow
at a point.
may be
Fig. 17.
by
means of a spring-tapper
and Compression
E is connected with the upper point A
a sudden bending down causes tension of
A.
When connection is made with A' the
same flexure causes compression of a'.
Both induce galvanometric negativity.
the
lifter
at
downwards,
is
T.
projecting
provided with
It is
length, projecting
regulated.
the height or
this
As one
be varied.
-.
and
its
passes through S R.
a screw-thread, by
s is
head
rod
which
18), in
(fig.
When
delivered
may
of the
and
delivers
The
the stroke,
Fig. 18.
tap.
the
is
lift,
and
intensity of
measured by
graduated
shown
and
sharp
height of the
therefore
released,
scale
not
in the figure.
We
first,
by augmenting the
by a
give
superpose a series
isolated
single
taps,
or
is
sliding catch.
We may
in
rapid
may
Stimulation, again,
or pin in
may
pricks
Or
It
of
Successive
may
the specimen
is
torsional vibration
at
27
The
one end.
A makes
is
applied
(a)
stimulation
that end
gal-
direction
outside
the
towards, and
away
the
current
being
circuit
the tissue
in
Vibration of
from, A.
tivity
current
now
of
(fig.
it
will
is,
Fig.
is
19.
Torsional Vibrator
The plant
A and B.
(b)
be
ap-
is
clamped
at c,
between
This method
when we wish
excitation, the
The
(a)
In the cases
described,
plied directly.
B is stimulated^
the
response being
reversed.
just
19),
to study the
good conductors of
here,
therefore, inapplicable.
given
agent sulphuric
or hydrochloric acid
is
applied at
X at a short distance from the proximal contact. The transmitted excitation is now again demonstrated by the induced
galvanometric negativity of that contact. It will thus be seen
that, whatever be the effective form of stimulus employed,
it gives rise to a definite and invariable electrical response
whose sign is always one of galvanometric negativity.
It was shown, then, in the course of this chapter that
the excitatory change in sensitive plants is characterised
by contraction, negative turgidity variation, mechanical
*
depression
of the
leaf,
'
and by the
electrical'
response of
COMPARATIVE ELECTRO-PHYSIOLOGY
28
galvanometric negativity,
all
the leaf
is
physically restrained.
The same
is
electrical response of
galvanometric negativity
plants.
And
'
ordinary
it
was
and
electrical
induce
the
same excitatory
CHAPTER
111
Method
of
response
Torsional
block Effective
intensity
vibration
as
a form of
of stimulus dependent on
Uniform
electric responses
method of confining
excitation to
one contact
Tetanus.
in
the last
tissues, in
of stimulus.
it
will
it
is
thus a sign or
be secured,
is
Assuming
physiological modifications
induced by various agents will be manifested by a corresponding modification of response. The conditions essential
to such application of stimulus are, then, (i) that it should be
capable of uniform repetition
by
(2) that
it
should be capable
of increase or decrease
the
electro-physiological
difficult to
meet.
definite
investigation
Chemical stimulation,
depends, are
for
very
example, cannot
COMPARATIVE ELECTRO-PHYSIOLOGY
30
galvanometric
electrical,
results,
But
it is
Indeed,
disturbance.
obvious that
if
we
response
the
as
is
almost a necessity.
is
The
fidence.
it
struck
subjected to
is
is
true,
thus undergoes an
I
stimulation caused
repeated,
unknown
variation.
of stimulation, in
which
these
all
The Vibratory
Fig. 20.
difficulties
modes
have been
Stimulator
Plant P
is
stops, s
s'.
successfully overcome,
possible.
These are
torsional
it
must be
condition to be
effect
fulfilled.
If
and
(2)
the
vibration,
we wish
to
is
still
obtain
another
the pure
unknown
This may,
true,
it is
effects
relative depression or
method of negative
variation, so called,
am
about to describe,
in
interposed
According to
clamped in
either end is practically precluded from affecting the other.
Stimulation is brought about by means of torsional
The stem or petiole is fixed, at its middle, in a
vibration.
vice, V, the free ends being held in tubes, c c', each provided
Fig. 21.
Complete Apparatus
for
Method
Stimulation
torsional vibration
may now
vibrational
mental investigations
may
be carried out,
is
given in
fig.
21.
COMPARATIVE ELECTRO-PHYSIOLOGY
32
and
B.
there
is
E,
in
make
effects
an
R,
heating
electrical
coil,
of temperature,
For the study of the effects of different gases, there are inlet
and outlet tubes, which enable a stream of the required gas
or vapour to be circulated through the chamber.
motive variation takes place, which afterwards subsides gradually. If next the
torsioned end be suddenly brought
back
Oy
motive response
first.
*"
tive,
Influence of
Suddenness on the
Efficiency of Stimu"^
responses' to
are
of the
vibrations
30.
ampntude,
In a the
in the case
of a to-and-fro
ci
Fig 22
same
Hence,
is
vi-
returned to
its
be equally
is
effective,
may
rapid in
^^"^^
i^"
stimulation to plants,
to
is
dis-
depend
vibration
constant,
conditions.
First, the
This
is
amplitude
done by means
33
must be uniform.
This last condition is
by an arrangement shown in fig. 23. The torsionkept tense by means of a stretched spiral spring, s,
vibration period
effected
head
is
made
of
From
steel.
this
B,
is
by the
S.
Fig. 23.
B,
acting
torsion-head
there
projects an
made
is
thus
The amplitude
is
at the
The arrangements
But
to
Q.
for certain
second
striker,
may
may
it is still
effect of
in con-
be excited simultaneously.
By way
of avoiding
it is
now
But
this
this, therefore,
by the additive
is
found
COMPARATIVE ELECTRO-PHYSIOLOGY
34
to
become
seen in
effective
fig.
24,
large response
rapidity-
substitute for
mated
Additive
Fig. 24.
Effect
(a)
single stimulus
of 3 vibration pro-
duced
little
effect
or
no
^e uniform.
stdk of
ment of
series of responses to
is
a distance of
following
is
t-x>
^
for
a dead-beat instru^i
type.
Ihe
sensitive-
deflection of
metre.
of
mm.
at
method of obtaining
form
found to
is
,
ampere causes a
uLiixf'
the
he galvanometer used
a
D Arsonval
ness of this
large
accurate
devised
these experiments
rj,,
./.
effect,
duced
and a
records,
response-recorder.
The
^^
Fig. 25.
Response Recorder
record,
is
driven by
clockwork.
Different
speeds
of
35
it
The
galvanometer spot
is
by moving the
light,
carrier
which
will'
is
The
recovery.
the
ordinate
the
in
these
curves
represents
We
a small
time
is
The
determined
surface
recording
by the
moves,
distance
in
unit
through
time.
In
which
this
the
simple
by
wrapping a photographic film round the recording drum.
of responses taken from the
I give in fig. 26 a series
root of radish {Raphanus sativus), in which the stimuli were
It
is
applied
at
intervals of
one minute.
It
may
may
be rendered,
if
proper
P3
COMPARATIVE ELECTRO-PHYSIOLOGY
36
out, that
for
in this
electrical responses
represented by down-curves.
Fig. 26.
large
volt.
Organ
Specimen
Root
Radish
Raphanus
sativus)
Geranium {Pelargonium)
Stem
Vine
Vitis viniferd)
Amaranth [Amaranthus)
Horse Chestnut {/Esculus hippocastanutn)
Turnip {Brassica naptis)
Petiole
Eucharis
lily
{Eucharis amazonica)
Peduncle
Arum
Fruit
lily
{Ricardia africana)
high as
strong electrical
shall see in a
plant which
autumn
we
intensity
in
The same
3/
or winter.
may
Again,
and, lastly,
is
killed, the
my
in
'
Plant Response
'
that
Hence,
if
by
a tissue be surrounded
and
variation of temperature,
will
be subjected to a sudden
Now
experiments
through
flowing
duration
the
the
wire
and
if
of
intensity
maintained
be
not
in actual
to the tissue,
make
contact with
by
scorching.
it.
This
The
in
current
constant,
the
The thermal
is
successive
the
current
is
is
any injury
so adjusted as to
is
for
effect of a
may be
of temperature
is
tissue, if care
be
not so great as to
it.
The
difficulty of
individual shock
actuated by a
is
metronome
(fig. 27).
second rod
is
attached
carries a bent
COMPARATIVE ELECTRO-PHYSIOLOGY
38
the
figure, is
no thermal stimulation.
The
is
and counts,
say,
of
the
five
strokes
metronome,
after
which the press-key
is
again opened.
this
mated
effect
In
sum-
way, the
ob-
is
thermal shocks.
process
as often
This
repeated
is
as
desired,
the diagram
is
given
in fig. 27,
was
contact
important.
is
here
stimulation
confined to one
from reaching
prevented
I
is
circuit.
shall, in
the
distal
is
Hence
made with
if
this tissue,
It is true that
contact.
The arrangements,
then, being
may be
uniform stimuli.
How
in the
manner de-
made
fig.
28, in
which
is
may be
given a series
(fig.
28)
39
We
uniform stimuli.
shall
In
.LLL\^i
Photographic Record of Uniform Response in Petiole of Fern
Fig. 28.
to transmitted excitation
animal tissues
it is
and
same
is
necessary to choose
is
which exhibit
of these
first
little
the
specimens
fatigue.
In the
fall
of
the
height h
the
third
curves
(fig.
striking-lever
from the
and
from
3h.
The
29) clearly
response-
show the
Fig. 29
1:2:3:4
producing
in-
stimulus.
variations.
COMPARATIVE ELECTRO-PHYSIOLOGY
40
Angle of vibration
2-5
5
7-5
044 volt.
075
090
TOO
106
IO
12-5
'.
,,
,,
induced
with
increasing
stimulation.
is
always
diminution
of
Fig. 30.
(Cauliflower-stalk)
Vertical line
stimulus.
manner
But
The
fatigue.
this
-i
volt.
this
is
merely a secondary
following records
may be brought
(fig.
about.
31) will
effect,
show
due to
what
in
first
Every response
in
this
series
therefore,
starts,
increases
with
increasing
stimulation.
In
the
on stimulation
is
20
/U
Fig. 31.
Two
Specimens
of Stalk of Cauliflower
In
{a)
stimulus of the
recovery
series.
is
complete, in
{d) it is
incomplete.
is
only partial
is
In
if
we measure
the
tilted
upwards.
heights of successive
COMPARATIVE ELECTRO-PHYSIOLOGY
42
no allowance
made
is
Ordinarily, how-
way
its
In this
physiology as the
cardiac
muscle,
all
or none
which
intensity of stimulus
is
In the case
principle.
'
example, there
for
is
certain
of
minimal
When
a tissue
is
muscle, for
stimulus
is
disappear.
complete
added
The
to that of the
result
and these
is
first,
maximum.
attain a
With moderate
tending
to
become more
and
way
In
of the second
effect
is
incomplete,
52).
the
have obtained
mechanical
In fig. 33 is given a photographic record of tetanus, taken from the longitudinal motile
The
fig.
34 form an example.
observation of electrical
different
43
Fig. 32.
Record
ways.
In the
this
first
in
two
different
(b)
Fig. 33.
Photographic Record
of Genesis of Tetanus in
Mechanical
Response
of
Datura
alba)
Plants (Style of
When
Fig. 34.
Fusion of Effect or
Rapidly Succeeding Stimuli
(a) in
muscle
{h) in carrot.
it
position of stimuli.
CHAPTER
IV
Instantaneous mechanical
Response - curve showing general time - relations
stimulation by electro-magnetic release Arrangement of the rheotome
Tabular statement of results of rheotomic observations Rhythmic multiple
responses.
fairly
response of
electric
high sensitiveness
is
plants,
required.
One
by
stimulus.
rheotomic
mode
in
it
of investigation.
is
necessary to employ a
An
account of
this,
and
specimens.
some
cases, recovery
in duration in different
is
complete
much
in a
very
For the
purpose of forming a general idea of this difference two
response-records are given here, one of which was taken from
a stem of the quickly-reacting Amaranth (fig. 35), and the
short time
in others
it
takes very
longer.
It will
be seen that
of these
first
was
the
latter,
in
forty-
it
still
far
Thus
shown that
recovery
is
in
it
much more
be
will
nerve,
The
physiological modification
is
Fig. 35.
Response of
{h\ ct^
{b)
of
summer than
This difference
in winter.
mechanical response
as already stated,
is
period
is
it
Amaranth
is
demonstrated by
it
is
in
summer
the
as long as eighteen, or
overcome the
as already said,
devised
variation
by Bernstein.
The
relative
values
may
of
jthe
we may
by
using,
originally
electrical
impact of
COMPARATIVE ELECTRO-PHYSIOLOGY
46
galvanometric
of
contacts
equal
period
the
at
required
intervals.
The
difficulty in
this
investigation
lies
the
in
instan-
unknown
advisable
if
possible, a non-electrical
it,
responsive
variations in the
might give
effect.
electrical response to
And
form of stimulus.
rise
Hence,
it
to
is
employ,
it is
only
method, that
be overcome
is
itself,
Another obstacle
to
unknown element of
the responding
successfully
itself.
All
by employing
met
stimulation, which
am now
these difficulties
have
mechanical form
the
about to describe.
of
We
have
be imparted
we
by the
other.
The
intensity of this
against the
stop
P.
is
made
to rest
is
is
to subside.
The
vertical thread
shown
in
a soft
iron
fig.
37,
armature
moment when
is
seen to hold
at the
At
the
stimulation
is
magnet
is
interrupted
by an
be described
later.
By
'"
^'^^.iLirirr.o^
the break of the current the Torsion-head set by string against stop Q
is suddenly let go by electro-magnetic
armature is released and a
release seen in
fig.
37.
head
is
FiG. 37.
A,
General arrangement
for
P.
Suc-
Rheotomic Observation
pensator.
intensity
may
thus be applied on
may
be desired.
COMPARATIVE ELECTRO-PHYSIOLOGY
48
The next
point
an automatic arrangement
to secure
is
tissue, for
second.
are to be effected
contacts
short-lived
these
in successive
The
animal.
much more
are
tissues
sluggish than
those in
for,
two
is
seen in
fig.
striking-bars,
The
37.
A and
B,
the
therefore, are
this
revolving rheotome-disc
of which
is
fixed,
and
magnetic
the galvanometer G
is
short-circuited
by key
Kg,
and
is
it
striking against Kg, that the responsive current can act on the
galvanometer.
is
disc
is
rotated
the
The rheotome-
perfect
one second.
The
is
One
100 cm.
The breadth
also
cm. and
course of
disc,
'oi
it
second.
of the striker B
is
to the
and Kg which are adjusted along the same radius of the disc.
Kj is a balanced key, one end of which carries a two-pronged
brass fork, both prongs of which are normally dipped in cups
of mercury, thus completing the particular
By
by the
The
is
striker
tilted
striker B then
A on
electric circuit.
a projecting
rod
is
Kg.
kept
is
down by
a spring
S,
The
that
to say, in 01 second
is
(fig. 38).
cession, can
by
increased,
increasing
The key
said, controls
magnet,
the electro-
on
its
instantaneously
the
mechanical
Fig. 38.
stimulation of the
The
B.
already
which,
release,
effects
A and
K,, as
tissue.
Keys
K3
is
is
(fig. 37),
By
made
the speed
till
the break-action of
A on
Kj
first
postponed.
is
effected.
short
definite
before,
the rod B
adjust-
in relation to A, enables us to
is
of a second, at
at a distance of
increasing
cm. from A,
The
definite intervals.
When
stimulus
circuit, as stated
intervals
required
is
for
observation.
In
a second
COMPARATIVE ELECTRO-PHYSIOLOGY
50
'01
is
time.
may
There
in
be a pre-existing
contact, Ni
and
the
Ng.
of potential
difference
may
not be a source
that
The
employed.
it
in the
slider of the
compensator
is
so adjusted
specimen.
this
is
give below
two
which
tables
And
series.
summarise rheotomic
By
stood the
mean
interval
that
is
Thus,
between the
the removal of
for the
mean
interval of -02 of a
B,
whose breadth
is
on which
will
Specimen
Mean interval
after excitaton
I
second
>>
Specimen
Galvanometric
deflection
70 divisions
100
,,
'
Mean interval
after excitation
'I
second
-2
"
,,
310
220
i-o
,,
104
70
2-0
,,
15
-5
i-o
,,
2-0
6-0
II
Galvanometric
deflection
26 divisions
100
70
64
56
42
24
4
maximum
was attained
in three-tenths
of a second.
in
In
reached in two-tenths
how quickly
maximum, and how
39 shows
fig.
of
electro-motive effect
maximum was
first
There apppears
no latent period, the induction of the electromotive effect being apparently immediate. This will be
made evident by the results given in the next series.
rapid
is its
to be practically
"
100
'
8o|
60DU
40-
20-
'
^ ^^
^=;;
>
Fig. 39.
'
iT-
]i:'
3:'
4?
y:
\':
second.
abscissa, time.
;
(Petiole of cauliflovvrer.
Large
The
intensity of stimulus
I
often gives
of both mechanical
52
inertia of the
By
the
was
Galvanometric deflection
01
second
OS
15
20
63 divisions
77
82
68
76
75
104
86
67
30
40
60
70
80
It will
01
second, a
of a
110-
100-
\
^
80-
>
<
^%
\/
70-
'
'
P'iG. 40.
"
1"
i"
15,
'.
-3r"
already
^V"
been induced.
-3,
The
in
-i
Stem of
first
and the
maximum
third
"
-i
second.
occurred after
after -6 of a second.
ORS'ERVATION BY
We
responses,
there
the successive
is
rhythm
in
maxima being
these
here
53
multiple
found
to
second.
The
third of these
owing
mum
variation
this period
either of a
is
may
be
made
still
shorter
intensity of stimulus.
by the employment
tissue, or
Strong stimulation
is
of a
greater
CHAPTER V
of
responses
turgidity- variations
opposite signs,
Indirect
characteristic
excitation -
of
positive
Wave
of increased hydro-
Method of separating
metric positivity
Having now
I
shall
of response
response Separation
of physiological block.
is
characteristic of excita-
electrical
means
In-
induces galvano-
and negative
factors, in
in
the
And
tradictory results.
it
and by ascertaining the conditions under which each invariwill be arrived at in the field of
We
With
regard,
first,
that
to say, of
we may
The
cut end
test.
is
in
is
filled
fixed watertight
other
POSITIVE
55
is
connected alternately with a vacuum and a force-pump, by means of which a diminution or increase of hydro-
end
static pressure
may
be induced at
and
its
away
In this
will.
from, or force
it
way
it
is
variations
When
at will.
turgidity
is
is seen to fall.
This is what happens also
under the ordinary negative turgidity-variation induced by
excitation.
When turgidity is increased, on the other hand,
indicating leaf
the leaf
is
erected
(fig.
41).
In the
Fig. 41.
Artificial
Hydraulic Response
whole
of
loses water.
Mimosa
The
from
the
lower and
the
upper.
In the
case
and
in
it
is
again
the
turgidity
characteristic
electrical
presently.
It has already been mentioned that the direct application
of stimulus at a point causes a negative turgidity-variation
of that point.
shall now see whether, under any
We
COMPARATIVE ELECTRO-PHYSIOLOGY
56
stimulus will
circumstances,
If
variation.
stem of Mimosa^
on
stimulus
the
at a certain distance
stimulated.
is
a hydrostatic disturbance
clear,
And
with
of increased
hydro-mechanical disturbance
this
great
relatively
pressure, as
we have
Now
velocity.
it
pressure.
be transmitted
will
such an increase of
the
pulvinus,
which the
consequence of
in
ought to be erected.
although this hydrostatic
And
leaf
disturbance
sumed
very
transmitted
is
con-
is
in
After
hydrostatic
^y^
passage of
the
wave,
the
follows
there
Fig. 42.
given
to
and
Direct
'"'"^^''''"'
^^
i^vlf
Thermal stimulator
at s
In-
^'""^ ^
''''^'''''
'^''^
of'erectTon!''
arrangements
inductions
may
when
pulvinus,
^
the
prcvious
^
movement should
excitatory
glve
depression,
the
erectile
place
or
fall
tO
of
the leaf
In fig. 42 are seen the
an experiment on Mimosa by which these
be verified. Moderate thermal stimulus is
S,
at a certain distance
latter is attached
traces
the
When
the stimulus
by a thread
response-record
is
And
variation.
for
applied at
This
turgidity
produces this
an up-curve.
When
fall
by a negative
of the
leaf,
turgidity-
seen in
a moderate stimulus
is
fig.
43
applied
POSITIVE
57
This
followed by a responsive
is
(fig.
fall,
when the
true excitation
b).
43,
It
it is
two
discriminate
to
different effects
It
factors
and
its
We
expansion.
direct excitation
and
negative turgidity-variation
of
fall
the
Unfor-
leaf
Mechanical Responses
Fig. 43.
of Leaf of Mimosa
{a)
of stimulation which
is
applied
at a distance.
And
it
has not
as
the
true
excitatory
effect
responding
intervening
organ.
tissue
When
is
the
effect.
highly
as
see that
if
tissue
But we
shall
non-conducting or
effect
is
is
COMPARATIVE ELECTRO-PHYSIOLOGY
58
This
and expansion,
distinguish
therefore
as
the
Indirect
shall
Effect of
I
contradistinction
in
term
to
the
Indirect Stimustimulus,
lation, as it is generally used. The last-named, however,
latter effect of
shall
positive
turgidity-variation
MITTED Stimulation.
Fig. 44.
If
the
title
of
TRANS-
intervening tissue
be of
Stimulus was applied at some distance from the responding leaflet. And
the preliminary erectile twitch is due to the prior arrival of the
Thick dot represents moment of application
hydrostatic disturbance.
of stimulus.
fig.
44
is
and transmitted
effects
of the indirect
leaflet
of
Biophytum.
POSITIVE
59
is
the case
order,
methods.
separate
to
them,
we may employ
various
is
In
make
the
Or we may choose
the
quickly
a direction
Thus
example,
S^ (fig.
leaf, it
is
record
lus,
that
is
mind that a
fall
till
3 "45
It is
even after the arrival of the respective waves, before the turgidity-variation
is
The
This, as
we have
done on the
tissue
For the
is
latter causes
represents
the impulsive
tissue,
COMPARATIVE ELECTRO-PHYSIOLOGY
6o
the tissue
or
The outward
negative
respectively expansion
shall see,
but
is
is
that
is
to say,
manifestations of
these
and contraction.
The
positive, as
we
The
turgidity-variation,
Fig. 45.
fast-
In observing
positive
stimulus,
turgidity-variation,
consisted
opposite sign
to
of
that
due
to
the
indirect
effect
of
the
negative turgidity-variation,
itself in
fall,
of
POSITIVE
the
We
leaf.
between the
exists
now
shall
see
6:
and
negative turgidity-variations.
In carrying out this experiment, I took a specimen of
Biophytum and applied stimulus at a distance from the particular leaflet whose responses were to be observed, arranging,
same
at the
and
electrical responses.
It will
And
this
is
the contractile
by
its
positivity.
opposite
namely,
fall
excitation.
It will
its
Besides
galvanometric positivity.
movement of water
in
the
to
tissue
gives
rise
its electrical
this,
expression,
mere physical
the
tion
this
can
and
positivity,
be detected,
still
The
killed.
how
is
question of
to discriminate
what
of
associated
turgidity,
found
decide.
it
very
But
at
Electrical Responses in
first
ultimately
'
by bringing a plant
a
condition
just
Biophytum
difficult to
to
short
of
death, and
thus
abolishing
its
COMPARATIVE ELECTRO-PHYSIOLOGY
62
indication was
positivity.
On
found
be one of
to
considerable electro-
order
excitatory
exhibit
to
and
ordinary plants,
the
expression
electrical
hydro-positive
effects
took a petiole
one
of
of
of
the
stimulation,
cauliflower
connection,
the
in
made
and
proximal,
it,
In order
was sud-
downwards.
This
was repeated once more, and the
same effect observed.
I
next
represented
"ok
of Cauliflower
a, hydro-positive ;<5,di-phasic;
'
<r,
Static
Shortly
followed
by the
experiment on
-"^
is
seen
Miimsa,
in the record, as
This
a di-
POSITIVE
From
animal.
it
63
will
in reality due,
applied nearer
that
we
47, c)
Owing
proximal contact.
stimulation, the
(fig.
is
masks the
It
is
seen
thus
that, as
method of
to apply stimulus
is
at a point so distant
is
sufficiently
reaction completely
static
then,
that
ductor of excitation,
at
we must
effect
of
excitation
hydrostatic wave.
tion,
will
it
first
may
obvious,
electrode, in order
that
behind
the
sufficiently
be the indirect
and very
lag
Similarly, in a
It is
the
respectively.
if
be very
strong.
In
each
other,
effect
as
'
shall
in
the hydro-positive
from 'the
true
future
effect,'
excitatory
refer
to
by way of
effect,'
of
that
hydrostatic
differentiating
negative
it
turgidity-
a high
or
less
tissues
which
exhibit
this
excitation.
When,
therefore,
little
in
power of transmitting
experimenting with
this
COMPARATIVE ELECTRO-PHYSIOLOGY
64
so near as
response of galvanometric
tivity
(fig.
was
nega-
obtained
case
this
in
48).
it
given
by transmitted
excited
is
said,
induced
one
of
due to hydro-positive
and the other of negativity,
positivity,
action,
due to
true
excitation.
the stimulator
is
near
to,
When
or co-
the tissue
is
subjected
to rapidly
and the
elec-
the
Photographic Record
Fig. 48.
of Electrical Responses of
Potato-tuber
a, Positive response to stimulus
ability
near.
is
The
positivity.
resultant
electrical
response in general
is
is
the galvano-
precedes
it.
From
this,
it
is
effect,
clear
that,
as
regards
the
stimulation, there
may
POSITIVE
65
Constituent Factors
Resultant Response
Ne >Ph
Nk <Ph
Galvanometric negativity
Galvanometric positivity
Excitability great
Excitability diminished
or nearly abolished
We thus
for
may undergo
antagonistic
diminution, or even
of simplicity, assumed
the sake
act
effects
may
time-relations
may
phasic
stimulus
Some
may
Hence
be seen, giving
it
two
the
be subjected to change.
effects of interference
We
reversal.
that
by
various
diphasic
rise to
negativity, or the
may
even be multi-
will
responses.
sequent chapter.
I
shall here
by means of a
the
contained
ordinary response
currence
of
the
effect
in which,
we can unmask
of positivity, from
The
Mimosa
galvanometric negativity.
of
excitatory
contraction,
in
the
ocfor
leaf,
depends on a
If this motile
fall
of the
excitability should be in
say,
true
these circumstances,
opposite point
we
S^ (fig.
et armis, to
produce
is
COMPARATIVE ELECTRO-PHYSIOLOGY
66
by the
loss
such
as
may
Similarly,
be selectively blocked,
anaesthetics,
without
the
appreciably
agents,
the
affecting
cm.
below
the
proximal .contact.
The
first
three
H/-y^
Fig. 49.
First part of record shows normal negative responses ; second part shows
positive response unmasked by selective physiological block of chloroform ; in the third part is seen the abolition of response when stimulus
is
itself.
of chloroform
is
POSITIVE
6/
the
The
abolition of
we have
If
now
the stimulator be
area, the
On
doing
this, therefore,
hydrostatic positive
Response
point.
to
is
effect,
its
we
find
consequent
is
We
two
shall consider
effect
there
is
first
namely,
the
direct,
In that which
we
or true excitatory
is
indicated
turgidity-variation.
on a tissue
is
generally speaking of
much
effects
It
may
be transmitted even across tissues of which the excitability has been depressed or abolished, whereas the transmission of true excitation
is
arrested
by the intervention of
The occurrence of
nature.
It
electrical
changes took
F 2
68
'
COMPARATIVE ELECTRO-PHYSIOLOGY
That
this,
variations.
CHAPTER
VI
growth-response
Increased
internal
ment of
spiral tendril
We
have seen
in
external stimulus,
positivity.
These two
in
and
internal
their general
expressions.
But while stimulus from outside, impinging on an excitable area, thus caused an expenditure of energy at that area,
COMPARATIVE ELECTRO-PHYSIOLOGY
70
may suppose
Thus, we
The
the leaf
now be
fall
of
making them
over- turgid,
and raising
pulvinus, which
their
The
variation.
merely
of the plant.
this internal
energy which
is
it
consists of the
sum
total
may be
very various.
imparted by
thermotonus
light, or
;
by
We
have
phototonus
for
instance
by favourable temperature,
by internal
or by the presence of favour-
tonicity, as
We
and
different
modes of
by many different
manifestation.
And
such a
if
we should succeed
in
7I
still
is
'
Thus
it
will
be seen
the
movement
and
For
the
leaf, in
of the
leaf, in
of growth.
This
parallelism
is
external stimulus
is
the depression
of the
motile
leaf,
or
ment
is
leaf,
or enhance-
We
of internal energy.
One mode
is
COMPARATIVE ELECTRO-PHYSIOLOGY
72
Growth response
Contraction
Contraction
Diminution of turgidity
and concomitant depression of leaf.
Action of internal energy exhibited by
(a)
Recovery
[b)
energy of a plant
is
Diminution of turgidity
and concomitant depression of rate
of growth.
Action of internal energy exhibited by
{a) Recovery:
Re- establishment of turgidity and
gradual return of organ to normal
rate of growth.
{b) Increased hydrostatic pressure
Increased rate of growth.
:
by a moderate
rise
of temperature.
Thus, when
Mimosa
is
And
by the
raised in tem-
all
its
leaves
Thus
is
exalted by rise of
in a
three
As
we have
on the motile organ is one of depression.
The following record (fig. 50) shows that it has a similar
influence on the rate of growth.
The first part of the curve
shows the normal rate of elongation. But after the applicaregards external stimulus, on the contrary,
seen that
its effect
there
is
growth
is
of stimulus, the
established.
normal
rate
of growth
On
is
the cessation
gradually re-
in
73
effects
in
period
the
of variation, as a stimulus;
temperature
Fig. 50.
itself,
or
its
The
steady,
the
condition
will
(fig. 51),
The normal
rate of
74
Or
the
same
of mechanical
In
may
difference
response.
this
A
to
be demonstrated by means
Mimosa
there
placed
is
sudden
a small
in
of temperature.
rise
a preliminary excitatory
is
rise,
by
above
the
carries
leaf
original level.
its
effects
in
We
take
citable than
is
outer
more ex-
and
cave,
the
this,
or convex surface
of
tendril
spiral
In
Passiflora.
external
stimulus,
sponds
Record of Growth in
Crinum at Temperature of 34 C.
and 35 C.
Fig.
51.
The
The
rate of
stant
be-
the
to
excitatory
of internal
increase
fall
The response by
of Mimosa.
energy
is,
this.
When
vv nen
purlinp^urung.^
the
ine
tendril
lenani
is
by a movement of
curling,
when
the
The
85 (Chapter X.)
sudden variation of
In
fig.
temperature, acting
had
opposite
the
effect
that,
75
namely,
of
thus
is
that while
seen
the
characteristic
tive
is
is
of increased
effect
of
to cause a nega-
internal
energy
The former
of
of the
leaf,
and
is
in
in
The
rate of growth.
latter, or
growth.
We
isotropic organs
1.
Effective
stimulation
induces
contraction
and galvano-
metric negativity.
2.
Increase
of
internal
galvanometric positivity.
energy
induces
expansion
and
CHAPTER
VII
external
critical level
level
Sub-tonic,
Outward
exceeded Three
is
typical
cases
when
critical
and response
than stimulus Investigation by
Instance of sum of work, internal and external, performed
by stimulus constant Positive response of
characterised by feeble
protoplasmic activity or sub-tonicity Enhancement of normal excitability
response equal to stimulus
less
growth-response
tissues
We
shall
find, in
this
by the
festation
is
of even
moderate stimulus
will
then
prove
condition
is
sufficient
to
tissue in this
If,
on
Here, during
effect.
the
first
however,
it
is
at
work
part
As
of
application,
a matter of
fact,
rendered
effective.
is
favour-
sion.
on recovery, return to
its
original
induce the
stimulus will
first.
These responses
owing to the after-effect of
as the
if,
fatigue
molecular con-
following chapter.
It
will
It
is
is
be
namely, of great
excitability of the
or
little
The
no outward
incident stimulus
of internal
increase
expression
And
may
the
find
galvanometric positivity.
In the case which
tissue
is
we have
Hence the
any
loss.
The whole
responsive expression will in this case consist, not of contraction but of expansion.
is
In other words,
it
will
be exactly
In order to demonstrate
this, I
took a seedh'ng of
Tamarindus indicus which had been cut off from its supply
of external energy, and was consequently sub-tonic.
Owing
of
internal
energy,
its
growth
to the insufficiency
had, in
fact,
come
to a standstill.
On now
to
organ
in
is
exactly opposite.
COMPARATIVE ELECTRO-PHYSIOLOGY
78
is still,
we may expect
a case
In such
Hence,
if
we
the
first
factor,
we
shall in
with
its
and enhanced
positive turgidity-variation
rate
of growth.
this
accumulating income,
now
pre-
dominant, resulting
traction,
in the
It
it is
necessary to have
This
The
upon a second
mirror, which is subject to a compensating movement. When
the balance is exact, the spot of light, reflected from the two
Lever.
falls
When, however,
there
is
balance
a
is
upset.
movement
is
accelerated,
in
in
one
the
of external stimulus on a
of Crtnum
horizontal line
and
it
will
(fig. 52).
took a flower-bud
now
applied,
But by
this absorption
itself,
condition
the tonic
citability,
it
now
as seen
now
by the downward
cut
shows
is
to say,
The
the
after-
after-effect
mere
not a
thus
curve.
the
of
the
The
effect.
is
and
off
portion
dotted
curve
That
re-
due to the
which
remains latent.
It was
only when this was exhausted that the normal
rate of growth was re-
rate of growth,
energy
increased
established,
seen
as
in
And
curve.
now
was
in
condition
tonic
renewed
of
did
light
full
the
application
of
as the tissue
stimulus
not again
hancement of the
growth,
contraction and
tion.
rate of
Its
Growth
effect
during
retarda-
phenomenon of multiple
fully dealt
Fig. 52.
rise to
the further
Now
COMPARATIVE ELECTRO-PHYSIOLOGY
80
the tissue
itself,
and we
traction
Thus,
in this
intermediate
negative.
(3)
And,
lastly, that
when the
tonic condition
is
already
and
Both this and
by the experi-
we
effect
sub-tonic
lily.
of incident light
It
was
into
condition
the
being raised,
excitatory negative.
ternal
find
it
clear,
from
considerations already adduced, that the lower the tonic conthe greater will be the proportion of stimulus held
The nearer is
latent for the performance of internal work.
dition
On
component.
The
internal
subjecting
stration
growth.
it
We
have seen
that, of the
will
in the
two antagonistic
predominate
if
former
factors of
the excitability
be
in
We
We
have to bear
stimulus
is
called
shall here
sub-tonic.
COMPARATIVE ELECTRO-PHYSIOLOGY
82
possible.
The
depends on
its
tonic
incident stimulus.
condition
Now
may
it is
should
MT
be
the
cause
level
critical
exceeded,
slightly
if
there
C to
to
is
that
quisite to
in
evoke response
will
be greater
itself is low.
tissue
is
ST
Diagrammatic
Fig. 53.
Representation of the
Tonic Level
N, normal ; ST, sub-tonic
HT, hyper-tonic ; and
c, the critical level.
the
sti-
Thus
will
afford
If the latter
us a measure of the
stimulus will
precipitate
spontaneous
activity.
ENEkGY
IN RESPONSE 83
if
we
This considera-
evoked
shall
show
in
in cells of epidermis,
in general to
be positive.
Even
when cut
off
sub-tonic as to give
how
positive
response.
dition
is
allowed to
fall
effect,
show
shall here
when
tissue
somewhat withered.
its
are at
into
COMPARATIVE ELECTRO-PHYSIOLOGY
84
may
also be demonstrated
tinuous stimulation
positive
is
or
tetanisation,
The
Any
of galvanometric negativity.
citability
Fig. 54.
tive
whether
by lowering of
diminution
is
one
of the ex-
Photographic Record of Abnormal Positive passing into Normal NegaResponse in a Withered Specimen of Leaf-stalk of Cauliflower
When
causes
may
will bring
culminate
in
about a decrease of
actual positivity.
this negativity,
Thus negative
which
is
not
becomes
latent
and
this in
its
condition
When
tissue.
positive,
is
When
the absorbed
its
stimulus
its
is
is
the
may
observe
many
effects
in
different
will
types,
If,
response,
positive
The
present
chapter has been that of the energy received and given out
by the
tissue,
stimulus,
by
following chapter.
CHAPTER
VIII
dis-
in
effects
in-
A, B, C, D,
effect
tinuous stimulation
raises tonicity
and
than stimulus
Instances
excitability
pression.
According
in a state of equilibrium
cesses of assimilation
stimulus causes a
and
down
is
maintained
dissirriilation.
or
supposed that
is
On
this theory,
each other.
is
falls
below par
for
it.
is
supposed to be further
fatigue-
stuffs.
lies,
however,
of
rest.
In
isolated
is
effect,
8/
no active circulation
and its removal after
enough
case
which
arise
but
when we come
in
still
cardiac muscle
The
enhancement of amplitude.
in
direct opposition
we have
which
in
exhibit a
gradual
to
those
greater are
for
in
are
this
to
sive negativity.
In actual
characteristic response
For
the case.
in
fact,
is
is
by
galvanometric negativity
sponse.
It would be difficult again to conceive of assimilation
and dissimilation in the case of inorganic matter. Yet even
in inorganic
met with
in
and
all
fatigue.
namely, uniform
Response being
we can
COMPARATIVE ELECTRO-PHYSIOLOGY
88
One
anomalies hitherto
encountered
phenomena
responsive
most important
of the
factors, then, in
are
all
our
in
shall
have arrived
the
fact,
it
For a
full
exercises on
though
From
of
referred to Chapter
the
interpretation
response, however,
is
The numerous
itself.
at,
the reader
is
XLII.
every type of response
that
to
is
be
borderland
phosphorus
in
in the
the yellow.
Under
chemical activity,
being
less active
than
it
is
geneous
When
living
tissue,
when
unstimulated,
is
homo-
iso-electric.
is
The
result
is
an
electrical
current
attended
may be
pressing effect on
as, after
is
activity of the
may have
tissue.
a de-
Hence, just
it
89
be necessary.
tion, will
mental
chemical events
may
how upon
menon
thus seen
It is
the funda-
way
we can avoid
that
And
it
is
the
many
contradictions
with
we
case.
the
But
process
entire
this
is
of response,
while
stimulus
is
amplitude.
perature,
We
shall
thus
by enhancing the
often
find
a diminution of response.
shall
now
exercised on response
by the molecular
shown,
(XL
in
its
conduction.
the Chapter
1)
condition.
It will be
on the Modification of Response
it
COMPARATIVE ELECTRO-PHYSIOLOGY
90
first,
we have
By
where response
transition,
Above
normal.
tion
is
now
the substance
which
From
the stage of
molecular transforma-
a condition
is
is
is
very rapid.
absorption
this
less
to the stage C,
Further
stability.
D and
Here
E.
Stimulation
great.
applied
in
this
is
has
condition
very
little
here reduced to a
is
distorted
position,
absorption
obtain
abnormal
is
we
long-continued
Intense or
positive.
of tissues.
somewhat
here
is
sub-tonic,
From
rapid.
cules of the
is
still
distorted in the
proportion
is
thus utilised in
will
now
Thus
repetition of
at the
This
fact
stage
we
obtain a stair-
that by the
may
after-effect
be incipiently
effective
by
in
stimuli individually
repetition.
The
ineffective
dis-
finds
being
made
made
91
the
all
The
In
staircase effect
fig.
is
is
55
increase in the electrical response of a vegetable nerve
in
has,
is
been
said,
here, for a
The molecular
income.
by
condition
give
Fig. 56.
Staircase Increase
in Electrical Response of
Petiole of Bryophyllum,
rendered
sluggish
by
cooling
by
hibited
different
responses
electrical
by the root of
In
tissues.
to
radish.
uniform
Fig.
fig.
57 are seen
uniform
The
a
is
succeeded
by stages D and
condition of over-strain.
form
'
An
responses
to
In
uniform
fig.
59,
stimuli,
E, representing
are
applied
shown
at
uni-
intervals
XXXII,
COMPARATIVE ELECTRO-PHYSIOLOGY
92
of one minute.
An
inspection
of the
record
shows that
is
tion.
other.
in
Fig. 57.
of half a minute instead of one, while the stimuli were maintained at the
Cfig.
same
59, b) that
intensity as before.
It will
now
of
be noticed
much
smaller
.Ll-L\^i.
Fig. 58.
shows
that,
when
An
inspec-
greater frequency of
strain.
The mole-
There
its
thus a diminution
is
93
of
height
in
the
resultant
u
()
(c)
(^)
Fig. 59.
Record showing Diminution of
Response, when sufficient Time is not
allowed for Full Recovery
Fig. 60.
Fatigue in
Celery
enhanced (Radish).
response.
restored,
increased response.
Residual strain
is
thus seen to
celery
(fig.
there
that,
molecular
60).
It will
owing
recovery,
be noticed
imperfect
to
during
the
the heights
Patigue in Leaf-stalk of
Cauliflower
Fig. 61.
rest,
a photographic
Stimulus
Fig. 6
record
30 vibration at interval
^ "' "^^""'"
of
and
fig.
62 of fatigue
in
inorganic response.
It is
will
be greater
if
This
is
COMPARATIVE ELECTRO-PHYSIOLOGV
94
seen in fig. 6;^, where the first set of these responses, A, is for
an intensity of mechanical stimulation of 45 vibration, and
the second set, B, of augmented amplitude, for an intensity
of 90 vibration.
On
reverting, in c, to the
first
of
intensity
stimulus-
45,
the
re-
first set, A.
This change
due to the
is
But
excessive stimulation.
we should expect
effect
that the
of such over-strain
continuously
for
period of
In order to
rest.
re-
(at
the
several
newed stimulation
Days
intensity
of
45)
fifteen
minutes after
fatigue
far
45'
B
Fig, 63.
The intensity ot
Successive stimuli applied at intervals of one minute.
stimulus in C is the same as that of A, but response is feebler owing to
Fatigue is to a great extent removed after
previous over-stimulation.
fifteen minutes' rest, and the responses in d are stronger than those in
The vertical line between arrows represents -05 volt. (Turnip
c.
leaf-stalk.)
One
that,
first
owing
is
relatively large.
these curves
little strain,
is
the
The succeeding
95
The
are similar.
first
response in
because
we
The
intensity of stimulus.
first
6^y A,
fig.
The
rest.
first
shows
first
this,
because
of B shows
it,
time to an increased
of c does not
show
it,
because
And
stimulation.
minutes'
Of
the
first
it,
because
rest.
And
its
may
it
it
response
is
be the maximum
contraction and galvanometric negativity.
the
in
first
effect
will
negative
response,
fatigue
is
of the response.
very great,
as,
for
instance, in
certain
The normal
contraction
now
muscles,
(fig.
64, a).
by
rise to results which are essenbe noticed that after the responsive
fall of the leaf it returns to its former erect position, in spite
of the fact that stimulus is still being continued.
Here also,
tion
electrical
tially the*
same.
shocks gives
It will
we have
the usual
fig.
64,
d,
This
is
seen
electrical response
COMPARATIVE ELECTRO-PHYSIOLOGY
96
is
effect of rest in
photographic records
curve
will
The
66).
first
which
attains
first
amplitude,
large
it
declines,
minutes
after
in
Another
fatigue-reversal.
two
(fig.
in
is
of electrical response,
the response
The
now
were
Fig.
65.
Photographic
Records
of
figure),
Fig. 64.
Response)
and
to
Continuous Stimu-
two minutes'
rest.
The response
first.
in this case is
The same
seen to be
as
before
97
shows
greater
which
is
maximum
contraction
Fig. 66.
is
\ea.( of^
induced,;
AfzmosaJ
In the
This
is
first
followed, however,
with re-erection
of the
by a
leaf
namely, expansion,
According to the chemical
reversal
assumed
to be
IL
COMPARATIVE ELECTRO-PHYSIOLOGY
98
Now we
about.
response
may
of
may
is
of a continuous income,
expression
sion
be the result
is
the recipient
its
responsive
Thus
Taken by method
of conductivity variation.
(fig.
widely present, as we
the response, not only
ing record
is
shall see, in
each
Such a phasic
substances.
The
follow-
a large
number of similar
may
alternations.
may
99
I
often noticed
refer to
mechanical and
both
electrical
responses.
The
simplest
d,
c,
responses.
record
the
Fig.
(fig.
leaflet
of
Desmodiuni gyrans.
Photographic
69.
Record
of
Periodic
Periodic Fatigue in
FiG. 70.
Pulsation of Frog's Heart
modium gyrans
(Pembrey and
Phillips)
(fig.
70).
we
(fig.
71),
already understood,
observed.
assimilatory
occurs.
'
recuperation,
we
which pulsating
We
phenomenon of
H3
TOO
COMPARATIVE ELECTRO-PHYSIOLOGY
individual
response to
individual
stimulus,
into
that
of
The excess
stimulation.
stimulus
is
subsequent expression
in
XVII).
There can be no doubt that these effects of periodic
alternation of phase are due to two antagonistic reactions,
becoming effectively predominant by turns. Thus the continuous impact of stimulus on a tissue may first give rise to
But by the continuous
the negative phase of response.
absorption of incident stimulus, the internal energy is inpositive variations in series (cf Chapter
with
creased,
opposite
its
of
reaction
positivity.
gradually
will
diminished,
of
two
these
Fig.
curve.
less
sents
horizontal
state
line
repre-
balance
of
balance
is
disturbed,
by a sudden
When
this state
cessation
of
the
is sometimes seen,
with examples of
meet
followed by
retina and certain
of
responses
this in, among others, the
Such facts it has been
vegetable tissues under light.
suggested afford a demonstration of the two antagonistic
We
shall
really to
be accounted
for
will
met with
258 and 383.)
figs.
lOI
in the
response of
we have
We may
augmenting
force.
The
imagine a
from a gradually
be considerable.
But
this will
will
seem
to
may
spiral
soon reach a
first
beyond
limit,
have but
further
responsive distortion.
we may
the stage
reached
or E.
its limit.
Fig. 72.
In a
But the maximally distorted molecule has now a great tendency to revert to the position of
equilibrium, and the shock of stimulus, instead of inducing
little
further distortion.
That
plained by
is
response
of
this
is
to be ex-
(fig.
thermal
to
This
stimulation.
progressive
whose
decline.
and
to one of expansion.
shall
response
another record
in
COMPARATIVE ELECTRO-PHYSIOLOGY
I02
The
under
reversed
fatigue,
am
tissues
may be
experiment
of an unexpected character.
It
is
usually
supposed that fatigue is typical of such tissues as muscle,
and absent from nerve. But I shall show with regard to all
none of these
is one
of degree and not of kind.
The same intensity and duration
of stimulus which is efficient to cause
fatigue in muscle will not be enough to
do so in the case of nerve. But even
the various types of response, that there
which
is
distinctive of
any one
nerve
display
will
fig.
73, a
difference
fatigue
cessively stimulated.
in
is
The
tissue.
when
ex-
particular
nerve
of frog
Thus a
none which
of any one
Fig. 73. Reversed Response of Fatigued
is
exclusively characteristic
is
Were
tissue.
it
otherwise,
Nerve
in
effect.
For at first
74) shows that this is not the case.
exhibits a characteristic staircase effect the responses are
record
it
is
of
(fig.
and
to
be
seen,
muscle, in
fig*
75
is
is
we
see fatigue, in a
staircase response
is
lastly,
in stages B, C,
and
The
nerve,
D.
and even
in
inorganic
in
skeletal
substances.
In
The
I
103
its
is
is
In
responses given in
fig.
74, the
same
is
served
in
sponse
of
When
the
however,
phenomenon
similar
the
is
is
ob-
mechanical
re-
in^
Fig. 75.
crease, followed
Preliminary Staircase,
Fig. 74.
followed by Fatigue, in the
Responses of Muscle (Brodie)
(Resistivity variation
method)
mode
of record
is
by
(fig.
64
b).
resistivity-variation,
variation of resistance at
fig.
first
is
And
found gradually
finally,
we
when the
find,
by the
Instances have been given, in which a portion of the incident stimulus has been seen to be held latent to
work.
And
from
this
it is
do
internal
is
quite un-
COMPARATIVE ELECTRO-PHYSIOLOGY
104
tenable.
incident stimulus
is
in
This
manifestation externally.
strate
while a single
moderate stimulus
single
is found to give
have shown, not
stimulus
This
only
in
and the
in electrical response,
subject
latter
up
Thus
in plants.
in
be taken
will
a subsequent
in detail in
chapter.
And, lastly,
what has
that incident
follows
it
from
been
said,
stimulus
need
on
but that,
it
may
par.
example
which incident
was seen to find
bifurcated
in
to a
fig.
The up
j6
expression.
is
given
responses
the
in
of standstill.
stimulus
applied
must,
The
in
increasing
tonic
find
that
internal
energy,
or
the
sixth
the
in
the trend
portion
the
first
organ,
thus
condition,
response
stimulus
on
we
now
and in
downwards of the
of
five
first
however, from
which
style
In
photo-
of
above
it
describe an
in
stimulus
76.
now
shall
Photographic Record of
Responses of Style of Datura alba
Fig,
contrary,
the
actually raise
hitherto
first
base-h'ne.
the series
to
tonic condition
I05
sixth of
But, as a favourable
is
countered
taking
are,
on the contrary,
satisfactorily explained,
on
full
molecular condition of
the
responding substance.
From
however,
it
and
exalts,
From
at
Instead of
this,
depresses, the
subsequent
re-
sponses.
plained
is
we have
is
with
its
greater excitability.
undergo a diminution or
The molecular
reversal.
a continuous
conditions
D and
E.
In the
stage, there
is
no true ex-
by the
abnormal positive variation. The substance is next transformed into stage B, where response exhibits a staircase
character.
In the next stage C, the respon.ses are uniform.
COMPARATIVE ELECTRO-PHYSJOLOGY
I06
Under
D and E
are
reached,
fatigue.
propriate conditions.
In a future chapter,
we
shall
study
from which
in
light
tissue,
CHAPTER
IX
by
in
differential excitability
Responsive
current from
the
more
to
the
less
we know,
sensibilities
stimulus, to the
and, indeed,
it
all
these diverse
movements
we
same
we shall
will
the differential
factor in the
An
give
These apparent
find, to
tissue
problem
on
investigation
COMPARATIVE ELECTRO-PHYSIOLOGY
I08
demands
that
we
first
discover
some means
tissue.
As
may
take a
compound
strip of ebonite
and stretched
india-
whole
is
the
is
more
contractile,
we
Of
the india-rubber.
lower
side.
In
fig. "JJ is
the
shown a
concavity of the
smoked
surface
lever.
by
excitable
If
^''cJntraSlf'^'Re!
we
being
that
more
which under
becomes
moderate
stimulation
diffuse
^o^ally
side
concave.
stimulus
apply very
on the upper half of the pulvinus,
sponse of Artificial
we
^^"P
traction
of this
half,
the leaf
though of greater
is
raised.
intensity,
is
stimulated
When
in this
more excitable of the two. This experiment may be carried out very easily by using the stimulus of
Fig. 78 gives the results observed {a), showing the up
light.
movement consequent on stimulation of the upper half; {b)
that caused by equal stimulation of the lower half; and [c] the
In the
resultant fall when the two are excited simultaneously.
that this half
is
the
we
find
it
excitable.
We
shall
is
is
the electrical
09
mode
of
citable
sides.
of the electrical
which might
contacts
possibly take
it
shifting
place
were allowed to
if
fall.
are
straws
fine
filled
with
normal
On
saline.
now applying
a series
of thermal stimuli, on
the
near
petiole,
pulvinus,
the
obtained the
responses given in
It
79.
be seen
will
{a)
that
the
current
Fig. 78.
fig.
responsive
flows
in
the
tissue
from the
tively
more excited
rela-
excited
upper, half
of the organ.
We
and
electrical
Diffuse
induces
greater
contraction
and
may
1.
On simultaneous
excitation of
two
points,
and B, the
responsive current flows in the tissue from the more to the less
excited.
no
COMPARATIVE ELECTRO-PHYSIOLOGY
Conversely, if under simultaneous excitation the responsive
2.
current be from
two
to A, then
is
points.
Fig. 79.
Mimosa
The
block.
petiole
is
local,
but diffuse,
On
is
now
necessarily
A and
B,
III
For
and
the
it
this
is
identical.
And
here
Fig. 8o.
made
at
A and
B,
and
tissue subjected
to vibrational stimulus.
This
is
true only
long continued
its
excitability.
excitability.
By the
unilateral action
of light, then, the organ has been converted from radial into
anisotropic, the lower side being that
which we
shall
expect
more excitable.
On mounting such a stem in the vibratory apparatus
(fig. 80), and making diametrically opposite contacts on the
to find the
find that
on applying vibration
112
tissue,
as
we
The lower
side.
it
will
be seen that we
is
applied directly,
thus,
is
it is
Moreover,
we next
longitudinally,
it
we
we may
Two
moistened rags,
in
pieces
inner
at
posed tissue
are
surfaces
current
the
diametrically opposite
is
is
now
excited
two surfaces
points.
outer
When
the
simultaneously,
and the
and
inter-
its
two
responsive
concave to the
outer convex surface, proving that the former was the more
excitable.
We
by
exact measurement.
may be done
present
I
will
be shown
a future chapter.
For the
method of stimulation which
bring to great perfection, by which
in
113
every
Variator,
way
satisfactory
and
by which stimulation
convenient.
is
will
be found
The Thermal
The
cm.
current
is
electrical
sent,
is
circuit,
Fig. 81.
The Thermal
Variator
The
time,
by means of a metronome
variation
(fig.
81).
The thermal
closure,
ebonite
clip, c,
heating-spiral.
adjustment.
contacts
fixed on a slide,
This slide
is
S,
experimental
tissue, these
the
e;
and
E'
COMPARATIVE ELECTRO-PHYSIOLOGY
114
ment
is
made, the
slide is
coil.
pushed
When
in, till
the circuit
the tissue
is
well in
completed, for a
is
to the
disturbance.
The
will
be no thermo-electrical
responsive
tion,
That
it
is
the
when
the tissue
is
seen
from the
fact
that
is
opposite
is
This
is
Musa.
The
may
sheathing
use the
mounted
in
B,
petiole of
for
purpose, because
practically
many
A.
required piece
is
cut and
have mentioned
I
no sign of fatigue
Musa
when
as suitable
to show
There are
which would doubtless answer
find
it,
in its
fresh,
responses.
it
is
found that
the responsive current flows across the petiole, from the inner
uniform
(fig.
82).
This,
it
subjected to the
will
be seen,
in-
and concavity of ^
rpotije
more
excitable side
the
is
is
more
II
that which, on
diffuse
stimulation,
it
be
spicuous
mechanical response.
P^-lUJ-^
^ly
Fig. 82.
First series,
normal
after application of
chloroform subsequent
depression.
has
many
sequent chapter.
J?
CHAPTER X
THE NATURAL CURRENT AND
ITS
VARIATIONS
Natural current in anisotropic organ from the less to the more excitable External
Increase of internal
stimulus induces responsive current in opposite direction
energy induces positive, and decrease negative, variation of natural current
Effect of sudden
Effect on natural current of variation of temperature
variation
Variation
of
physiological reaction
positive,
natural
Agents
Extreme
CO2 and
of alcohol
current
by chemical
agents,
referred
vv^hich
to
induce the
Action of
Effect modified by strength of dosevapour Natural current and
variations
excitation, the negative variation
Na2C03
its
unreliability of negative
variation
so-called
as
test of
excitatory
We have
seen that
by an external
when
the pulvinus of
stimulus, there
is
Mimosa
is
excited
is
half,
with a con-
Increased
to
show increased
the
its
Hence
is
made
we
In
which also we
power of emission
to be
energy,
We
energy
of
emission,
we observe
II7
in order to
absorption
of excitability,
ITS VARIATIONS
is
On
necessary.
its
would have
been necessary,
excitable
condition,
there
an electro-motive difference
is
between the upper and lower halves of the pulvinus, in consequence of which a current flows across the tissue, from the
less excitable upper to the more excitable lower half, which
is
We
it
opposite
effect
namely,
Under
we may expect
metrically
the
positive
the
rise to precisely
galvanometric
relative
to
be galvano-
to be galvano-
metrically negative.
tissues,
we
We
saw that
in
find
the
same
convex
side.
side,
The concave
is
more excited
effect of
of internal
stimulus,
energy
we should expect
is
it
an
Since the
COMPARATIVE ELECTRO-PHYSIOLOGY
ii8
83,
pulvinus
motile
fig.
electric
organ.
We
increased
or
diminished
internal
of
internal
energy,
we have
As
already seen
temperature,
rising
diminution
the case of
The
gradual
natural current, N,
ascent
or
In
Mimosa^ we
depression.
In
of increase
crease of internal
or
de-
energy
descent,
Musa
rest,
thermal
Upper and
is
by
temperature.
falling
Parallelism of Natural Current
Fig. 83.
in Pulvinus of Mimosa and Sheathing
Petiole of Musa
its
con-
being,
secured
trariwise,
face of latter.
effect of
variation of current.
The specimen
of
is
and external
take
first
surfaces,
and led
chamber
fig.
is
this gradually
To
It will
be
i.
When
the chamber
is
return
to
To
study the
ture, the
the
to
diminution of current
is
effect,
chamber
is
ITS
temperature of
it
is
the
II9
room, this
annulled.
temperature,
VARIATIONS
found
(fig.
84, right
means of the
On
cooling
"
annulled.
is
That these
effects are
is
exactly
tiometer.
It
was specially
temperatures
apply
tions.
thermal
ascent,
we
about an increase of
the existing current.
But since sudden variation of temperature acts as a stimulus, we shall, in the preliminary stage,
obtain an excitatory reaction, which will cause a transient
have seen that a steady
rise
brings
rest.
when
is
the current of
rest.
This
steady,
give here
(fig.
will
by an
be followed,
increase of
85) a photographic
COMPARATIVE ELECTRO-PHYSIOLOGY
I20
curve
we observe
In the
effects.
movement
a sudden
rise
part of the
first
This
of temperature.
is
field.
so
We
we
and
enhanced above
On now
normal.
the
chamber
is
down
to cool
allowing the
to the original
more or
normal value.
We shall next study the effect
of chemical agents on the natural
less to its
current.
Fig.
Photographic
85.
is
The mode
the contacts at
steady Rise
of Temperature on Natural Cur-
the substance
tent, I
Musa
in
natural
current
place, as
shown by
up curve
when
temperature
steady there
variation,
first
rise
of
becomes
is
as
a positive
shown by
the down curve ; on return to surrounding temperature, the normal current is restored to its
original value.
procedure
of
the
be
liquid,
applied by a pipette.
the specimen
is
same
If
it
on both
time.
If
can
be
it
be gaseous,
placed in a chamber
to
effects
obtain
stream.
of
various
which
results
is
observing
In
agents
are
at
we
first
For example,
certain
and others an
moreover,
is
increase.
The
found to be modi-
fied
in a
dose.
current,
may
often
well
is
to
be
modified
ITS
VARIATIONS
by the amount
121
of
the
dose.
Much
we
light appears to
is
it
of,
induces an
If now we apply it in
on the two diametrically opposite
contacts of
Musa, we
shall
reagent this
is
negativity, resulting
On
rest.
to say,
it
in
the application of
that
to the less
excited convex.
It
is
effects,
we
are able
to
induces the
We have
seen that
excitatory effect
of
or positivity.
which
in
agent,
may, when
Now
a chemical substance
If the
might be
positive,
to be succeeded
application
We
COMPARATIVE ELECTRO-PHYSIOLOGY
122
of rest
tion
would give
rise
its
action, to a reversed, or
These inductions
rest.
am now
about to describe.
Fig. 86.
of
of
Musa
P^iG. 87.
Effect of
CO, on
Musa
Natural Current of
cent, or
is
solution of
If
now a
dilute
And
if,
lastly,
we
we
We
fig.
87
In
It will
ITS VARIATIONS.
in
23
the
first
by a
followed
diminution.
We
have here an
Vapour of
that
is
is
now undergoing
to say,
it
of the
alcohol
induces a
the natural
surfaces,
reaction.
made on
iso-electrical surfaces,
electrical contacts to
be
But
The
resulting disturbance
may
be
dis-
if
the
contacts,
is
indicative ot
a positive variation
positive
an intermediate dose.
of the
at
existing
current,
is
due
COMPARATIVE ELECTRO-PHYSIOLOGY
124
will
in-
It follows
and
1.
may
variations
its
Under normal
from the
the tissue
phenomena of the
be summarised
in
natural current
general as follows
less
is
more
In other
excitable.
less excitable.
Increase of internal
3.
The
rest.
ditions have
now been
its
studied.
may occur.
may thus
or less persistent.
It
in
an
By
this excita-
may remain
is
it
of galvanometric negativity.
We
negative variation
things
we
shall
distinguish
frequently happens,
stimulation, with
appearing at
first
of the
current
of
as the
however,
its after-effect,
in
rest.
primary
This state of
condition.
It
consequence of previous
ITS
direction,
We
VARIATIONS
rest,
125
current of response.
understanding of these
observed
shall
effects,
in
differential excitability
and
We
differential
which occur
more
in the
shall here
changing
effect,
stimulation.
is
positive
the
tion,
current
is
up
t.
we have
down
seen, the
to
On
i.
negative and
stimula-
response
the
We may
now suppose
about the B
At
zero.
this stage,
ability of a tissue is
annul
will
now be
enhanced.
be up
will
t.
We
condition
tissue to the
relatively
t.
will
The
half.
we have already
seen,
is
zero.
bringing the
stronger excitation,
still
C condition,
of the lower
upwards
of
so-called
current of rest
is
greater
it
is
thus
still
follows that
is
upwards
t.
COMPARATIVE ELECTRO-PHYSIOLOGY
126
the resting-current.
And,
we may imagine
finally,
lower
citability of the
half,
owing
The
or E.
t.
to fatigue, has
now become
upwards,
this
As
1.
a negative variation of
appear as
will
it.
Condition.
rest.
Current of
Primary condition
P'eebly excited
Moderately excited
Strongly excited
The
rest.
Negative variation
Positive variation
Negative variation
and
various
accidentally in
Variation of current of
fatigued
re-
sponse.
conditions
mentioned
may
may
be
induced
be brought
about by the excitatory effect of experimental preparagive here the records of certain experiments
1
tion.
is
The
Owing
is
from below to
we observe
pegatiye,
J\
positive,
having
As an
formerly
The
after-effect of stimulus,
was
88).
strong stimulus,
first
above,
there
moment
(fig.
now
surface,
which
become
relatively
rise to
a response
But
first.
after
this,
VARIATIONS
ITS
owing
to
the
127
greater
From
it
will
be
seen that hardly could any standard have been devised for
the study of excitatory reaction, so likely to be prolific of
confusion as
the
in
displayed,
cases
three
this,
For
current.
first
we
now
appearing
effect,
as a
In
rest.
again,
the
abnormal
case,
actually
is
it
current of
fourth
the
which
response
But while
responses appear to
variation
tive
these
Natural current
is
nevertheless con-
The
stant.
direction of the
responsive current
is
always
less
excited.
normally
is
consequence of
It
in
in
The first
strong externj},! stimulus.
two responses are normal, i.e. current
Strong
being from below to above.
stimulus is here seen to induce mulAfter the second
tiple responses.
response on account of the greater
fatigue induced in the lower half of
the pulvinus, the direction of the
responsive current is seen to be
reversed.
Thick dots represent
moment of application of stimulus.
which
4,
down, reversed
the
course
conditions
of the
the
the less to
current
the
of
flows
rest
more excitable
that
in
increase of internal
its
diminution gives
rise to a
negative variation
that
those which
resting-current
of the
128
the
external stimulation
There
are,
many
In this way,
CHAPTER
XI
REAGENTS
Induced variation of excitability studied by two methods: (i) direct (2) transmitted stimulation Effect of chloroform Effect of chloral Effect of formalin
Advantage of the Method of Block over that of negative variation Effect
Response unaffected by variation of resistance Stimulating action
of
of solution of sugar Of sodium carbonate Effect of doses Effect of hydroConversion of normal
chloric acid Di-phasic response on application of potash
negative into abnormal positive response by abolition of true excitability.
KHO
This
is
by the
is
demon-
to
thus seen
It
killed,
This particular
respond.
to be a concomitant
electrical
response
physiological activity
But
in cases in
steam or by poison,
it
is
is
brought about by
is
of physiological efficiency.
W\\\,
the
It is also
impor-
way
it is
and
still
others^
response.
COMPARATIVE ELECTRO-PHYSIOLOGY
130
An
is first
to obtain
takable manner.
in
Chapter
III.
may
!l
Before
After
Fig. 89.
of Block.
is
transmitted stimulation.
We
And
first I
vapour
is
blown
by
chloroform,
is
to
be studied,
fig.
21).
the
In
131
contact
A and
experiment
is
B,
and
their close
in
out by
carried
first
The
neighbourhood.
obtaining a
series
of
agent
by being blown
It will be seen
84 how rapidly chloroform induces depression of
If the speciresponse, and how the effect grows with time.
men be subjected for a short time only to the anaesthetic,
is
applied,
from
fig.
depressing
the
action
proves
longed
air.
transient,
passing
off
on
abolition
of
response,
I
give below
(figs.
90, 91),
effect of chloral
llll^
^A.
Before
Fig. 90.
After
In the
COMPARATIVE ELECTRO-PHYSIOLOGY
132
in
91,
fig.
is
to
have postponed
now
till
some
difficulty.
we read
effect
this
Now
galvanometric deflection.
the
in
circuit
galvanometric deflection
the
of
heightened
\\\\\^^^^^^^
Before
After
Fig. 91.
excitability,
in the tissue.
the tissue
may undergo
But by
the resistance of
The observed
without any E.M. Variation.
variation of response may thus be partly owing to some
be induced
unknown change
E.M. Variation.
This
difficulty
galvanometric
deflections
being
An
now
proportional
actual experiment
will
to
the
make
EXCiTABlLlTY
Taking a
carrot as a specimen,
found
33
its
The
ohms.
to be 20,000
reduced this to 19,000 ohms. The resistance of the galvanometer used was 1,000 ohms, and the high constant external
resistance interposed
was
million ohms.
The
variation of
a thousand.
In studying the variation of excitability in animal tissues,
is
employed.
But
may
by
employment of the Method of Block instead. For,
in the method of negative variation, one contact being
injured, the chemical reagents act on injured and uninjured
unequally.
It thus happens that by this unequal action the
resting difference of potential is indefinitely altered. But the
intensity of response in this method of injury may to a
here draw attention to the advantage which
is
afforded
the
certain extent be
when
introduced which
may
is
is
itself.
If further
proof be desired
now obtain
A and B.
it
I
by the
give
obtained in this
below
way from
(fig.
92)
we can
two ends
a record of responses
This petiole
was somewhat conical in form, and owing to this difference
between the A and B ends, the responses given by one were
slightly smaller than those given by the other, though the
stimuli were equal in the two cases.
A few drops of a
10 per cent, solution of NaOH were applied at' both ends.
The record shows how quickly this reagent abolished the
the petiole of turnip.
COMPARATIVE ELECTRO-PHYSIOLOGY
U4
response of both.
(fig.
93)
is
given a photo-
KOH,
and
order to show
in
way
affect
After
Before
Fig. 92.
NaOH
Therefore,
responses
the
change
recorded
NaOH.
in
may
the
amplitude
of the
be taken as due
E.
M.
entirely to the
In
the
experiments
just
described,
the
stimulus was
By
the application
of
the tissue
receiving
modified,
stimulus,
also
EXCITABILITY
of the
effect
135
jointly.
the receptive from the responding point, and apply the reagent
on the
This
latter.
may
Before
Fig. 93.
Af,er
The two
KOH
this,
responding
the chemical
point.
It
reagent
will
be
is
applied
seen
that
is
taken
locally at the
the
receptive
excitability
area alone.
I
shall
now
COMPARATIVE ELECTRO-PHYSIOLOGY
136
In
Fig. 94.
fig.
94
is
1*5
The thermal
Fig. 95.
in
(fig.
95).
EXCITABILITY
137
that
a strong solution,
a solution of intermediate
Thus the
variation.
variation
positive
to
in
by a negative
last-named
the
be indicative of increased
excitability,
The
is
application
inducing excitation,
of a strong solution
we know,
the excitability
is
depressed.
large
quantities,
will
be found
to
induce a profound
may be
is
In
carrying out a
growth response,
found
stimulating in solutions
of,
say,
"2
to 5 per cent.,
per cent,
Another
fact,
tion, is that
general
mind
in
is
The
here seen to
range of safety
sphere of action.
is
is
difference
lie in
the
very narrow.
in this
connec-
thus
With copper
cause depression.
'
COMPARATIVE ELECTRO-PHYSIOLOGY
13^
is
different.
Fig. 96.
The
plant.
effect of
is
often
induce an
to
Fig. 97.
per cent.
KHO
per cent,
electrical effects
we
occur in the
and the
One
of these
of
we have
The
As
the latter
masked,
if,
positive effect
seen,
negativity
(p.
is,
relations of the
its
KHO
In
resultant
application
made
of
here given,
we observe
The
KHO
ability, as indicated
Later,
per cent.
of fern.
the
subsequent abolition.
KHO,
be un-
also
and
97)
excitatory effect
may
FiG. 98.
fig-
true
This positivity
66).
a resultant
is
by abolishing the
other.
the positive
is
39
metric negativity.
negativity.
we observe
is
first
seen
to
reduce
the
excit-
is
delayed.
I4O
Hence
Its
existence
twitch
in
is
now
is
seen
a di-phasic response,
downward
preliminary
in
to positive
this positive
response
is,
however,
CHAPTER
XII
of phase
Interference
through
intermediate
application of
effects
causing
di-phasic
KHO Effect
Diametric
balance
Effect
of
unilateral
of unilateral cooling.
ceed to describe a
new and
method of making
interesting
it
will
be found
remain
obscure.
delicacy, enabling
This method
the
is,
investigator
moreover, of
to
by any
detect
it must
extreme
the slightest
agent.
'
'
If
now
A and
B,
be excited simultaneously,
effects
E and
^^
Now
if
their time-relations
of equal
opposite signs,
COMPARATIVE ELECTRO-PHYSIOLOGY
142
'
up,'
also
be produced
the
if
B be depressed.
of
excitability
ment of
B,
'down.'
If,
same phase, we
The
curves.
resultant zero-response
may
thus be converted
into di-phasic,
is
capable
shall
now proceed
necessary additions
(fig.
99).
The angle
of
brass, B, projects
stroke
is
given to
this,
The amplitude
elastic piece of
When
a single
is
by the attached
induced,
spring,
S.
x^fter
deciding on a
angle on the
left,
we proceed
and taking
same
the
on the
as that
right.
I43
two
But
in
in the
be slightly
to
different
effect
therefore,
exactly equal
The two
on the
striking-rods,
left, R',
one on the
right, R,
in
the
same
Fig. 99.
B, elastic brass
vertical
The
left
other.
rod
When
is
is set,
same
pointing to zero of
vertical plane,
and the
same moment.
and
left
The
reproduced
the two
separate
by
excitatory reactions on
and
In
equal
fig.
100, a,
constituent
The
COMPARATIVE ELECTRO-PHYSIOLOGY
144
*
left,
up by the right. On the simultaneous excitatwo points, the resultant response was zero {b).
the excitation of one say, the right be increased by
and the
'
tion of the
But
if
response
found to be
is
'
up.'
It is
differential
effect
Fig.
[a] Isolated
{b) null-effect
We
when
shall
the stimulation
difference of phase.
left.
If, on the other hand, the
rod be set at a minus angle, the excitation of the right will
be later than that of the left. Under these circumstances,
become
is
less
increased,
and
the
neutralisation
of
we
shall
the phase
effects
will
In
fig.
loo,^.
seen
the
I45
of the
left,
di-phasic
elements of the
spondingly.
It is also clear
from
this that,
corre-
having obtained
if
'
'
'
We
how
thus see
resultant
down
'
'
negative or
up
'
reagent.
We
also see
how
and di-phasic
effects
(fig.
tin,
of the excitability of
its
reagents.^
of
Interference,
the various
we may,
as
reagents
we have
by
the
Method of
and
left,
and Non-Living,
p. 115.
146
been given
is clamped
at
stimulus
vibrational
will
excitation at
the two
given agency
is
induce equal
and
A and
The
points
tested
by applying
B.
it
simultaneous
effect of
locally,
say at
{a)
(d)
A,
I
any
and
shall
(.0
Fig. ioi.
We
have seen
in the previous
a tissue.
of Bryophyllum and
right
'
up
On now
'
and
left
took a petiole
first
'
'
Strong solution
I47
of potash
'
'
same
time
of
suppression
response
at the
by
increased
the
opposing
this
102).
(fig.
use of the
Method,
gate by
of
the
its
Diametric Balance
undertook to investi-
means the
lowering
of
on excitability.
ture
influence
tempera-
For
this
Photographic Record of
103.
Response of Petiole of Cauliflower
by the Diametric Method
Fig.
Photographic Records.
Di-phasic response of petiole
of Bryophylhitn^ the up component being due to the excitation
of right side. Strong application
of
on the right abolished
this responsive component, giving
Fig.
102.
{a)
application
sultant response
local
KHO
rise
in
{b)
enhanced
to
'
'
down
ice,
the
re-
response
purpose,
of
became converted
down
normal
to
up '-response
was restored on allowing the tissue
In this instance,
'
up.'
B.
The
point
its
L 2
COMPARATIVE ELECTRO-PHYSIOLOGV
148
of one minute.
restoration
It will
(fig.
103).
We
In a subsequent chapter
we
shall
by
variations
CHAPTER
XIII
of
theories
Different
Reymond Electrical
theory of Bernstein
injury
distribution
Hermann's
in
Pre-existence
Alteration Theory
strating
stimulation
Residual
Distribution
sectioned
cylinder
of
True
Experiments
potential
in
vegetable
tissue
with
significance
of
made
'
Positive
'
one
end
Apparent
current of injury.
transverse
contact will be
shall
demon-
negative, as
I
Bois-
If a section be
is
Du
Electro-molecular
electrical
Electrical
theory of
a muscle-cylinder
have occasion
found to
in the present
be
galvanometrically
contact.
explanation of this phenomenon and of the excitatory negative variation of the current of injury.
It
is,
therefore, only
The
that
Pre-existence Theory of
smallest
the
particle
Du Bois-Reymond
had
the
supposed
same electro-motive
each other.
longitudinal
cylinder
and
this
is
zone
is
motive difference
between
one
electrode
Thus the
placed
electro-
on the
COMPARATIVE ELECTRO-PHYSIOLOGY
ISO
is
moved
is
further
The
section.
of
the
distribution
symmetrical
is
with this
the
of
side
left
equator
on
tension
electrical
On
104).
(fig.
theory
Reymond
Fig. 104.
Distribution of Electrical
Du
of
^
smce been found
.
Tension in Muscle-cylinder.
adequate.
Bois^
tO
be m-
shall later
distribution of
all electro-
disturbance of equilibrium by
up and down chemical changes.
It is my intention to show in the course of the present
chapter that
the
over-stimulation.
due
to
current
And
molecular
of injury
since
upset,
we
is
shall
and
its after-effect
after-effect
it,
of
fundamentally
is
best
an
excitation
understand
if
we
first
the
study
factors
as
assiniilation
and dissimilation
is
We
have found
for
its
length.
In
the
first
when
a portion of
it
was subjected
to
151
any molecular
The
disturbed areas.
the
was strong
105).
(fig.
This
is
shown
in
the electrical
upwards
The
response
which
of
the
is
tin
as
same
persistent
volt.
"i
after-effect,
the
sign
of
change.
When
torsioned
wire.
molecular
distortion
covers
its
torsion
is
limit, it
the
slight,
is
exhibited mechanically in a
torsion
the
is
released
wire
quickly re-
of equilibrium.
COMPARATIVE ELECTRO-PHYSIOLOGY
152
even after
it
free.
Recovery
is
thus, in such
Turning next
to plant response,
we
find
a similar per-
And
stimulation.
first
we
shall take
the
simplest
case
that
in
Vertical line
'i
volt.
as seen
in the first
of the series
is
complete,
complete
(fig.
stimulation,
is
06).
seen
in-
exhibit
an after-effect of residual
is
really
due to incomplete
was applied
directly.
We
is
shall
in
53
which excitation
due
to strong stimulation.
We
know
and the
effective intensity of
such
of section,
we
greatest
at
is
lessened
as
shall
find
the point
the electro-motive
that
and
of section,
we recede from
it.
change
progressively
is
This induction
may be
verified experimentally
petiole, at
be as shown
in
'i
volt
5
1
It will
cm.
)>
,,
E.M.
in
Specimen of
100 divisions
difference
"i
Cclocasia,
volt.]
between indifferent
50 divisions
40
33
29
27
further away.
COMPARATIVE ELECTRO-PHYSIOLOGY
154
I
shall next describe the more sensitive galvanometric
method of investigation. The resistance is here maintained
constant by having the permanent contacts at A and the
(fig.
107).
Fig. 107.
ment
Fig.
decreasing distances
from
108.
Recoids showing
in-
for
A.
give
a series of records
injury
effect
is made nearer and nearer A, the permanent afterbecomes greater and greater. From observations made
of a similar experiment,
in the course
results,
given
value, with
in
tabular
form, which
lessening distance, of
metric negativity.
this
galvano-
55
25
220 divisions
180
cm.
I-O
,,
120
1-5
,,
75
40
2-0
3'o
,,
,,
Fig. 109.
series of
to
fig.
A,
lines.
It will also
be seen from
COMPARATIVE ELECTRO-PHYSIOLOGY
156
as between
the
and
sections a
equipotential
section
further
a!
equatorial
;
point
A and
away from
distribution
is
two terminal
galvanometrically negative, in
is
the
It will also
it.
reference
be seen that
to
one
this electrical
fig.
104.
Thus, without the postulation of any electro-motive molecules so-called, these experimental results afford a simple
injury, as
Fig.
10.
Ends Sectioned
The Current
after-effect,
due
of Injury
to
is
twenty-four.
The renewal
of the cross-
The disappearance
of negativity,
This
is
by re-stimulation.
owing to recovery, only
effect
when the
injury has
not been
excessive.
57
If,
however this has been too great, the injured tissue will then
But
pass gradually into a condition of permanent death.
change
the electrical
positivity, as
show
shall
one of
is
Thus the
in
may be brought
of an existing current of
is it
necessary
And
secondly
why
is
variation of
With
it ?
reference to the
already seen
that
of these questions,
first
when two
points,
A and
B,
Eg.
be
will
zero.
From
a resultant response,
bility
simul-
response
If,
we have
are
it
this
is
is
two
we can
we must
This inference
may
being
that,
there
COMPARATIVE ELECTRO-PHYSIOLOGV
158
owing
yet,
to the
irresponsive condition of
B,
there
is
When
the point B
a more or
is
injured, there
less
is
generally speaking
A.
But we saw that the direction of the action-current
was opposite that is to say, from A to B. This will explain
B to
why
reason
the
method of doing
this is to
points.
be such as to
If
this
excitability
One
then
its
Or by causing
we may simply depress
the
the
kill
tissue,
permanently abolished.
is
the
shall
now
negative variation.
area on
Taking the
surface, say B.
its
petiole of turnip,
current
is
we
injure an
now observed
to
The induced
of
the
plant,
described,
its
volt.
experiment here
In
the
between A and
B,
and a sudden
by -026
volt.
volt.
The
(fig.
iii).
with time.
This
is
often, as has
59
It is in
is
of injury
necessary,
is
by whose induced
it
no longer
exists.
But
variation
we may
effects.
supposed that
is
its
antecedent con-
shall
Tffni""^^''^'"^'
Fig.
III.
The
first three records are for stimulus intensity I ; the next six are for intensity twice as strong ; the successive responses exhibit fatigue.
The
vertical line to the left represents 'l volt.
The record is to be read
from right to left.
in the
positive.
when the
fact,
For we
current of injury
may
is
obtain the
zero, or
even
of resultant response
is
a simple explanation
injured,
it
is
often applicable.
die.
When
In fact,
the tissue
is
have often
COMPARATIVE ELECTRO-PHYSIOLOGY
l6o
found
that, in order to
of thermal
section
temperature
is
ensure death
necessary.
by the application of
heat,
we have merely
find that
exces-
But
away
is
more or
less restored,
The
Hence
contacts,
And
response, will
a fresh injury
found to renew
is
its
now
This
amplitude.
cause,
is
due
It
maximum when
reason
of
injury,
unopposed
that
is
abolished,
variation,
the resultant
electrical excitation
when the
it
due
the excitability of B
is
effect
at A.
be at
totally abolished
its
by
may
to hydro-positive action.
When
this
happens
may
excitability,
The effect
of
this, in
in the case of
conducting nerves.
As an
A,
it.
well seen
by injury reaches
point.
is
its
is
and causes
after-effect of this
l6l
response.
We
common
effect
of injury
is
in
to
We
stimulus
is
unknown
reason,
galvanometrically positive.
As
a result
response
is
The
may
first
be.
abnormal responses
is
is
is
different
The
The
points.
discussion
of the
first
of these
upon
here,
though
its
detailed
As an example
we may take
is
more
is not uncommon.
For example, we may have a
muscular tissue terminating in a glandular. Owing to this
anisotropy, the muscular and glandular surfaces are unequally
tropy
excitable,
and
it
will
generally speaking,
it
be shown
is
in
For further
details, see
When
such a
Chapter XLII.
COMPARATIVE ELECTRO-PHYSIOLOGY
l62
preparation
is
it is
found
From
this
it
but
all,
se,
In
is
the
excessive
its
tissue
is
We
for
transmission, and
physical injury
it is
is
obviously incapable of
is
con-
we
and
on
owing
to excitation
If,
next,
points of
surfaces,
it
will
point, relatively to B,
This
become
is
and consequent
more excitable B.
excitation
becomes galvano-
acts as a
strong stimulus.
It
shown that
such
after
stimulation,
there
As
negativity as an after-effect.
effect
is
Hence,
very protracted.
persistent
is
163
galvanometric
it
be
will
From
this
it
neighbourhood of the
cut, to
from the
excited end.
strong stimulus.
occurs in a muscle-cylinder
is
fact
two sectioned
ends,
and that
equator.
The
this
decreases
progressively towards
the
that
diffuse
its
the
excit-
stimulation, causing
it
injured
and uninjured.
This
is
the
explanation
of
An
'
positive
'
CHAPTER XIV
CURRENT OF DEATH RESPONSE BY POSITIVE VARIATION
Anomalous case of response by
variation
positive
and dead
tissue
living Explanation
We
have seen
response
by
in the last
negative
variation,
it
experimental
tissue,
the
lation,
kill
say by scalding.
is
customary among
one end of the
is
It
is
generally sup-
negative to living.
On
stimu-
now
tissue,
gation,
hitherto, as
we have
it.
variation,
is
In the course of
my
investigations
mode
on the response of
variation.
was
165
'
The abnormality of
08 volt.
fact, that
become
reversed,
cordant with
discovered
which the
and
con-
it,
positive variation
on,
now
(fig.
112).
many
Later
instances in
killed
to the unkilled.
Since, then,
it
is
itself to
spontaneous
variation
means
as
our
only
my
Fig. 112.
Response by Positive
Variation of Resting Current
The
the
causes
of
these
current of injury,
shown by
anomalous
tive.
reversals.
The
as to
My first
positivity
to study a case in
my
ceeded
in
we
find
Thus,
in the leaves
The innermost
of these
COMPARATIVE ELECTRO-PHYSIOLOGY
66
may be
patches
living tissue
is
quite dark
and discoloured,
while, as the
And beyond
this, again,
we
find
the
Proceeding thus
tissue.
from
its initiation,
towards
we
shall
somewhere
if
made with
various points
The
be found to be at
its
variation of elec-
succession through
in
The
be found to increase.
first
attained on
is
tissue,
little
beyond
inward from
decrease,
we come
till
iso-electric
with the
maximum
this, this
On now
On now
dead
point being
passing further
difference
to a point in the
living.
The maximum
found to
is
tissue
which
is
to the
we
still
find that
tissue.
The dying
and yellow
or
left
is
is
it,
the dead,
has been
most rapid along the
living.
It
CURRENT OF DEATH
On
radial line.
we
167
potential surfaces
The
first
which
be selected
which
tonic condition, on
modified in any
way
level
its electrical
we
depends,
is itself
which cannot be
that of complete death.
This may
is
ment
is
The only
change.
is
to be used as a standard.
is
condition
The method
level.
of experi-
will
abed, and
so on,
mm.
The
is
completely dead, to
first
placed on a and
The
b,
then on b and
^,
and
</,
and so
forth.
them-
in the construction
of
these, a
is
The
existence of such a
easily tested
two electrodes
in
Any
electro-
will
now
This
difficulty
may
be overcome by
first
taking special
COMPARATIVE ELECTRO-PHYSIOLOGY
68
In
saline.
eliminating
variations
cyclic
means
of
succeeded
by subjecting the
differences
all
of alternating electro-motive
commutator, without
Pohl's
in
electrodes to
By
force.
cross-bars,
the
E.M.F., and
of
the
resulting
variation
in
E.M.
the
intended to test
by
difference,
turns.
the electrodes
intensities of
for
this purpose.
was gradually
raised
The
this
machine
afterwards as
Original difference
After first cycle
After second cycle
After third cycle
It will
E.M.
Galvanometric deflection
Condition at starting
009 volt
360 divisions
40
difference
OOI
the
leaf.
was placed
hour,
till
and dead, of
The
electrodes,
along a radial
line
manner already
it
will
mm.
the
be remembered, were
CURRENT OF DEATH
Galvanometric Deflection
Position of Electrodes
05 mm.
5-10
20
25
30
10
15
15
20
25
30
40
40-45
It will
169,
division
20
45
140
140
no
-h
20
10
,,
M
,,
,,
maximum
being at 30
10
mm. from
20
15
25
30
35
40
45
gJMMillllll
50
100
150
HllillllHIIIIIIIIIIIIIi
HHIIIillllliili
11
200
iBWiiiiiiiiiiiii
250
I^I^HHiiHIiliH
350
400
HIHIili
dead
tissue.
coincides
with
This
point
the visible
Beyond
of
maximum-negativity almost
this,
however, there
is
an electrical
COMPARATIVE ELECTRO-PHYSIOLOGY
I/O
becoming increasingly
An
positive, as
(fig.
com113)
boiling water.
extremely
tissue.
easily,
are
It is
extremely
immersion
may
say that
it
is
tissues,
resistant,
in boiling
throughout.
this, I
difficult to
and
immersion would
cause
to
be
killed,
death
to
encroach
is
such prolonged
upon
those
This
difficulty in
tissue
difficulty
exposing
all its
hot water.
CURRENT OF DEATH
each of which was nearer than the
end, which was to the
peated,
last
by
The same
left.
171
mm.
to the
process was
dead
now
re-
successive
the
At each
or living end.
was balanced
difference
by
potentiometer.
the
straight
form
of
potentiometer
had
This
scale
thousand parts
and When
its
(fig.
Fig. 114.
AB
114),
terminal
adjusted to
volt,
Straight
plant.
each
was equal to
E.M.
Form Potentiometer
Distance from
maximum
( )
or
'oo
volt.
1*2
S'o
i8-2
i-o
,,
2-5
1-5
55
47
22 -O
23-0
2-0
2-5
3-0
23-3
3-5
4'0
4-5
5-0
The
cm.
6-8
lo-o
12-8
15-6
i6-8
i7'5
i8-o
55
5,
,,
55
we
find a point
is
portion
however,
is,
Thus two
iso-electric.
points,
and another
much
COMPARATIVE ELECTRO-PHYSIOLOGY
1/2
'01 8
is
dead
volt,
tissue
is
that
of the dead
is
-0233
volt.
Hence the
0053 volt.
We
Fig. 115
Nymph(va
alba,
The
maximum
difference, followed
ClJRkEKt OF DEATH
173
of
apparatus,
the
recorded.
An
may
the
animal,
concomitant
electrical
also be obtained, in
same phenomenon
the form of an electrical spasm of
record of the
galvanometric negativity.
Succeeding to
this rigor
of the
Now
it
will
in a tissue
all
expected.
alive,
be understood that
we
must
necessarily
through
pass
various
zones,
dying,
maximum.
Compared with
tracted living
is
this,
abnormally
relaxed dead.
The
death-frontier,
however,
is
not fixed.
The
line
It
of
is
con-
maximum
rigor
the
is
proceeding
so that a
COMPARATIVE ELECTRO-PHYSIOLOGY
174
what
is
known
as the
'
In the
injury-current.'
of these
first
hitherto been
current of injury.
Of
considered.
when
these,
made
is
we
find
shall
dead
it
And
contact.
thirdly,
be positive to the
contact.
We
the
first
first,
or
at
tissues,
living
on a
is
or living
first,
positive.
Taking the
contact
of these
first
negative
is
the
that
by a
injured
response
in
negative variation
to
of the so-
which the
in
action-current,
is
In the third,
zero.
injury.
1
here that
In the
up-line.
variation.
The
may state
record, in
first
The
by an
116,
represented here
almost neutral
that
is
to say, on
away from
the
are,
living contact.
point,
which
thus
became
positive
record
In the third
kill
the
the
injured
living.
The
75
is
It will
,vW^
-^{ifu r
Fig. 116.
In the
first
by negative
effects
namely,
a negative or a
variation of the
positive
injury-current.
give below
response which
injury-current,
is
a short
may
occur
summary of
when either
the diversities
(a)
^c
c
Br
m ^-z=
3,
^R
m^
*R
Fig. 117.
Current.
(a)
of
Specimen
originally isotropic
A, end slightly injured and negative ; C, current of injury ; r, actioncurrent, a negative variation of c.
(d) A, end killed and positive
C, current of injury ; r, action-current, a positive variation of c.
First
injured
we
(fig.
117, ^).
The
current of injury,
is
C, is
is
slightly
a->b, and
COMPARATIVE ELECTRO-PHYSIOLOGY
176
But when A
responsive
variation
current
117,
(fig.
Second
is killed,
also
is
-^ A,
constituting
^ A,
the
positive
d).
than B or
is
B'
(fig.
118,
The primary
a).
less excitable
is
be A->
effects
b'
and A
that
If stimulus
-> B.
is
positive variation of
A -
B.
(a)
-<:
(h)
> C -<
'
T
M
B'
l^^
and a
=^B'A'
p^^
tD
*
'A'
Fig. 118. Typical Cases of Variation of Current of Rest and ActionCurrent ; intermediate point naturally less or more excitable than
either of terminal.
(a)
current oi
Intermediate A, less excitable, shown by vertical shading
A->B and A->b' ; when right-hand point x excited, action-current
R from right to left, gives rise to negative variation of a->b', and
[b) Intermediate B, more excitable, shown
positive variation of a-^b.
by horizontal shading ; current of rest a^b and a'-^b ; action-current
on excitation at x from right to left, giving rise to positive variation
of a'->b and negative variation of a->b.
;
rest
Again,
we may have
The
or A.
118,
b).
Stimulation at x will
now
The
give rise to an excitatory current R, from right to left.
exactly
reverse
to
the
appear
be
however,
results here will,
of those in the last case that is to say, an identical current,
R, will give rise to a positive variation of A' -> B, and negative
Instances of these effects will be given
variation of A->B.
:
in
Chapter XVIII.
177
preparation.
current C
is
from the
in isolating
current c
is
-> A,
after-effect of excitation,
the specimen
reversed to B ->
by
section, the
such as
may
normal resting
still
->
of the current of
rest.
(a)
-^c
(b)
^^1
B
R-^
-^R
Typical Cases of Variation of Current of Rest and ActionAnisotropic organ, B end originally more excitable than A
Fig. 119.
Current.
(a)
R<
(C)
-<
response
(fig.
19, c).
how
intricate
the
resting
will
thus be seen
responsive variations
have
consisted
in
of
assigning
the
current,
positive
of absurdity
variation
the
to
negative
a
to
when we
find
COMPARATIVE ELECTRO-PHYSIOLOGY
178
Indeed
must be
it
said
that,
been found,
its
The
logical inquiry.
inquirer,
when
in the
way
of physio-
and mystical.
capricious
we
described,
find
the
phenomena of
have been
response,
however
no way
As
isotropic
injury,
tissue,
we must remember
excitation,
sign,
its
within
galvanometric negativity.
point
is
a naturally
limits,
is
one of
normal sign of response, from contraction to expansion, from negative to positive {cf. fig. 64)
The change at death, in which contractile rigor passes into
post-mortem relaxation, is analogous to this. Thus when one
end of the specimen is merely injured, that end becomes
rise to
a reversal of
more or
less
its
persistently
current flowing
away from
galvanometrically
it.
negative,
the
of galvanometric positivity.
In an isotropic tissue, then,
we may, by moderate
injury,
is
79
metrically positive,
which
is
analogous.
we have
a state of things
is
But
galvanometrically positive.
dis-
of the so-called
resting
is
On
diffuse stimulation
The
differential
excitability
or
anisotropy,
less
may
it
excited
is
A.
be either
isotropic tissue.
Numerous
illustra-
N3
CHAPTER XV
EFFECT OF TEMPERATURE ON ELECTRICAL RESPONSE
General observation of
effect of
temperature on plant
and Holly
Effect of
After-effects
rise
Effect of
rise of
and
fall
Enhanced response
effect in auto-
as after-effect of
of
temperature.
tissue
We
know
importance
condition.
be gauged by means of
further
in
this,
logical efficiency
is
is
electrical
response.
influence of temperature
the maintenance
There
favourable to
that the
its
is
of
of a proper physiological
If the plant
is
limits physio-
An
succumb
Certain
species
easily.
effect of
temperature
Desmodium gyrans.
fig.
l8l
and
shown the
The
ture.
effect
In
fig.
121
Fig. 120.
Desmodium
^jyrans
Normal
Specimen, and
in
it
is
temperature.
they
come
We
on the
Up-records represent
When
is
quickened, with
the temperature
is
raised
still
rise
of
higher,
to a stop altogether.
shall
As
effect of
temperature
COMPARATIVE ELECTRO-PHYSIOLOGY
82
electrical
appeared.
response of this plant to have practically disA few specimens were found nevertheless which
twenty-fifth.
On now
Fig. 121.
Photographic Record of Pulsations oi Destnodium during
Continuous Rise of Temperature from 30 C. to 39 C.
then placed
it
for
15
minutes
in
a cooling chamber at a
temperature of
C.
On now
83
response,
it
(fig.
122).
after-effect of
purpose
this
lowered
I
stem
the
(2)
of
(c)
(a)
chose
Ivy,
and
I took
(3) Holly.
their normal responses
17
an
ice-chamber
temperature
at
of
a
C.
The
for
24 hours.
specimens were
then
and their
responses under stimulation once
more retaken
out,
corded
these
From
(fig. 123).
will
it
that while
be
seen
the respon-
(b)
abolished,
of
hardier
the
that
is
revival.
interesting fact
I
that
Diminution of Response in
Eucharis by Lowering of Temperature
(a)
{b)
Normal response at 17 C.
The response almost disappears when
plant
subjected to - 2 C. for fifteen minutes.
Revival of response on warming to 20 C.
is
One
which
Fig. 122.
plants,
(<:)
have noticed
when a
plant approaches
its
death-point,
by reason
is
its
re-
is
is
due
to the
unmasking of the
positive,
This
by the
effect
abolition
COMPARATIVE ELECTRO-PHYSIOLOGY
84
We
shall
effect
on the
electrical response
The
plant-chamber to the
in
determinate
various
raising the
temperatures
great difficulty of
required.
was
able,
2i),
fig.
In the process
will.
ct
Holly
Ivy
Fig. 123.
a.
After-effect of
b,
Eucharis
later, acts as
a stimulus.
The
diffi-
moments of
the
way
a
obtained
the
petiole of Eucharis
effect of
following
lily,
temperature on
In this
affording a general
response.
It
will
be
idea of the
seen
that
85
while
On
fall,
hanced
(fig.
124).
The heating
ments
arrange-
were
this case
in
was made to
what rapidly.
some-
rise
be
It will
here
not
disappeared,
even at 65 C, though,
as
we
shall
see
the
in
is
at about 60 C.
anomaly
due to the fact that
the plant, which is a bad
conductor, was not allowed time fully to attain
This
apparent
is
temperature
the
We
surroundings.
see
that
perature
when
is
raised
about
minute and
the specimen
its
shall
the tem-
slower rate
in 1*5
of
at
1 C.
Fig. 124.
in
given below.
Temperature
rising
Temperature
65
(9)
51, (10)
when
I
give below a record of the effect of temperature
varying from 30 C. to 50 C. on the response of the stem
of
Amaranth
it
is
(fig.
125).
COMPARATIVE ELECTRO-PHYSIOLOGY
86
fatigue,
and
to
It will
it.
this plant is
In
this
case,
but
from
subject
little
continuously-
is
30 C. upwards.
the
C. in
1*5
time for the plant to attain the temperature of its surA tabular statement is given below, showing
roundings.
the effect of temperature on amplitude of response, in two
different
specimens oi Amaranth
Specimen
Speci MEN
I.
Height of response
Temperature
Temperature
II.
Height of response
1
30 c.
200 divisions
170
40
95
72
43
50
The same
,,
in
fall
no divisions
30 c.
35
40
45
50
'
90
40
25
10
From
muscle.
be
this
might
it
concluded that
rise
of
temperature beyond 30 C.
or so, induced depression of
But here we
by an anomaly.
excitability.
are
met
50 o
creases,
J
Fig.
Diminished
125.
Amplitude
of
35 C.
the
thermal
with
excitability,
the
is,
case
throughout
ascent,
optimum pomt
correlated
in
plant,
Up
tO
at or
of
the
an
near
to a certain extent,
phytmUy
for
at 30 C.
9*1
was
Thus
temperature.
mm.
a certain
in
8/
specimen of Bio-
that
is,
it
From
raised to 37 C.
in
became enhanced
to
these considerations,
would
it
an
that
increase
is
to say
galvanometric positivity.
we have
seen,
The
inference that
it is
by augmenting the
force of
increased
the
force
of
recovery,
we should
fall
be seen
in
fig.
126,
That
in the first
and con
part of which
is
given
COMPARATIVE ELECTRO-PHYSIOLOGY
88
siderably enhanced.
is
The diminution
its
original value.
may
Fig.
The
pulsations to the
the room, 29 C.
lowered to 25 C.
to cause antagonistic
metric positivity.
seen in
first
figs.
of these
in the
next chapter.
The
In the second
(fig.
133)
is
Amaranth
now
clear that
when
89
be
tissue
first
120).
(fig.
surrounding temperature,
its
If similarly a
plant
electrical
responses to suc-
When
is
force
factor.
tiveness
This
seen
is
its
appearance as
its after-effect.
in
fig.
124,
COMPARATIVE ELECTRO-PHYSIOLOGY
igO
When
the temperature
killed,
is
is
and
its
This
is
Before
first
demonstrated visually
record
(fig.
127).
in
In this
into the
After
Fig. 127.
response dis-
electrical
effect of
Steam
in abolishing
Response
The two
records to the
left
line
volt.
in
It will
be seen that at
first
But
was
and
in
five
plant
minutes the
this quickly disappeared,
effectively killed, as is shown in the waning and final aboliThis experiment affords us a qualitative
tion of response.
demonstration of the abolition of response at death under
a transitory augmentation ot excitability was induced.
191
we
of death.
It is
There
response of plants.
which response
is
is
abolished.
at this temperature
it
is
a temperature-minimum below
If the plant
apt to be killed.
which
is
highly susceptible of
response
is
per-
disappears
also
at a
certain
The
maximum
electrical
response
temperature con-
CHAPTER XVI
THE ELECTRICAL SPASM OF DEATH
symptoms Accurate methods for determination of deathpoint Determination of death-point by abolition or reversal of normal electrical response
Determination of death-point by mechanical death-spasm
From thermo-mechanical inversion By observation of electrical spasm (a) in
Different post-mortem
anisotropic organs
inversion
{b) in
radial organs
It
will
last
since above
it
life
been experienced
passes
in the
is
in
the case
which
is
critical,
into
death.
Much
has
difficulty
critical
no sure criterion of death was hitherto availsuch as would furnish an immediate and reliable indica-
point, because
able,
tion of
as
its
occurrence.
The
drooping, withering,
various
discoloration
and
escape
the
of
Even when a
later.
not
till
after
symptoms
To
it
some longer
may
and
it is
are seen.
make
itself visible.
symptom
on which
of drooping,
this
it
is
depends cannot
layers
we
moment
at
93
and
becomes marked, the temperature has already
passed several degrees above the fatal point. I have found,
for example, that the colour of the milk-white style of Datura
alba has changed to brown by the time that the temperature
after
death
by the time
is
it
The
by purple
normally at about 6% C.
In
all
finally, in
rings, loses
its
fila-
which
colour
Hence,
is
all
that the
symptom
But
mortis.
meant by
is
it is
the death-point.
In animals, an
does
not
synchronise
throughout the
change
earlier
than others.
it
as the point
sign of death.
shall
death-symptoms and the modes by which they may be detected with certainty.
With regard
where
to such detection,
that, theoretically,
a determination
it
should be possible to
else-
make such
effect
COMPARATIVE ELECTRO-PHYSIOLOGY
194
some effect which at the moment of death underwent a sudden reversal to its opposite. There would not
here be even that minor degree of uncertainty which is indetection of
a waning
effect.
And
by
my
effect,
we have
When
the plant
is
is
killed,
in
the
After this, a
second set of responses, to the same stimulus as before, was
In this way observations were made with each plant,
taken.
till
give below
(p.
From
these experiments
it
in these
should
be stated here that this investigation was carried out in the
winter season in England, and it will be shown later that
It
death-point by about 4 or 5 C.
I
method of
195
Temperature
Specimen
100 divisions
17 c.
53 C.
70
4
17 c.
53 C.
160
17 c.
50 C.
100
17 c.
55 C.
80
17 c.
40
60 C.
17 c.
55 C.
decrease of
0*7 volt
60
response, tending
towards
its
abolition,
it
with
under-
40^
50
58
ii
60"
Fig. 128.
The
at various temperatures
critical
temperature of 60 C.
went actual reversal from the normal galvanometrie negativity to positivity (fig. 1 28).
This was due to the fact that
02
COMPARATIVE ELECTRO-PHYSIOLOGY
196
on reaching the death-point, the contained positive component in response was unmasked by the abolition of the true
excitatory
But
effect.
It
this positive
be seen that by
will thus
method
this
is
the tissue
is
should
is
it
necessary,
thin, this
if
longer
But
fatal.
exposure to
is
to
it
be killed
effectively.
I
For
mechanical responses.
at a certain critical
moment
the
ot
obtaining the
symptoms
afforded
by
The experimental
plant
for instance
was placed in a bath of water, whose
temperature was being raised gradually, at a uniform rate of,
say, 1 C. per one minute and a half, until the death-point was
Mimosa^
reached.
of the
During
leaf, for,
a gradual
all this
we have
seen,
it is
fall
and not
a sudden variation,
rise
This gradual
stimulus.
rise,
a vigorous
Mimosa
the death-spasm
or very near 60 C.
is
found to occur at
is
followed
after
is
physiological
activity
also.
Thus,
53 C.
latter
value,
will
it
I97
56
filled
In
all
show
death-point
is
The
florets
characteristic
found to be at or near 60 C.
to the radial organs of ordinary plants,
Turning next
onset of death.
thermo-mechanical
specimen, while
it is
curve
is
recorded
by the
The
rise
of
ordinate
the case of
Mimosa
is
moment
elongation, up to the
When
this point
is
The expansion
described in
turning point
is
very abrupt.
The curve
as
This
a whole
is
filament of Passiflora.
here at
59*6
is
C.
(fig.
129).
very
COMPARATIVE ELECTRO-PHYSIOLOGY
198
similar
in
till
less
hesitation, as
occur in
55 C.
the
it
were,
record
seen to
is
or
at
near
The
first
or
down
part of the
Two
there
maybe
P'iG. 130.
in
other
is
60 C.
takes
When
place
at
the
usual
temperature of about
is
enfeebled, or has
namely,
it
may be mentioned
some
will
4 or 5 C.
it
lower
99
here
it
by
by as much
as 19 C.
I
moment
it
occurred to
me
that
it
by an
spasm.
electrical
It
may
be well
temperature raised
The
gradually
up
the death-point.
to
may now
be expected to
But
and similar at
and there will be
since
A and
little
they
B,
will
effects
are
marked
a
in
resultant effect,
In order to
we must have an
then,
are different.
which
equal
obtain
organ
excitatory
induced.
may
be either natural or
may
take a specimen
We
specimens described
find to
answer admirably
Any
of these
for the
purpose
of this investigation.
Taking the
and
tudinally,
divided
it
longi-
COMPARATIVE ELECTRO-PHYSIOLOGY
200
outer
convex.
Electric
contacts
be raised continuously.
This was
by an incandescent electrical
lamp which was placed in a second chamber, vertically below
the plant-chamber.
There was a wooden partition between
the two, by which the light of the lamp was excluded from
the specimen (fig. 131).
For the radiation itself will be
satisfactorily
accomplished
J-
^-1^=^
Fig. 131.
Electric
lamp
in the lower
E, e', electrodes
compartment
raises
the specimen
all
air
itself.
was enabled
included
the lamp-circuit
in
made
about
The
its
it
to
possible to
average being
per minute.
natural current through the petiole
circumstances, from
the
less
that
is
excitable
is,
under normal
outer to the
is
more
galvano-
201
This
metrically positive.
made
to section
gradual
rise
for the
of temperature, as
turgidity of the
of
we
Mimosa, and
increased
this
was exhibited
But the electrical
turgidity
that
is
responses
occur equally in
its
We
galvanometric positivity.
electrical
we
surface,
obtain, during
This
positivity.
the rise
is
marked by
posed
it
in
is
causing galvano-
necessary that
For
fluctuation.
reason
when
is
the rheostatic
For
this
resistance inter-
made
at the beginning
investigation
movement of
experienced
At the commencement
much trouble from the
galvanometer
the
spot
of
of
and
light,
my
erratic
the
almost hopeless.
the attempt
regulate
to
the
slide.
rise
of
It is for
for
at the beginning.
in this
manner,
obtained,
curve at
found, in
all
the death-point.
59'6
or
60
C.
rest,
to
occur accurately
the
same
COMPARATIVE ELECTRO-PHYSIOLOGY
202
to occur at
or about
And
55 C.
was lowered
to this degree.
I
give
below an
inversion at death
electrical curve
(fig.
It
132).
more
we have seen,
These responsive
variations were very large,
excitable, as
took
therefore
photographic
and 6;
54 C.
C. only.
The
increasing
galvanometric
tivity of the
surface
of
tivity
specimen.
temperature, had
Photographic Record
Sudden
previously,
posi-
P'iG. 132.
first
part of the
it
is
to
The
posirise
of
been going on
be understood,
menced.
metric
corresponds to
filament
of Passijlora,
variation of turgidity.
all
alike being
degree of temperature.
The
These interruptions,
As soon
made
curve
the
as the death-point
at
132),
(fig.
thermo-mechanical
curves
is
203
curve
(fig.
129).
Each of these
contraction.
which there
to
abolish
is little
differential excitability,
excitability of
the
by previous
it is
necessary
one
scalding.
lamina.
On
the
more excitable
increasingly
petiole
positive.
The
became
photo-
(fig.
took
55
C, and
place
^
at
133).
^^'^-
^ltr<Zh
another
means of detection of the death-point, afforded by the conversion of the response to external stimulus from the normal
be interesting,
be applied at
if in
the
COMPARATIVE ELECTRO-PHYSIOLOGY
204
lateral leaf
excitability of
by scalding.
The
electrical curve,
The
existing electro-motive
in
difference
40
50
1.
A
55
58
\/
Fig.
134.
by dotted
Stem of Amaranth
4.
increase, reaching a
maximum at
the death-point.
Meanwhile
mechanical
uniform
to
to pass
205
The
off.
stimulation
which
is
As has been
We
may
be mentioned here,
anticipation of a future
in
may
also be obtained
this
in
by the
and
electrical resistivity,
way
already given.
I give below a table showing the death-points
determined by various methods
:
Death-
59
59
59
Specimens
point
C.
I.
2.
Movement
of uncurling
Spasmodic
lateral
Morograph
J5
mens).
Each gave
9-12. Style of Hibiscus (four speci
mens).
Each gave
13-18. Coronal filament of Passiflorc
(six specimens). Each gavt;
Anisotropic organs
19. Petiole of Cauliflower
20. Scale of Uriclis bulb
21. Petiole of Musa
22. Radial
organ
petiole
movement
60
...
...
55
60
60
Electric inversion
5J
J5
J>
)J
55
}>
55
55
59-6o
60
60
60
Amaranth
23-
It
is
Petiole of Afnaranth
thus
seen
Resistivity variation
that
employing
different
and others
death-point
is
59-6
59-6
methods
60
and
flowers, bulbs,
determined which
COMPARATIVE ELECTRO-PHYSIOLOGY
206
is
gamous
plants.
which depends
Among
on
the
the
same
mechanical
phanero-
methods, that
thermo-mechanical curve,
is
Of an
by the
and
is
for all
is
giving
specially accurate.
also that
And
which
it is
in
CHAPTER
XVII
xMULTIPLE
Multiple
Biophytum
in
tissues
Continuity
of multiple 'and
pulsation of
Desmodhim
leaflet
under physical
restraint
We
single stimulus of
intensity,
sufficient
is
Growth-pulsation
acted on by a
exhibits a
it
single
to
effect,
find appropriate
expression
It
later.
response reaches a
limit.
It
may
The
incident
energy
in
such cases
is
held
latent for
rhythmic manner.
this
no
less
In
fig.
leaflets.
For more detailed account see Bose, Plant Response, pp. 279-357.
COMPARATIVE ELECTRO-PHYSIOLOGY
208
The average
may
is
seconds
but this
to one minute.
vary
in different cases
The
imperceptible volumetric
Fig. 135.
In
fig.
136
is
seen such
effects
by means of electro-motive
response.
In
given
by the
leaf
of Biophytum^ the
fig.
from
five
was
to eight
thirty seconds.
is
individual thermal
here
137
responses,
stimulus
gave
It will
rise
to
From
the corresponding
mechanical
it
will
to
by
vinules
by
animal,
the
are
tissue,
almost
all
also
the
by
exhibited
kinds
in
cardiac
of
tissues
show
which
of
this
in
the
vegetable
different
case
organs
In
lation.
r
fig.
139
is
given
responses
a series of multiple
^
^
have
obtained
which I
from
frog's
stomach.
have
^^^-
136-
Response
^^?
<;
Multiple
in
Electro-tactile
Stem
^^"S^^
of
Miviom,
Strong Thermal
Stimulus
(Original record reduced to ^.
also
Fig.
137.
We
COMPARATIVE ELECTRO-PHYSIOLOGY
2IO
In the
cases
stimulus.
Fig.
(a)
The
138.
may, however, be
Forms
of
increased,
and
multiple
response
may
consequently
be
temperature
the
This
is
seen
the
of
in
the
plant.
following
of pulsatory responses
record
as
young leaflet of
Biophytum^ when the temperature
was raised to 35 C. With the
induced
in
increase
of
internal
turgidity
of
the
energy,
the
was en-
tissue
induced
(fig.
immediate
pulsations
similar
pulsations
are
in-
autonomous
heart
Fig. 139. Photographic Record
of Multiple Electrical Response to Single Thermal
Shock in Frog's Stomach
brought about
sources
or
..
ir
itself in
1.
pulsatory
from
in
;
the tissue,
or from the
There
and autonomically
responding plants. Biophytuni, which under ordinary circumstances belongs to the former of these classes, becomes
converted into the latter under exceptionally favourable
tonic conditions.
That is to say, it responds by a single
response to a single moderate stimulus, and by multiple
excessive accumulation of such latent energy alone.
is
thus a
continuity between
Fig. 140.
at
multiple
tonic conditions,
however,
Under exceptionally
it
favour-
exhibits spontaneous
or
stimulus
When
energy,
is in
This
is
it
to a condition of
(fig.
autonomous response.
141) of the response
P3
COMPARATIVE ELECTRO-PHYSIOLOGY
212
The
was
leaflet
in a
quiescent
amplitude.
On
ex-
to the increasing
for
some
time.
It is
it
and these,
showed a staircase enhancement of
absorption of energy,
owing
is
may
Fig. 141.
shown by the
may
stimulus
give
responsive
either
responsive expression, or be
latent form
for
give
indications.
partly or
an
to
rise
An
wholly reserved
subsequent manifestation.
'
plant
or
animal
is
in
Inner stimuli
ex-
immediate
held
is
held latent in
21
when
this is
moderately high.
rapidly
having reached
maximum
depressed position,
it
There
its
From
somewhat
next a rather
slow
/^
/^
ft
M)
?1
This up-and-down
motion is in some cases
tion.
approximately
straight.
of the leaflet
twisted after
and
the
is
slightly
its
descent,
corresponding
more
In
circular.
view of certain
peculiarities
be ob-
to
y y y
Mn
response of Desmodium^
it is
time-relations
of
its
phases of
in
in
is
is
movements
stated elsewhere,
p. 5.
Fig.
142
COMPARATIVE ELECTRO-PHYSIOLOGY
214
photographic
gives
record
of a
autonomous
of
series
Desmodium
the
also
leaflet
during
movement
rate of
different phases
its
of
have
of
series
recording-
separated from
arm of the
lever
The
thread.
silk
current
coil is
the
in
primary
Ruhmkorfl"'s
broken at intervals of
Hence
seconds.
five
the other
by a long
succes-
represent
Fig. 143.
Spark-record of Single
Pulsation in Leaflet of Desmodium
seconds
terval
successive
dots
5 seconds.
(fig.
in
of the
in the
of
five
give here
obtained
manner, of a single
this
mechanical
pulsation
of
of Desmodium.
be
understood
It is
the
that
down-movement
leaflet.
These
tractions
movements
An
impulse
pause
is
are
lower and
of the
alternately.
after a
in
developed
in the
lower
each.
143) a record
leaflet
to
up-movement
intervals
movement
to
particular case
position
is
attained
which
7 mm.
there
is
is
half,
which
this excitation
a pause.
excitatory
gradually
The maximum
reached.
per second.
is
gives rise
is
in
The up-movement
much
This movement
21
slov/er rate.
by a
moderate excitatory contraction of the upper half of the
pulvinus.
1 give herewith a table showing the characteristic
rates
of
due to natural
is
movement
in
aided
recovery,
the different
pulsation.
Average
rate
Maximum
rate
'61
mm.
.7
Duration of pause
45 seconds
per second
'
,,
,,
,,
Total period
Average rate
-4
'5
Maximum
rate
Duration of pause
40 seconds
Up-movement
mm.
,,
70 seconds
per second
,,
,,
35 seconds
and
we observe
first
This
exhausted.
during the
first
is
seen
ten
brief
and quickly
seconds,
action, the
and up motions.
latter
gives
impulse
may
up-movement.
maximum
gauge
The former
=1-52.
We may
The
intensity
of
=1*4
the
occasions
the
the
downward
and a-half
up-movement.
down
COMPARATIVE ELECTRO-PHYSIOLOGY
2l6
The
down-movement
much
again
is
less
We
have seen that a single excitation has a single conWe have also seen the multiple
electrical responses corresponding to multiple excitations.
comitant electrical pulsation.
It
have any
I
electrical concomitant,
and
purpose
of what nature.
if so,
experiments
For
this
movement of
the
leaflet.
straight line.
was
reaction
at its
maximum.
which the
joints to
merely as indicating
made one
leaflet,
These
flags.
electrical connections
normal saline
The
now gave
there
solution,
and
electro-motive
rise to
correspond-
On
taking
was surprised
to find
in
circuit.
followed
In
large
principal
a given
case,
p. 299.
of this
peculiarity.
responses
electrical
it
was
photographic
a sensitive
film.
light,
The
The galvanometer
vertical
movement
stood
as
converted
into
spot of
horizontal
by
was
of the spot of
is
to be under-
reflection
from a
12.
The
be seen that the minor electrical wave took place while the
position
movement
positions.
It will
its
highest to
its
lowest
the
movements.
The
moments of
its
excitatory up and
following considerations
make
it
down
easy to
COMPARATIVE ELECTRO-PHYSIOLOGY
2l8
understand
why two
electrical
mechanical pulsation.
change
at a given
First,
waves
we know
correspond to one
an excitatory-
that
concomitant, an
its
Second,
And
change.
excitatory
lastly,
on the cessation of
excitation there
an elec-
is
trical recovery.
Now we
the spark-record
(fig.
143)
subjected to two
is
vinule
upper
and
of
pul-
the
in
lower
halves
alternately.
two
these
which give
electrical
And
Photographic Records of
Simultaneous Mechanical and Elec144.
trical
Pulsation of
a b (upper
Desmodium
figure) represents
of leaflet
sponding
a b (lower
Leaflet
up-movement
figure)
corre-
wave
down-movement
electrical subsidiary
is
rise to the
disturbances
two
of
negativity.
different
ampli-
by the
fully
accounted
different period
the
It
excitations
galvanometric
Fig.
its
excitatory
will
the
impulses.
It
be remembered that
impulse
excitatory
ing persistence.
On
is
we
we
find that
still
is
certain
is
complete,
galvanometric
residual
short-lived
The same
downward-movement
reaction
finds
electrical
negativity
there
is
tically complete.
it was found
up impulse was
down
approximately as
i'5
is
to
i.
And
to the
it
is
interesting to see
(fig.
electrical
waves
is
wave
is still
higher.
give
below two tables showing the absolute values of the electromotive variations in two different cases.
Table
Number
of observation
E.M.
Table
I.
E.M. variation.
wave Subsidiary wave
variation.
Principal
0014 volt
(XX)55 volt
0013
00051
Number
f
'
tt
r>.
II.
^^^
at
M. variation.
E.M.
variation.
Subsidiary
wave
-0024 volt
0016 volt
-0025
0015
,,
0014
00054
0025
0016
0015
00054
0025
0017
0016
,,
0026
COMPARATIVE ELECTRO-PHYSIOLOGY
220
ments
v^^ere
restraint
(p.
prevented
20).
electrical responses
modes of expression of
a single funda-
I first
Fig.
Restraint of Leaflet.
The
part of this record shows both mechanical and electrical pulsaIn the second part, leaflet was physically restrained, as seen
Electrical pulsation
in the mechanical record, becoming horizontal.
now seen to persist with even greater vigour than before.
first
tion.
the leaflet
(fig.
145).
leaflet
was
restrained, as seen in
But the lower record, which gives the electrical response, still shows the double electrical pulsation unimpeded.
Indeed, so far from the mechanical response having been
the cause of the electrical, we find that on its arrest, at
the latter becomes very
least in this particular case,
line.
much enhanced.
In
appears as
fact, it
the fundamental
if
energy.
We
give us a
on a
the
tissue,
mode
same
of
indication enables us
further
Q06.
f\
N^
005-
existence
the
strate
demon-
to
dof-
may
which
mechanical
pression
ex-
the
in
so-
autonomous
movements.
called
sent in
by
plants
all
IV_
001
means
of
highly magnified
given
by
shown
'^'
'
'
003nn
80
'
KA/
1
'
'
003-
the
Q02-
re-
801-
/V
/
the
have
20"
40"
eo"
8 0"
this to consist
Fig. 146.
I'"
)"
Crescograph.
_j^.-"''
is
of growth, and
cord
/\
603-
002-
multiple
minute
pulsatory movements.
Growth -responses
The
146,
growth responses
is
COMPARATIVE ELECTRO-PHYSIOLOGY
222
is
positive
its
positivity, if the
two occur
rapid succession.
In growth-
negativity.
to exhibit, galvano-
This consideration
may
in
the
seedling
of
Pisum sativum^
indifferent cotyledons.
are
negative, as
CHAPTER
XVIIl
RESPONSE OF LEAVES
of Burdon Sanderson on leaf- response in Z>w^ Leaf-andcurrents Their opposite variations under stimulus Similar leaf-and-
Observations
stalk
stalk currents
shown
True explanation
Opposite-
of these resting-
and
Ficus
lamina
of
religiosa
Dioftaa
Subsequent
results Experimental
arrangement with
Explanation of various
of
Musa
results.
II.
phenomena
in plants
'
As
it
has
'
in
the course
it
which the
and thus we
shall
of such cases as
offer
an explanation
Before doing
the principal
effects
COMPARATIVE ELECTRO-PHYSIOLOGY
224
These observations
by the
tonus; and
secondly, to
the induction
action of electro-
of variations of a
the lamina.
As
prolongation
the
Fig.
(a)
shall distinguish as
'
its
the
this
147.
the leaf-current
On
(fig.
{a)
147
'
).
But
That is
stalk-
RESPONSE OF LEAVES
No
225
offered,
directioned currents
of
rest,
anomalous
one case
or the apparently
result, that
in
As
seen
regards
that
176),
(p.
logically
peculiar
these
than either of
more
This
excitable.
of
we have
rest
an intermediate point be
if
excitable
less
currents
is
physio-
two terminal
the
from the
less to the
It is
point of junction
religiosa^ for
The
is
The
currents
here also,
of junction
the leaf-current
towards the
We
same
tip
of the
(fig.
147
leaf,
and
{b)).
may
currents
in
rest
inwards.
This
decumana
(fig.
147
{c)
that
the
effect
in
direction,
in
be the case
in the
and flow
leaf of Citrus
).
be reversed
will
find to
to
and
leaf
of stimulation
stalk,
is
of these
we must remember
to
give rise to a
excited.
true
If then
When
right
is
left.
the
lamina to the
in the case
of Dioncea^ induce a
(fig.
147 (^)).
The same
thing
is
seen
COMPARATIVE ELECTRO-PHYSIOLOGY
226
latter
phenomenon may be
we observe
a positive variation
of the
leaf-current,
(fig.
146
and a
This
{/)).
is
Another
from
left
there
now
to right
is
thus
By
reversed, being
now
algebraical summation,
147 (^)
occurs a negative variation of the stalk-current,
(fig.
).
leaf-current, in Dioncea
and
in Citrus.
which an investigator
it is
tion,
or the
left.
It is
now abundantly
is
is
one uni-
which flows from the more to the less excited portions of the
If there be already an existing current, the responsive
current is added to this algebraically, and induces, according
tissue.
Much
confusion, and
avoided,
if
many
namely
RESPONSE OF LEAVES
227
more
to the
Another
effect
petiole cut
left,
In Nature (vol. x.
p. 128),
phenomenon.
is
off, in
in the leaf
Thus the
'
electrical conditions
on opposite sides
other.'
He
by section of the
would serve to enhance the intensity of the leaf-current.
Taking the ordinary leaf of Ficus religiosa, I have myself
been able to obtain results precisely similar to those described
in Dioncea^ by making successive sections of the petiole, at
shorter and shorter distances from the point of junction.
The leaf-current at each section underwent an increment.
The parallelism of the two sets of effects will be seen from
gressive removal of the antagonistic element,
stalk,
Ficus Leaf.
Length of stalk
Galvanometric deflection
7 cm.
16 divisions
2*5
,,
36
)j
1-25
50
,,
50
>>
0-6
65
60
>
o'3
90
Length of stalk
cir.
Galvanometric deflection
40 divisions
Burdon Sanderson's suggested explanation that the sucdue to successive removals of the antagonistic element, by section, is
Q2
COMPARATIVE ELECTRO-PHYSIOLOGY
?28
He
quite untenable.
contrary,
due
the
to
increasing
excitatory
Similar results
sections themselves.
may be
on the
of
action
the
obtained, even
and nearer
to that of junction.
cut acts as
from the
summation of
it,
left
to the right.
Algebraical
this
of
is
in
As
the lamina.
the section
must become
And
lastly, in
that
if
two
same time
that the
would be increased,
were
the
in
diminished,
effect, as
if it
with
the
leaf- current,
and
similar
be studied in
the
extra-polar
We
were
will
(Chap.
if
same
detail,
effbctsr
induced
XXXIX.).
have already seen
that,
by means of induced
varia-
caused by section of the petiole, it is easy to obtain an unmistakable indication of the nature of the true excitatory
electrical
change.
acts
as a stimulus.
His
investi-
RESPONSE OF LEAVES
229
'
The
results
Fig. 148.
applied
stimulus
Electrical
on
distal
Fig. 149.
Parallel Experiment in
Sheathing Petiole of Musa
Thermal
more
internal surface, /,
than lower,
side
Upper or 'internal'
more excitable than lower.
led-ofif circuit.
surface
excitable
7n.
Burdon Sanderson's
fundamental
in fig.
148.
In
fig.
149
According to Burdon-
the
right
lobe of Dioncea
(fig.
current
150).
is
induced
This means, of
lobe has
become
this
first
phase,
when a
certain
interval
had elapsed, he
COMPARATIVE ELECTRO-PHYSIOLOGY
230
became
relatively negative
in, in
to the
'
of excitation also
made
itself evident.
'
'
150.
Time-marks
by
The
following
is
does
for
it
The
excita-
RESPONSE OF LEAVES
is
its
23
mode
other*
suddenness of
the
origin,
rapidity of
its
propagation,
and the
incidence,
its
is
identified
excitable
it
of animals.
tissues
of
the
have
process in the
nature of this
preliminary disturbance (to which alone the term excitatory variation ought to be applied,
we know
'
action-current
nothing.
The
'
alone being
it
of animal physio-
to
that
indicate
in
excitation,
excited
is
cells
such as
become
parts always
become negative
to unexcited.
its
The
ap-
explanation
in
motive
surfaces.'
which differed
from those referred to above. He had previously found that
usually speaking the upper surface of each lobe was negative
to the lower.
Later, however, he came to the conclusion
that in the leaf of DioncBa in its prime,' the upper surface
was positive to the under. On repeating his fundamental
experiment' moreover, with these vigorous leaves, he found
that instead of the pronounced positive response which he had
previously observed, he now obtained a short-lived positive
effect succeeded by a strong negative (fig. 151).
He was
unable to offer any definite explanation of this difference
between the two sets of results, but suggested that it migKt
arise, in some way, from changes of the resting-current.
certain results,
'
COMPARATIVE ELECTRO-PHYSIOLOGY
232
'
In the
leaf,
phenomena
those of the
are linked
by
indissoluble bands
excited
togrether
is
In
making
this statement,
'
by a negative
place
We
highly misleading.
For,
owing
this
supposition
in
is
fact
due to prepara-
may and
frequently does
The assumption
negative variation
Indeed,
it
is
its
prime
'
For
(upper
'
'
had
first
returned to positivity
its
'
{ibid. p. 447).
prime,'
Thus, though
and of the
'
modified
'
vol.
179, p. 446.
RESPONSE OF LEAVES
The
due, in
233
condition of the
the term
'
leaf,
modification
of the phenomenon.
explanation
response,
of
then,
this
still
'
in
is
any way
satisfactory
reversal
of
remains to be
found.
So much
experiment'
for
the 'fundamental
The
next
experi-
Burdon Sanderson
consists of a leaf
Fig. 153. Experimental Connections with Dionaa according
the
to
Second
by symmetrical
contacts on the under surfaces of
If now the
its two lobes (fig. 153).
right lobe was excited, by touching
which
is
led
off
Experimental Method
Burdon Sanderson
of
contact,
it
became
first
Fig.
154.
results
to the
give
'
rise
to
the
positive
response
COMPARATIVE ELECTRO-PHYSIOLOGY
234
positive
first
These
And
changes.
positive
latter
again, after
lastly,
using
experimental
the
No
theory
effects respectively,
presently describe,
shall
will
it
results
I
The fact
able.
which
all
nated
(p.
6i).
had been
elimi-
liability to
positive
plexity
arising
from
responding organ.
the
It is
effects,
dis-
differential
excitability
of
the
it is
positive effect.
The
various electrical
phenomena which
are possible in
by the greater
of the organ.
fall
of the
leaf,
contraction of the
RESPONSE OF LEAVES
235
upper half.
point,
we have an
then
response of the
This
leaf.
or
erectile
mechanical
positive
is
The corresponding
more
electrical response
more
this
will
excitable.
Between these two extremes lies that experiwhich stimulus is applied at some intermediate
point, the consequence of which is that the hydro-positive
wave, with its greater velocity, reaches the responding organ
earlier than true excitation, thus bringing about a prethe organ.
ment
in
citatory
response
electrical
followed
by
fall
would
wave of
be
is
in
we have found a
men,
prime,'
we
by
is
We may
great con-
distance
With such a
speci-
its
maximum
point, at the
positive
so great as to allow
diphasic,
this
imagine that
ductivity,
The corresponding
leaf
therefore
negative.
only possible
the
of the
But
negative.
if
pre-
we took a
at the same
would
fail
due to hydro-positive
way,
may
wave
and we should see
in
action.
other positive
alone
this
difference
COMPARATIVE ELECTRO-PHYSIOLOGY
^36
undergoes such
'
We
effect will
of true excitation
is
By means
of
stimulation.
Thus,
in
found that
now only
per second.
The diminution
that in
mm.
was about
per cent.
18
1*54
excitation
may
shall
at a
certain stage
It is thus easy
positive effect alone being then transmitted.
to explain how it was that in Burdon Sanderson's experi-
as
'
'
results arrived at
by Burdon Sanderson.
*
Plant Response,
p. 244.
RESPONSE OF LEAVES
I
shall
237
'
The more
leaf of Dioncea.
of the pulvinus of
mode
thermal
of stimulation.
momentary heating-current
passed through a thin platinum wire gave the thermal variation required,
form of stimulus.
distance of 16
stimulation
tissue
'
leaf of Dioncea
(fig.
excitatory negative
(fig.
155
151).
ip)
).
This again
a Dioncea leaf in
is
the
its
same
'prime'
Musa
the
led-off circuit
COMPARATIVE ELECTRO-PHYSIOLOGY
238
is
masked by
it,
(fig.
i55
(^))-
of the
This response
is,
as
will
above the
be seen,
di-
phasic, its
the
first
under-surface
representing
phase
is
its
clearly
hydro-positive or
due to the
indirect
earlier
effect
transmission
of the
from the
was supposed
of excitation,
stimulated
RESPONSE OF LEAVES
of
under-surface
may
surface
of
true
excitation,
initiated
above.
vertically
wave
the
239
may
or
not be helped
by the induction of
due to the transmission of the hydroThis view is supported by the fact that in a
positive effect.
corresponding experiment on an ordinary leaf, in which the
second contact was at a distance too great to allow of the
of any hydro-positive wave, the
effective transmission
positivity at the distal,
the
case
this
the
arrival
its
concomitant
negativity.
It
now
is
clear
among
that
the
various
results
of Dioncea,
true excitation at
phases which
there
all,
are
due to the
Burdon
Sanderson at starting fell into the error of wrongly identifying the true excitatory electrical effect with that which was
due to the hydro-positive effect, and vice versa. We have seen
is
hydro-positive
that there
is
effect.
We
also
seen
that
leaf of Dioncea^
excitable
have
In fact
the
it
latter that
intricacies
in the
240
COMPARATIVE ELECTRO-PHYSIOLOGY
shown
current which
rise
to an excitatory
response
of
arise,
the
leaf-current.
concave surface
not from
any
ot
specific
CHAPTER XIX
THE LEAF CONSIDERED AS AN ELECTRIC ORGAN
Electrical
organs in fishes
Typical
instances,
Torpedo
and Makptenirus
servations
effect,
Multiple
by
excitations Multiplication
pile-like
of terminal
electromotive
lily.
response in
plants,
is
it
possible
to
obtain an
Among
animal
organ of certain
trical
offers
unusual
fishes
to
difficulties
attempt to show
structures, there
the
is
investigator.
But
shall
in the
we
organ
arranged
a voltaic
in
is
as type,
series,
of numerous electrical
one over the other, like the plates
Each
pile.
consisting
plexus
of
nerve-fibres
arrangement
sity,
in
Though
is
small, yet in
COMPARATIVE ELECTRO-PHYSIOLOGY
242
number of
From
the
point
organs
electrical
containing
some of
of
general
in
these plates in
50,000.
modified
constitute
The
nerve-endings.
electrical
muscles,
known
fish
as
an exception to this rule, inasmuch as morphological evidence goes to prove that in its
case
than
glandular, rather
is
it
is
The
organs
in general, is that
it
It
that
tried to
of the dis-
direction
He
organ.
Thus
posterior^ surface.
in
the plates
aspect
the ventral
ventral or anterior, to
is
from the
In
The
axis.
tail-aspect,
Pacini's
all,
is
from
is
tail
If these
generalisation, as regards
the
-nervous
non
posterior
direction of discharge
it
some attempt might have been made to offer an explanaphenomena. Unfortunately, however, this is not
tion of the
so, since
Malepterurus presents a hitherto inexplicable exIn this fish, though the anterior or
nervous surface
is
head
The
that
is
in
Gymnotus
towards the
difficulties in the
way
very great.
Is the activity
these
alone,
fishes
phenomena
in
Or
Against
243
it
is
in
it
related to the
excited muscles
is
this
instance of Malepterurus, in
standpoint, the organ
is
which, from
is
the
a morphological
of opinion.
There
is
under a
of electrical responses.
We
this
that
if
organs
we could succeed
any
we should then be
in discovering
which
cases
so
much
showed
among
similar
vegetable
characteristics,
phenomenon
in
it
entering into an
electrical
plate,
nervous surface, in contradistinction to the jelly-like substance in which it is imbedded, forming the opposite, and
here indifferent surface of the plate.
is
closely imitated
Now
by many ordinary
this
arrangement
which the
leaves, in
I have discovered
bundles of plants (see Chap. XXXII.)
elements which are in every way analogous to the nerves of
I
in the fibro-vascular
R 3
244
COMPARATIVE ELECTRO-PHYSIOLOGY
animals.
plate of Torpedo^
we may take
is
to the electrical
example of
this
may
be found
tissue.
An
in
characterised
is
by
dry and leathery. Thus the nerve passing into an elecof Torpedo corresponds with the petiole attached
trical plate
two cases
alike,
it is
the
elements.
In
it
is,
the exceptional
as
we have
Malepterurus, on
the
other hand,
electrical element.
analogue
Dillenia
is
found
in
both of which
In point
of the electrical
Malepterurus respectively.
that
is
to say, whether,
on stimulation, the excitatory current in the type of vegeby the leaf of Pterospermum is or
not,
versely, whether
in
by the carpel of
is
organs,
two types
is
245
governed entirely by
The
is
many
In
modifications
In this
way
it
induced by
may
be said
of Bryophyllum
and of Coleus aromaticus that when vigorous, and
of the
calcineum,
in their
first
of these
direct stimulation
Under
shocks.
discharge
is
may
either
excitatory
is
Turning then to
first
type,
shall
The
records
(fig.
156)
whose direction
COMPARATIVE ELECTRO-PHYSIOLOGY
246
Of
importance
great
Du Bois-Reymond
out by
was the
on the
electrical
directions.
investigation
effects
of
currents
in
the
in
different
introduced
carried
induced
by
Du Bois-Reymond,
the terminology
in
homodromous, and
as
Polarisation -effects
he distinguishes as
'
'
as absolutely
A polarisation-current
negative.'
in
and
'
relatively positive/
negative'
tively
polarisation.
It
polarising-
homodromous
or
same
Fig. 156. Electrical Response
of Lamina oi Nymphcea alba
due to Transmitted Excitation from Petiole
Direction of responsive current
from anterior or lower to
posterior
and upper
of the effect
surface.
the
in
He
charge.
to the
dis-
will
and negative.
This
own
dia-
polarising-current.
In
the
upper
the
Reymond,
to
two
opposite
is
homodromous
current,
relative.
then,
is
de-
The
polarisation-effects.
galvanometer
The
positive
The heterodromous
resultant
polarisation,
current,
on
According to Du Bois-Reymond,
polarisation.
heterodromous shocks induce no relatively positive
On
part of figure).
Saxton's
from
down curve
lower
in
machine, he
positive
accounted
he
This
effect.
by
for
both
cancel
each
polarisations
tive
directions
other
positive
to
heterodromous
the
to
be
so
small
as
to be practically negligible
to
be
posi-
alone
effective.
Du Bois-Reymond
failed
excitation
phe-
these
in
What
calls
he
positive polarisation has been
shown by subsequent workers
Fig. 157.
due to
citation.
But
The
nomena.
to be
as to
in
how
to
the
question
polarising-currents
single-directioned
am
fully
and
by
Du Bois-Reymond
is here supposed
be from below to above,
i^
homodromous current | (upper half
of figure) is supposed to induce
natural discharge
to
rise
satisfactorily.
periments carried
The exout
on
leaves,
which
am
about
to
COMPARATIVE ELECTRO-PHYSIOLOGY
248
as that of Torpedo.
One
such
leaf,
When
that of Pterospermum.
in
is
one direction
in
that
is
to say,
Du Bois-Reymond
That
is
this
in Torpedo.
result
really
is
same
the
effect
found to occur
is
are used.
leaf of
thermal
shocks,
acting
SrrX:wLX;
Surfaces are Excited Simultaneously by Thermal Shock
,,
. i
T3
,
Resultant
responsive current^ from
been
^^^
fully
is
that
on
excitation
of
it is
electrical
electrical
result
is
These responsive
is
it
the more
or negative, but
itself.
established,
simultaneous
excitable.
more
is
to responsive currents
Photographic Records
Fig. 158.
rise
by the
The response
of electrical organs
may
in
general, then
its
more
more
we have
current which
to
is
form
now
of
clear
that
in
surfaces.
definite-directioned
homodromous
these currents
The
stimulus.
here seen
the
after-
polarising-currents, whether
it
two
Referring once
intensity
or heterodromous,
act
of
an
as
the
is
electrical
after-current
is
opposed by
to
excitatory
action
as
of the latter
such,
is
effect
yet
it is
actually
due
after-effect,
to ordinary
Du Bois-Reymond,
In
Torpedo^ for
the reverse
is
more efficient
Such diversity of results is probby the considerations to which I
have referred.
If
we
take, lor
is
in
which
and
posterior,
to be sent in a
Pfliiger's
at
break
Law
will
hold good.
We
at
sake of
COMPARATIVE ELECTRO-PHYSIOLOGY
250
simplicity, neglect
any
may
effects that
Under
homodromous
induction-
On
directions.
polarisation-current.
K,^^,
we
shall obtain
minus K^ on the
surface,
less
negative polarisation-effect
Under a heterodromous shock,
on the other hand, we shall have K, on the more excitable
anterior surface, minus
on the less excitable posterior, plus
Even
this,
Pfluger's
Law
more
reversals,
The
may
according to the
relative
excitatory
of the
acting
break.
It
factors
will
may
thus
If,
is
excited by currents
then,
we wish
to study
method of
is
excitation,
negative or counter-polarisation
may be
we must
see
that
are eliminated.
is
to say,
These ends
to symmetrical
The
25
similar, equal,
if
the
and opposite.
equal,
if
shocks be so symmetrical as to be interchangeable. Whichever may be the factor of excitation will then act equally on
both surfaces.
solely
two
by
The response,
now be determined
therefore, will
surfaces.
It
in
and in point of time-relations. An ordinary makeand-break Ruhmkorff's shock does not fulfil this condition, since
intensity
the break-shock
two.
is
defects are
opposite.
Such
reversal of current
tension of a spring
S,
by the electro-magnet
current
in
is
accomplished by means of a
The key R
rotating reversing-key.
E,
is
No.
In
the
L,
wound up
the electro-magnet
experimental leaf
is
against the
coil
I is
When
the
the alternating
case just
described
the
sequence of the
current
K4, this
sequence
Employing
may
this
be made
method,
COMPARATIVE ELECTRO-PHYSIOIX)GY
252
at
Fig.
A,
The
leaf.
159.
galvanometric
after-effect,
Experimental Arrangement
for
at
any
short
Rheotomic Observations
secondary, of the exciting induction-coil ; N,, N,^, non-polarisable elecmaking electrical contacts with posterior and anterior surfaces
of leaf.
trodes,
is
the key Kg
(fig.
159).
by the
We
striker B
have seen
impinging against
that,
owing
to
the
to the
employment of equi-alternating
induction-currents,
how-
we
ever,
253
by any
(fig.
by
stimulation,
of
and
mode
In this
or posterior.
equal
shocks,
opposite
as
given
is
less,
Neverthe-
thought
surface
either
to
well
it
to
under shocks,
which the
in
alternating currents
primary
in the
were
and then
circuit
right-left-right,
first
left-
right-left.
In
the
electrical
Gotch
Torpedo
of
the
maximum
organ
found
electromotive
change to be attained in
about '01 second after the
application of the excitatory
shock.
In
leaves, again,
the
maximum
effect
which
is
at-
160.
Records of Two Successive
Responses in Leaf of Bryophyllum
calycinum
under
Equi-alternating
Electrical Shocks
Fig.
tained
to
nature
of
depend
the
on
the
tissue,
and
exciting shock.
as
'2
second.
It
should
may be
be remembered that
as long
in the case
vigorous leaves of
a
stronger
Nymphcea
electrical
'2
second
alba,
stimulus,
(p.
51).
With very
the
maximum
effect
was
COMPARATIVE ELECTRO-PHYSIOLOGY
254
much
attained in a
shorter
observations
made on such
is to say, in about
showing the rheotomic
time that
03 second.
It
should be remembered
Galvanometric deflection
01 of
20 divisions
63
a second
03
o<i
,,
,,
07
17
15
,,
,,
20
,,
,,
,,
i!
.>
'.
that
these
The maximum
results.
electro- motive
change took
place,
This
stimulus.
curve
shows
it
will
be remembered, after
40).
This point
will
be
In the course
excitatory
the
It
Fig.
of
Nymphcza alba
lower
surfaces
is
not
maximum.
said
that the
for
definiteness of direction on
the physiological
the
in
has been
organ.
In
anisotropy of
those
leaves
differentiation of the
upper and
marked,
differential
strongly
the
excitability of the
is
undergo
liable to
255
under
reversal,
by previous
the
In
stimulation.
however, which
leaf
of
Ptei'ospernium,
Here the
very persistent.
which
surface,
leathery,
is
excitability
the
posterior
its
highly excitable.
is
generally
is
a series of
is
is slight,
surface, with
The
of
give in
fig.
162
It will
and exhibit
no signs of fatigue.
We have thus found a
vegetable organ whose reuniform,
very
practically
and
We
type.
its
shall
next
responsive peCU^
^
of the vegetable organ
the
study
^
liarities
Responses given
^%^'
by Leafr^^f^%f
01 Fterospermum suberifolium to Stimulus of Equi-altemat-
^^\
J
whose responses correspond
1
Minute
It
is
regarded as
Among
still
more
perfect
if
we take
Here the
pitcher of Nepenthe.
is
mined
also saw, be
its
be
we
is
whether,
responsive current
is
in
these
leat
taken as the
Or
the analogy
The next
cases
also,
jecting
them
to
found
COMPARATIVE ELECTRO-PHYSIOLOGY
256
this to
be the
case.
The
From
this
is
seen
the
>fuKLk.UMLL
An
Fig. 164.
Photographic Record
of Normal Responses given by
Pitcher o{ Nepenthe, under Equialternating Electric Shocks
Responsive
is
the tendency
to multiple response.
is invested with a
glistening
membrane,
while the upper
dry
and
more
The moisture observed
or concave is moist and glanduloid.
inside each scale is in fact exuded from this inner surface.
or less
On
subjecting
one of these
responsive current
is
it
is
is,
as in the
of the
electrical
organ of fishes
may
be exemplified,
in
by means of the
The bulb
may
surfaces
It
will
understood that
be
the
here face
left,
while the
Thus the
all
the glanduloid
non-glanduloid
aspect of
left
this
are
pile
The
ment
is
of
the
bulb.
We
When
middle.
an
electrical
this
hollow peduncle
connection being
or outer, connection.
An
On
is
cut across,
made with
equatorial belt
it
allows of
numerous intermediate
additive.
We
how
how
response
in
258
'
COMPARATIVE ELECTRO-PHVSIOLOGY
in the othpr
and
finally
how, by a
serial
arrangement, the
fishes,
is
known
evident.
The
light
to occur in
Further con-
CHAPTER XX
THE THEORY OF ELECTRIC ORGANS
Existing theories Their inadequacy
uni-directioned, to shocks
organs Similar
electric
results
Uni-directioned
Multiple
responses
of
electrical
organ
Multiple
of
Musa
of electric
responses
of
Biophytum.
One
the
phenomena of
electrical
organs
other
electro-motive
themor
reactions
is
is
the question as
specific
falls
observed
that
to
to say,
phenomena with
is
animal
for
tissue.
and against
From
have described,
it
which
is
common
to
all
alike, that
we
contemplated.
in greater detail.
Before doing
this,
however,
we
shall briefly
S2
COMPARATIVE ELECTRO-PHYSIOLOGY
260
Bell, for
'
example, thought
it
moment
of innervation,
become
As
against
this, it
of a current of
rest,
first
Du Bois-Reymond
if
the nerve-current
there
is
'
is
Instead
in E.M.F.
to disappear in the
only an inessential
rest,
is
and
P.D.
during
always homodromous
Even had
the action
variation
Du Bois-Reymond
of the electrical
organ,
not by the
negative
in
the
it
is
plates transformed
is
now
held to be
invalidated.
subject,
I
this is
'
meant an
after-current in the
same
By
direction as the
261
It is, in, fact a new name for that phenomenon which Du Bois-Reymond indicated as positive
Du Bois-Reymond had also shown
polarisation-current.'
that this particular effect was most markedly exhibited when
exciting current.
'
activity
functional
Under opposite
or 'livingness'
conditions, again,
it
'
'
It is difficult, therefore, to
new name
for these
phenomena.
see
Dr.
an important
The
may
may
blaze.
quickly
live
is
may
they
sluggishly
muscle at
contraction
rest
blazing
is
smoulder
smouldering
oxygen
slowly or they
live
or
suddenly
a muscle in
its
.'
some explosive
is
dis-
in
Within a given
physiological " range of strength of current the negative
kathodic must, equally with the positive anodic, be designated an " irritative "
*
after-current,
*'
p. 74.
I.
p. 448.
Bledetmsinn, /ec^ro-
COMPARATIVE ELECTRO-PHYSIOLOGY
262
when
there
no
is
possibility of
shall presently
show
that galvanothis,
take place
The
fact,
however, that the excitatory after-effects dethe death of the tissue, has led Dr.
scribed, disappear on
'
non-living matter.
this,
is,
If
living.'
it is
He
stance which
the
'
is
blaze-current'
rion-vitalistic,
or
physical,
Hence, as there
reactions.
is
known
direction
also reversed
is
the case
to physicists.
is
but
of the
it is
after-current
maintained,
quite different.
is,
logical
it
is
so thoroughly explored
263
mystical
The very
vital istic.
moment
the
it
was
direction
either
in
due
to
the
to
previous
in the
response
uni-directioned
the
that
chapter,
The response
electrical
differential
of any aniso-
excitable, the
A being
A being
of
ability
And
negative.
positivity of
slight, or
the
in
A and
alone
becomes
excitation
induces
negligible, B
third
negativity of
case,
In
B.
all
of
the
from B to
first case,
A from
A.
The
that of B
in the
galvanometric negativity of
the third case,
at
to
be found
in
that
resultant current
it
at
is
B.
in certain
B,
second case,
that of
made
up,
consists of the
;
and
Examples of the
in
effect
of these will
last
responsive current,
probable
same
result
could
the
for
the uni-directioned
the
it
A being zero
Chapter XXII.
These being the conditions, then,
that
is
the
it
induction of
appeared to
be
me
obtained with
specimen were so
COMPARATIVE ELECTRO-PHYSIOLOGY
264
potentiality
galvanometric
negativity under
was
it
of the structure
other
became galvanometrically
negative, on
Chapter
T.,
excitation.
excitation
one surface
and the
have already stated, in
positive,
Thus
brominated lead
negative.
is
we
A B,
middle
in the
as
C,
is
then,
clamped
at
If,
represented
(fig.
upper diagram
165), and stimulate
the
right-hand end
in the
by
say
B,
mechanical
vibration, a responsive
165.
I7j>per
zero.
which
to-
cfimnlafpH
Slimuiaieo,
Lower /^urehek
flow
will
,j^
tne
^^^^^ ^.u^
11
Vanometrically
11
tive.
t)e
posi-
The same
the
case
will
,
With
on stimulation.
A,
When
is
it
being antagonistic,
will
But
fig.
first case,
B'
thus
then stimulation of
away from
b' will
the right-hand
The
effects.
the positive
265
of lead, of
strip
On
it
mechanical
now be
will
strip,
responding A
molecular
(fig.
166).
It will
as
the
disturbance
on
which
excitation
depends is caused by a
non-electrical
form of
stimulus
the
electrical
obtained
tional
is
in
its
unexcep-
freedom
'
by
the
polarisation-
positive, of lower
Resultant
negative.
from lower to upper.
surface
response
factor.
we can here
effect of
If inorganic
strip.
obtain
after-effect.
sending
bodies
only the
That
is
to
say,
response, from
is
When
these
found to be long-enduring
266
give
below
this
is
photographic
167)
(fig.
strip
shock,
homodromous
surface.
accepted, then
167.
that
the responsive
4,
is
of the 'blaze-current'
be compelled to refer
shall
It
to the positively-excitable
strip to the
Fig.
we
the
prepared as described.
heterodromous
t or
of
carrying out
upper
records
actually obtained on
responsive after-effects
in
is
to be
this metallic
a living condition
and
Heterodromous ^ Induction-shocks
Since
it is
direction of induction-shock.
is
is
is
It is also clearly
among
certain fishes
ability.
its
last
other
things,
depends
on
The phenomenon
terms
necessary to
namely,
call in
is
its
of
the
so charac-
electrical
differential
organ
of
molecular excit-
the molecular.
It
is
therefore un-
vitalism, or assimilation
and
dissimilation.
26!
homodromous and heterodromous shocks, the counter-polarisation-current is superposed upon the excitatory effect For this reason we see,
In taking these responses to
in
homodromous
the
that
164,
fig.
response,
is
which
in
smaller than
We
excitatory current.
the
leaf-organ,
have seen,
when
that
in
our experiments on
negative
this
polarisation
is
become
effect,
give here
(fig.
equal.
168) a series
of responses
of the
pared
strip,
these
lead
pre-
under
of equi-
conditions
shocks,
alternating
in
resulting
cur-
surface
the
to
upper,
as
before.
Having
strated
in
demonvarious ways
thus
mental
which
condition,
Fig. 168.
sponses
Equi-alternating
to
in
under
excitable,
electrical
and
excitation,
observe
give
are
known
whether
definite
to be
they
also,
uni-directioned
responses.
The
subjecting
shocks,
this
excitability
of
the
direction being
is
less
given
COMPARATIVE ELECTRO-PHYSIOLOGY
268
in
fig.
169.
sheathing petiole of
Musa
also,
we
is
now
structure of
an
electrical
question
any
single element of
is
Any
merely that of anisotropy.
which is anisotropic is
tissue
mw
potentially an
electrical
element.
origin-
becomes physiologically
anisotropic when it happens to
assume the recumbent posture,
ally radial,
owing
Fig. 169.
Response of Pulvinus
of ^//w<7j-a to Equi-alternating
Electric Shocks
of environmental
Such
stimuli.
^^
U^^ elec1^^
when
an anisotropic organ,
.
trically
excited,
^jirectioned
giveS
responsive
one-
current
which
also,
on excitation, gives
is
The extraordinary
269
discharge.
As
regards the
to be
first
Du Bois-Reymond
of these,
the
condition,
a general
as
that,
natural
true
in
the
current flows
in
the opposite
is
similar to that
is
Torpedo^
its
resting-current
be opposite
direction
in
find,
in
which
that while
its
to
opposite to
is
With regard
to
this,
is
that
electrical reaction of
of the plate of
excitatory current
this
is
In agreement with
Pterospermum, the
leaf of
the
have
primary
rule,
We
found, however,
found
little
doubt that
the excitatory,
if
could
it
and
of
rest.
In a highly
rest
observed
has not, in
all
in
an
the
fact
Torpedo^ this
after-effect
For
rest.
we have
seen
which reverses
current
preparations of the
electrical
organ
by
excitable struc-
that
while
reversed, or
in
This view
it.
the
is
supported
organ-preparation
ingoing-current of rest
is
of
con-
it is negligible.
It must be borne
mind that the fish is spontaneously excitable, and also
that there must remain a certain residual effect from the
organ-discharges.
excitatory discharge.
resting-current.
This view of
the
organ-current,
of
COMPARATIVE ELECTRO-PHYSIOLOGV
270
Du
Bois-Reymond,
as, in
reality,
a persistent after-effect of
made
by Gotch,
that
considerable
is
it
known
is
to
in
be very persistent
is
is
but in
rapid, this
excised
preparations.
We may
observed
turn
now
This
Schonlein.
seen
is
rheotomic curve.
multiple
the
in
of
apices
its
we have
the
seen, in
the electrical
in
given in
40.
fig.
responses of
the
With regard
effect.
first
is
is
un-
it
response
response
is
it
may
These, as
we have
Indeed
first
this
fact
that
responsive current,
it
is
causing a
new
excitation of the
tissue,
serially
arranged
that
is
of these leaflets
sensitiveness
is
27
And
by kathode-make or anode-break.
tion
this
value
is
wave
which, during
initiated,
is
leaflets
in
its
progress, brings
After an
serial order.
wave
is
found to
On
This,
it
of the
is
waves, however,
the
And
serial depres-
will
have found
be seen,
is
minimum E.M.F.
leaflet.
it
It clearly
first
The phenomenon
is, as we have
and not confined to elecexhibited even by ordinary
tissues,
We
and we
have seen
shall
find
and glandular
This
is
tissues,
interesting
it
in
are
subsequent chapters
actually
induced
in
that
nervous
in
made up
neuro-glandular elements.
of either neuro-muscular or
CHAPTER XXI
DETERMINATION OF DIFFERENTIAL EXCITABILITY
UNDER ELECTRICAL STIMULATION
its
flexibility Drawbacks due to
in
and counter polarisation-current Difficulties
overcome by employment of equi-alternating electric shocks Methods of the
After-effect and Direct-effect
Experiment of Von Fleischl on response ot
nerve Complications arising from use of make and break shocks Rotating
reverser
Motor transformer Response of Musa to equi-alternating shocks
Abolition of this response by chloroform Response records of plagiotropic
Cucurbila and Eel Differential excitability of variegated leaves, demonstrated
by electric response.
Advantage of
electrical
stimulation,
We
in
seen in the same place also that the more excitable point
of uncertainty.
The
electrical
form of stimulation
is
characterised never-
theless
points
many
counterbalanced by
These
them.
to
qualities,
sum
For the
of a
re-
point
by an
number of changing
factors,
however, are
serious drawbacks.
and anode-break.
There
is
The
physiological
due
to polarisation.
depend on the
are
said,
sufficiently
is
on
acting
factors
on
B.
complicated
added to them,
in
sum
sum
minus the
of the
These elements of variation
A,
but
still
another,
by
already
we have not
Pfliiger, that
shown
as
of the
the
The com-
complicated by
the fact that the result depends on the intensity of the electro-
With
in a later chapter, is
force.
feeble
stimulus
understood
COMPARATIVE ELECTRO-PHYSIOLOGY
274
and leaves
it
at B
Here the
will
then
true excitatory
effect is
much
At these
place at
At an
points
A and
we
If,
again, instead of
A from
that at
B.
These
strictly
over, can
We
that
thus see
by the employment of
this
method
B,
to
which
less
we
excited A, the
by
We
is
results
merely
which
in
How
mechanical stimulus.
and how
perfect
may
consistent,
due
be rendered, will
have postponed
till
now
The
mode
usual alter-
would be entirely
unsuitable for delicate and crucial experiments first, because
the excitatory values of the slow make- and quick break-shocks
coil
are unequal
These defects
have
been
which
we
an
actuates
current
is
obtain
reversed
an
induction-coil.
When
the
primary
induction-current
due
to
magnetic
varia-
minus n. When
the primary current is re-reversed, from minus to plus^
we obtain an opposite induction-current, due to magnetic
variation, from minus n to plus n.
It will be seen that if
these reversals of the primary current are made with equal
tions of lines of force, from, say, plus n to
COMPARATIVE ELECTRO-PHYSIOLOGY
276
The manner
galvanometer-circuit.
be understood from
fig.
170.
points
A and B of the
to be determined.
tissue,
We
in
whose
spiral spring
Any existing
Fig.
170.
difference of potential, as
After-effect of
M,
electrical
is
c.
Under these
K, the galvanometer-circuit
ential excitation as
may be
taken
qf a
series
excitations, at intervals
of,
In this
B.
the
of
way
after-effects
The
say, a minute.
records
of
brief
direction of
which
in
the tissue
is
but
for
its
perfect
One
have to be overcome.
working, certain
of the
first
conditions
The importance of
shocks.
this
will
be understood on
Ruhmkorff's
coil,
when actuated by a
vibrating
hammer.
of polarisation-effects
and
is
may
this drift
effect.
may
when we have
intensity
absolutely equal
alternating
The
turbing element.
will
The
effects
difficulty
is
arising
Still
COMPARATIVE ELECTRO-PHYSIOLOGY
2/8
is
the
disturbing
factor
last
element of uncertainty
is
the experi-
in
The explanation
break-shock.
hitherto
been
regarded
being
authorities
much
of
this
matter
phenomenon has
of great
difficulty,
In these
as
Ruhmkorff's
coil
The two
equal excitability.
have at
{a) feeble
value of anode-break
persistent,
That
is
to
any
case,
whatever
may
of nerve,
results secured
shocks, that
photographically, the
record
a certain
this
difficulty
is
electric
also to
In
light tremulous, and thus to spoil the photographic imThis may, however, be overcome by making the
pression.
now
means employed
to
may be
have been able to do in several ways, and^
among others, by using a Rotating Reverser. This consists
of an ebonite disc, on the periphery of which there are strips
of metal of equal breadth, and separated from each other by
obtain equi-alternating shocks of any frequency that
This
desired.
The odd
equal
distances.
strips
(i,
3,
5,
other, with all the even strips, with the negative pole of the
battery.
The
current
is
led
ofl*
by a second
pair of brushes,
P,
of an induction coil
(fig.
i/i).
Let us
disc
is
primary
coil
now
When
next pair of
the
strips in
COMPARATIVE ELECTRO-PHYSIOLOGY
28o
with the even strip and the lower with the odd.
Thus the
equi-alternating
This
coil.
currents
in
will in
the
turn
induce
secondary, the
Fig.
Method
171.
primary coil, P,
Duration of stimulas, secondary coil in series with
by metronome, M.
tion determined
means of an
electrical
The
fifty
alternations of current in a
When
the metronome, M,
-5
shown
key
is
is
in the figure, is
included in the
'5
circuit.
metronome
The key
second's duration.
lating shock of
so adjusted as to complete
When
this
gives a stimuis
now opened
28
The
is
alternating current
is
led to the
supply from
the
primary of a
The motor
street
mains,
is
Fig.
172.
speed
its
Ruhm-
driven by an
is
being
effected
by
an electrolytic rheostat,
a
R.
As
the
dynamo
is
provided with
nating current
its
armature.
number of
If the
lation,
It is
thus
day after day to use the same intensity of stimuand thus to compare the relative excitabilities of
possible
comMrative electro-physiology
2^2
different tissues.
the alternating-current
resonating index,
I.
When
same
the
At
is
the motor,
now
is
resonator
first
steel spring
is
made
The movable
at
is
till
plate
is
difficulty has
further
to
be overcome
dynamo
of the alternating
closed,
is
When
the main-
by the
coil,
in
interposition of
has
now
then, the
primary
to perform.
dynamo
coil, P',
Ruhmkorff's
is
is
is
the key, K,
is
P,
of the
P.
There is thus no
dynamo, and the duration of the
closure
dynamo
fluctuation,
When
coil.
nating current
circuit
which
In
may mention
In this
minute.
The
elimination of
in
usual
all
preliminary
test
sources of disturbance
The
stimuli of equi-
the
may
successful
here be
made
It
may
be
There
coil.
is
we have
a suspended
Having thus
symmetry of the
the
differentially excitable
Musa,
is
by
null action,
tissue,
interposed, with
its
On now
upwards.
surface
tested,
applying excitation
by
equi-
And
(fig.
171) be reversed by
if
means of key
K.
this deflection
The
direction of
on the relative
of
excitabilities
the
coil, S,
non-polarisable
surface
(fig.
be reversed.
electrode
in
up
it
does,
if
The
shown induced
But
if
we exchange
polarisable electrodes,
it
will
it
will
now be
tivity.
which
will
of Musa.
In these experiments the existing current of rest may
be balanced previously by a potentiometer. But this is not
absolutely
necessary.
give
below a
series
of
records
to
I73)> in
Musa
be more excitable
responsive current
than
the
under
outer or convex.
The
form of electrical
stimulus, as we found to be the case under mechanical and
is
seen
this
COMPARATIVE ELECTRO-PHYSIOLOGY
284
the physiological
subjected
The
on
many
mode
of applica-
difficult
problems,
ability
tissue,
perfect
To how many
vestigations
distinct
can be
it
in
ease.
in-
suc-
As there
chapters.
to prevent the
nothing
is
two exploring
on any two
distant, of the
Photographic Record of ReFig. 173.
sponse of Petiole of Musa to Equialternating Electric Shocks, before
and after Application of Chloroform.
points,
however
same organism,
seen that
we have
here
it
is
only the
differential
excit-
specimen.
shall,
We
in illustration
of the ex-
treme delicacy of
differences as
this
we
shall select
We
have
is
rendered relatively
fatigued
by the con-
side.
We
upper to the more excitable lower side, the responsive current under mechanical stimulation was in the opposite direction
now
we
Using
Fig.
was cut
eel.
off,
colourless skin
lower.
found to
Photographic Record
Fig. 174.
of Responses of Plagiotropic
Stem of Cucurbita to Equialternating Electric
of the
ventral or
flow
from
the
now
upper
Fig. 175.
Electrical Responses of
Eel to Equi-alternating Electrical
Shocks
Shocks
Current of
response from
ventral
surface to dorsal.
of Cucurbita.
Electrical excitation
o^ Cucurbita.
In
fig.
175
is
in
the
illustration of this.
ing
Another investigation which I thought might be interesthad reference to the variegated colouring of certain
foliage leaves.
striking
example of
this
is
found
in the
COMPARATIVE ELECTRO-PHYSIOLOGY
286
barred
by longitudinal
tribution of colour
vigorous leaves.
is
streaks
found even
The
of
milk-white.
in the
which
is
This dis-
tion
my
disposal.
mode
of investiga-
On making
electrical
leaf,
showed that
pallidity
physiological condition.
CHAPTER XXII
RESPONSE OF ANIMAL AND VEGETAL SKINS
in
tor
differ-
for
essentially different
General
formula for
all
skin Phasic
frog's
stimulation
successive
Mimosa
variation
in
(a)
of
current
grape skin
of
rest
{d)
frog's
induced
skin
as
(c)
result
of
pulvinus of
Phasic variation in autonomous mechanical response of Desmodiujn gyrans Autonomous variation of current of rest True current of
rest in skin from outer to inner This may be reversed as an excitatory
Electrical
Normal
By
it
factors
It
is
briefest
summary
still
is
Some
have
COMPARATIVE ELECTRO-PHYSIOLOGY
288
employed
of
the
results
animal tissues
And
also.
this will,
in
the
case
of
reliability and simplicity which it is thus possible to introduce into the investigation as a whole.
With regard to the electrical effects in animal skin,
epithelium, and glands, the inquiry resolves itself into the
determination
of,
(i)
of
rest,
the
'
ingoing
(2)
was found by
'
that
inner.
is
Hermann
He
in
found a
cells.
is
From
by Rosenthal
mucous glands of the stomach the current is also
ingoing, it was assumed that the observed electro-motive
The
forces were due to the glandular nature of the tissues.
and
of
the
fish,
the
glandular
and
skin current of the frog
that in the
the
same
There
inasmuch
cause.
is,
as,
in
this
view,
the frog's skin induces, a positive change, a similar stimulation of an unmistakably glandular surface is found to bring
about a negative. If then the electrical effect on the skin
of frog be the same as on a glandular surface, the dis-
have been recorded when stimulus has been applied indirectlythat is to say, through the nerve. This fact is not to
be wondered at, since the responsive effects are subject, as will
be shown, to numerous modifying influences. It is generally
made
is
it
for these
experiments,
electro-motive.
is
289
shall
is
And,
surfaces.
addition
in
to
these,
excitations of the
particular
still
two surfaces
of
distribution
is
the two
further compliindirect
that
is
will
be determined by the
Again, we
nerve-endings.
the
shall
is
see that,
to
liable
effect is
But
very
much
lowered, and
reversed to positivity.
For
all
these reasons,
if
we wish
to study the
specific
the current of
rest.
was 'outgoing.'
eel,
'
ingoing,'
meant that
by
single induction
Biedermann,
in the mucous
membranes of the tongue and stomach, obtained both
positive and negative variations of the current of rest.
shocks in either
Waller, using
direction.
single
induction
in
results
They would
to
direct
be
in-
COxMPARATIVE ELECTRO-PHYSIOLOGY
290
As
employed.
cally be
Engelmann and
others
theoreti-
used heated
we have
seen,
is
how
the
excitatory
these difficulties
shocks
in
effect.
may
have
already explained
is
made
to act
And
291
to,
if
be quite
different,
the
In
intervening tissue is great or slight.
they will consist of the transmitted effect of true excitation
in the latter of the indirect effect,
whose
electrical sign is
Fig.
176.
perfect
But for
method
method of
has
Chapter III., the stimulus being vibrational.
investigations on limp structures, such as skin, this
been described
is
in
now about
to describe
is
am
platform, P
(fig.
u 2
COMPARATIVE ELECTRO-PHYSIOLOGY
292
soaked
normal
in
When
saline.
the electrode E
the point A.
may
excitation in the
excitations
first
may
same way.
isolated
is
of
it is
necessary
or B should be stimulated,
For
slip.
in
reason
this
The extreme
bases
The
section.
is
tightly fixed,
The
is
is
collars,
by means
latter, again,
fixed.
The
inner
up or down movement.
pulling which rotation
A
is
string
is
also
produced.
specimen.
It
will
The next
to
variation
difficulty to
in
the
is
be overcome
pressure
of the
prevented.
is
that of liability
of the contact.
steel.
It
will
be
further held
elastic piece
and
in
time of execution.
The
number
intensity of stimulus
might
one end of the string wound round the inner brass tube
is
by
its
string
of tension.
When
is
fixed
of the
tasselled, after
is
293
is
its
other end
the tassel
is
is
ah'eady in a state
now suddenly
it
gives rise to a
rise to
the
same excitatory
reaction.
through a distance of
cm.,
and
let
by
negative
direction.
second knot
five rotations
in
be
five
in the
distance of
the pull
The
the
distance
To
approximately constant.
nullify
any accidental
galvanometer
When
circuit.
the
is
variation,
interposed in the
it is possible to use
an external resistance as high as one million ohms. It should,
however, be remembered that even if there be any unavoid-
is
discrimination of sign
response.
very great,
it
will not in
of the
any way
characteristic
affect the
electro-motive
may
The
repetition.
COMPARATIVE ELECTRO-PHYSIOLOGY
294
when
it
human
skin.
outer surface to
the
said,
from the
is
is
from
inner to outer.
skin,
way he found
In this
by passing induction
first
relation to
The
When
an indifferent
an induction shock
is
passed across
is
thus,
upper surface.
blaze currents
'
to electrical shocks.
The
by
Why
but even
should the
'
Vegetable protoplasm
accumulation,
in
shall,
tions from
295
which
it
will
surfaces.
We
then, say a
which
is
more
exposed
cells
walled, with
generally
little
become reduced
protoplasmic contents.
epidermal
much
their excitability to
degraded.
in size,
cells are in
We
same
and thick-
Hence, as regards
a physio-
be proportionately lowered
much
cuticularised
in
comparison
tube, protected as
rise to
protoplasm
is
in
'
glandular.
COMPARATIVE ELECTRO-PHYSIOLOGY
296
The degree
minating as
this
the extent of
its
may
diminution
of this
of
depend upon
In connection
transformation.
with this
it
two
surfaces,
necessary
is
it
For,
the stimulus
if
causing
thus
complications
The
changes.
the
in
intensity of stimulus
used
resultant
on
excitatory
may be
which
without
bringing
such complications
t
always a
is
will
conductivity
the
safely
about
depend
of the
Epidermal
tissue.
cells
are,
ductors, but
-\
in
this
matter
it
FiCx.
Diagram Representing
177.
Diagram
how
at B.
differences
in
this respect
tween different
absolute,
but
degree, and
may
question
to
circumstances.
of
a certain
be-
dif-
Thus a
conductors.
we
typical reactions.
possibility of the
left
of
fig.
excitabilities
177.
following
be repre-
its
become absolutely
The response
line.
by
B,
its
response will
line.
will
to say,
When
is
positive, as represented
IgJ
the
ingoing
'
'
'
outgoing
current
'
at
current at A,
B and
+ A, the
the
direction
if
by the diagram
The
that
is
the
less
to the right of
fig.
177.
then be represented as B^
A^,
We
negative A.
have thus
more negative B to
examined the two
more
show the
shall
a.
be recalled here,
taking vegetable
types,
skins
as
the
experimental specimen.
will
which
in
It
'
however,
am
latter
'
outgoing.'
justified, will
That
be seen from
the
Rotary
COMPARATIVE ELECTRO-PHYSIOLOGY
298
no compHcation due
when
is
by the
to
There
other.
is
here, moreover,
excitation.
response
if
This conclusion
is
tests.
Thus,
method
already
described.
shocks, according to
Again,
(3)
the
And,
finally, (4)
shocks
of
of
employed.
moderate
obtained by
Method
is
the
all
intensity
these
diverse
is
recorded.
methods
are
The
results
complete
in
concordance with each other, and fully support the theoretical inferences which have already been made.
I took the skin of a ripe
muscatel grape, such as are
On making
available in Calcutta.
was found
inner, just as
fig.
178).
to
flow
in
the
The grape
stimulating apparatus,
in
the
diagram,
now mounted
in
the rotary
The
distance between
the
same
namely, 2
On now
way
In this
299
It
galvanometric negativity.
give here
178, a) records
(fig.
B.
of the
tation
outer and
inner surfaces,
brought about
is
As
ance of the
circuit in the
maintained
same,
the
amplitude of these
gives
accurate
idea
re-
fairly
the
of
The
Fig. 178.
positive or
(a) A,
surface
is
B,
here
ingoing
'
effect.
JT
V.
faces.
the resist-
approxi-
mately the
sponses
added
The
fig.
179
it
will
is
negative, or
much
now
frog.
From
positive, or
'
same apparatus
frog,
ingoing.'
'
outgoing,' that
The amplitude
latter.
3^
Comparative ELEctRO-pkYsiotoGV
skin
alone
as
by supposing
for
active,
may
effective.
was not
that he used
possibly be accounted
my own
In
strong.
sufficiently
thinking, in-
experiments
opposite ways
in
though the
electrically,
effect
By comparing
of the two.
will
it
any way
Fig.
179.
Electrical
(a) A, positive
How
excitation
widely applicable
by rotary stimulus
is
the
be seen
will
is
quite unjus-
method of mechanical
in
an attempt,
successfully carried out, to determine the very difficult question of the characteristic response of the intact
This
will
be seen
in
the following
human
skin.
record
of the results
The
responsive elec-
The
responses
now
given, under
surfaces
of
stirriulation
grape-skin.
by
thermal
'
up 'that
is
fig.
301
after-effect of equi-alternating
current being
Fig. 180.
Photographic Record
of Electrical Responses of
Upper Surface of Intact
Human Forefinger to Rotary
Mechanical Stimulation.
Photographic
181.
Electrical Responses of
Fig.
.
Record
of
Grape-skin
to Thermal Shocks at Intervals of a
Minute
Down
is due solely to the differential excitatwo opposite surfaces. This is not complicated
in any way by the factor of polarisation, although the latter
could not have failed to be present if the exciting shocks
had been one-directioned. Under the conditions of these
equi-alternating
seen, in
shocks,
tremely characteristic.
when a
tissue
tion, its
certain
effect
is
is
often
is
ex-
We
Thus from a
one of
then,
maximum
it
may
pass into
COMPARATIVE ELECTRO-PHYSIOLOGV
302
changes,
record.
shown
the
in
fn fig.
182
very noticeable.
differential
this feature
We
here ob-
applied,
of
As
to the outer.
the
the
series
existing
of
the
stimuli
current
of
Fig. 182.
Photographic Record of Electrical Responses
of Grape-skin to Stimulation
by Equi-alternating
Electrical
tervals of a
Shocks
Minute
at
In-
to outer.
Note periodic
variation of resting-current,
causing shifting of base-line,
down and
rest
Fig.
up.
If this
had remained
In the
underwent an increase
then a
by another increase. Thus,
in the course of about ten minutes, it exhibited an alternation of almost one whole cycle.
In the next figure (fig. 1 83) I give a series of results obtained
to
inside.
decrease
This,
at
first,
to be followed, later,
303
Here we
the
find
usual
'
up
responses, showing
'
to without
that, as
from within
and here also we see the existing current of rest
is
two
now been
has
It
of skin
fully
is
its
surfaces,
greater.
upper,
is
lower, has
ability of the
been
fully
shown,
in a
previous
by experiments on
the pulvinus of Mimosa.
chapter,
long series
given
of
responses
by the last-named
periodic variation
resting-current
the
similar
to
and of
connections
FiG. 184.
Electrical
frog.
made
were
The
is
points on
lower
the
surfaces
upper and
of
this
to equi-alternating shocks.
the tissues,
surfaces
was
upper
And
excited.
record
it
merely, but
seen
responses
in
184,
fig.
of skin
is
the
general
In the particular
resemblance to
sufficiently obvious.
is
The
the
interesting
COMPARATIVE ELPXTRO-PHYSIOLOGY
304
of
stimulus,
cyclic
of
variation
conditions
relative
is
shown
in the resting-current,
have been
base
line.
able, further, to
even
have
already
ex-
that
of such pulsations,
to
similar
occur,
response of skin
to
obtained
or pulvinus.
pulvinus
half of the
If,
with
long-continued
for
of the lateral
will
be correspondingly
shown
variations
that
the
relative
anisotropic
organ
in
the record
turgidities
of
may undergo
tilted.
(fig.
the
two surfaces
a periodic
corresponding
the variation
of the current of
rest.
of an
change.
we have
The
seen in
This variation
may
Mimosa
is
from
the upper half to the lower, across the organ, this normal
direction
It
to be reversed.
it,
in the case
of the skin,
which determines the respective directions of the restingWe have seen that
current and the current of response?
frog's
skin,
from
outer to inner, is
in
the
rest
the current of
generally attributed to the possession of glands by the outer,
'
305
that
to,
an apparently similar
stomach.
Against
may
this
'
ingoing
frog's
which Hermann drew attention, that the skin glands are normally nearly closed to the external surface, and cannot therefore
other arguments.
First,
Second, the
specific
is
is
are,
moreover,
observed in the
no special glandular
response of
6 P.M.
There
relation.
a similar current
layer.
glandular surface
is
12
9 P.M.
-.^---^A/U^
A/VV^V^
i#^A..a#AtaM<Av^'^^'^
"n^'
6 A.M.
3 A.M.
Fni. 185.
The lower
definite,
record
and
is
is
by galvanometric
Note phasic
variation.
is
by galvanometric
of stomach
is
on
And, thirdly, we
the mucous membrane
positivity.
observed
in
rest,
to the
more
excitable,
opposite direction.
in the
COMPARATIVE ELECTRO-PHYStOLOGV
3o6
histological
modifications,
The epidermal
contents, and
may
may
layers have
be transformed
corneated or cuticularised.
tion
outer surface
the
excitability.
reduced
in
protoplasmic
ways, becoming
in various
The extent
is
little
of such transforma-
will as
tissue.
rest
if its
of the
slight, or will
It is
to be
specimen
will
be
remembered
is
that as
the
more
its after-effect,
Fig, 186.
Photographic Record
of Electrical Responses in Skin
of Neck of Tortoise to Stimulus
of Equi-alternating Electrical
Shocks at Intervals of One
Minute
The
proving in certain
cases persistent,
may
give rise
rest,
may
versed,
to outer.
owing
to the excitatory
after-effect of preparation.
we have
The most
is
liable
to
The
autonomous periodic
seen.
excitabilities of
two
excitable
outer, even
307
those
in
after-effect of
had
preparation.
is
As an
after-effect of
was
current
was stated
It
(fig.
86).
general
that
fact
the
is
And
the outer.
still
remain
this will
true,
even
is
galvanometriC response
Lower
Mechanical Stimulus
The
xiit
tivitv
\.i\ii.y.
f-vnpn*
c^ptu-
Ihe
in
^""^
natural
current of rest
inner,
seen
(fig.
187)
that
both
the
alike
surfaces
give
the
much
the
The
resultant
difference in
COMPARATIVE ELECTRO-PHYSIOLOGY
3o8
owing to
outer,
the
greater
from
still
is
inner
excitatory reaction
of
to
the
inner.
That the
fig.
i88.
The
is
actually
is
The
rently
record
by the
peculiarities
shows
negative
followed
responses,
positive after-effect.
appa-
(fig.
figure
189).
will
it
From
this
be seen that
of
mwr
After
sided.
proceeded
back
Photographic Record of
Series of Responses in Skin of
Tomato under Equi-alternating
Electrical Shocks applied at Intervals of
One Minute
reaching
to
records,
in
the
positive
direction,
Fig. 188.
now
In
zero.
the
similar
occurrence
of
and posi-
been
as
affording
visible
and
were to be regarded as indicating the
predominance, now of the one process, and then of the other.
That in the present case such an assumption is unwarranted
of a short-lived balance, while the subsequent negative
positive variations
we see
followed
309
process, such as
The
difference
their time-relations.
On
The
first
half
of the effect on
Fig.
189.
the
inner,
gives
rise
to
the
apparently
after-effect.
Thus, here the supposed tug-of-war
between two opposite processes of assimilation and dis-
positive
similation
is,
in
having different
reality,
between
time-relations.
It
two
is
normal
from a
responses
failure
is
to
not con-
occasioned.
COMPARATIVE ELECTRO-PHYSIOLOGY
3IO
The
result
which
have described
namely,
a greater
that
which occurs
in
is
with tomato.
may mention
few instances
in a
inner surface on
the
outer,
excitation
stimulus,
exhibited
a feeble
positivity.
Fig.
190.
But
will
An
abnormal positive
will thus
3 II
Its sciatic
Thus
provides
much
for
electro-physiological
investigation,
it
Fig. 191.
which was in a favourable tonic condition, I obtained the series of normal responses
to equi-alternating shocks, which is given in fig. 190.
The
skin,
favourable tonic
(fig.
period of tetanisation
response.
is
191) to be diphasic.
An
less
its
intervening
normal
'
CHAPTER
XXIII
and
epithelial,
membranes
secreting
tissues Natural
plant
in
Current of
Response of
snail The so-
response from
Dillenia
called
epithelial
Response
of water-melon
of rest
current
Misinterpretation
or secretory
arising
from
to
epidermal surfaces
Response
glandular
of foot of
surface
really
'
due
injury-
to
positive
variation
Natural
maximal stimulation
intensities
Differential excitations
Records ex-
of the
epidermis
similar
parallelism
the
glands
in
we
clear
If
we take
one made
other.
Uriclis
lily,
and,
is
The
little
may
If
now we examine
(fig.
192).
this inner
internal
very
differentiated
The
internal
mem-
its
cells
membrane
membrane continuously
313
from the top to the bottom, at the point where the peduncle
rises
ng (glandular
?)
layer
By
lower.
this,
secretion
is
filling
In
the
at
secretion
day.
and
walled
excessively
turgid,
extremely
are
simplicity,
interesting.
When we come
to
test the
we
find,
on making
nections with
the
electrical con-
external
epi-
tissue
to the internal.
cate
that
the
Transverse Section
192.
tissue of Hollow Peduncle
Fig.
was
internal
of
of
Uriclis Lily
the
This
confirmed
on the
application of simultaneous excitation to outer and inner
for the direction of the responsive current is found to be from
conclusion
is
and secretory
highly excitable
epidermal.
made
From
secreting
these
is
On
its
layer to the
experiments
we
less
excitable
see
that
the
COMPARATIVE ELECTRO-PHYSIOLOGY
314
epidermal
cells are,
and the
least,
of these secretory
Drosera.
cells are to
shall first
This is exemplified by
a single unripe carpel of Dillenia indica, already referred to.
When
this
carefully
is
connections are
must be borne
This
made with
in
mind
is
is
of
the
filled
with
inside
found to be
It
to
relatively
the
secreting
is
is
From
thus,
the
most
lily,
at,
excitable.
first
is
in
is in
experiment
The
definite direction
of the
it
did,
A similar
RESPONSE
.OF
was obtained as an
effect
shocks.
after-effect
of
The
nating shocks.
315
equi-alternating
effect of equi-alter-
was
In the
whose
fruit
we
of water-melon
interior cavity
is
filled
with secretion.
On making
outer epidermal.
Fig.
these responses.
From
vegetable
the
the
stimulus depends on
response
characteristic
of
When
thelium
tissues,
came
.1
^
iU
to the conclusion that
.
1.
'i.
it
Photographic- Record ot
Responses of Water-melon to
Equi-alternating Electric Shocks
Fig. 193.
Responsive
papillae.
One complicating
by the process of
isolation.
such stimulation
may
This
is
experiment carried
COMPARATIVE ELECTRO-rHYSIOLOGY
3l6
Waller,^
Dr.
the so-called
In
excitation.
tory
current of injury
'
the
'
case
reaction
we should have
The injury-current
case
this
happens, as
surface.
here, then,
the
is
necessarily
was obtained
in
the case
excitable
of the less
induced
from
becoming
side,
of
by injury
internally
diffused,
greater
citable distal
point
(p. 162).
is
connections,
we observe
On
a current of
allowing
electrical
rest, so-called,
This
which
is
not
galvanometric negativity at
the
An
independent experiment,
in
will
be described presently.
On now
citing
the
two
surfaces,
responsive current
is
The
consequent on section.
responsive current
'
simultaneously ex-
surface,
^.
loi.
positive variation of
From
current
of
current
the so-called
as
direction
rest,
317
constituting
it.
these experiments
is
it
is
the
vanometric
negativity
surface.
It
at
fixing our
If,
instead of this,
we had regarded
it
from
we must have
positive variation
interpreted
as apparently an
it
abnormal
^
;
such
sufficiently
may
recognising these
be originated
in different
condition
is,
may
as
distinct ways,
and that
This originates
current.
two
The
manners.
the resting-current
facts, that
in either of
in the natural
of ex-
differences
constitutes
because
positive,
current
injury,
a negative variation of
the
more
has
now become
which
of rest
may
such as
paration.
really, as
tent.
'
We
is
it
is
negative.
due
to
this.
point,
previous
excitation,
or
This current,
resting-current,
is
excitable
though
usually
regarded
as
rest.
become
the
It
persis-
p.
417.
COMPARATIVE ELECTRO-PHYSIOLOGY
negativity of the
same
in the
more
a positive variation of
at
now give
excitable, will
It is
it.
induced
its
direction
great
example of
Indian
this
garden-snail.
found
in
depression
of
may undergo
An
current
rise to a
originally
(p. 177).
occurred
was induced, with consequent great depression of exHence the responsive current was now found
citability.
fatigue
to
be
It
having,
reversed,
surface,
with
reference
to
the
glandular
ingoing.
rest,
This
was able
to verify
The
This
snail
surface, in the
in
normal
electrical
middle of
saline.
connection with
The
absolute electro-motive
had
condition, in
original
volt to
The induced
negative.
'0020 volt.
It will
from the
variation
was from
thus be seen that any
319
-0013
irritation
liable to
different
real.
proved indeed by
is
tissues, it is therefore
visable,
numerous experimental
case confronted,
may
difficulties
with which
we
ad-
The
are in that
How
practicable this
ments which
human
subjects.
We
likely,
is
partly
also to
its
imbedded glands,
richer
it
possession
appeared to
of
me
say, the
same
shoulder.
Fig,
194.
Photographic
Record of Electrical
Responses of Intact
Human Armpit
Responsive current from
armpit to shoulder.
Equi-alternating shocks of
one second's duration were applied at intervals of one minute,
and the direct effect photographically recorded.
The resulting responses were found to be ingoing as regards the
armpit, thus proving that that surface was the more excitable.
COMPARATIVE ELECTRO-PMYSIOLOGY
320
In order next to
are
relatively
show that
human
animal
we have
as
performed
Here
lip.
it
was
steady.
made, as seen
in
light
spring- contact
fig.
The lower
195.
Over
this
were tied
thicknesses
of
blotting-
paper soaked
in
On
solution.
zinc sulphate
were
again
this
soaked
zinc
in
spectively.
NaCL
lip
as
shown
in the
upper
figure, in
Fig. 195.
Lip.
Lower
made
Of
the
also
now
placed in the
The
On now applying
the
galvanometer spot
alternating
in
excitatory
the oppo-
321
site direction
was
The
two.
We
next proceed to deal with the response of the glanThe tongue of the frog has formed
a very extended
of
subject
the
Engelmann and
Biedermann.
careful
entailing as
isolation,
On
little
by
of researches,
series
very
injury
as possible,
it
'
'
As
unknown
excitatory after-effects,
rise
ap-
it
peared to
In connection
this, I
Our
is
Photo-
Record of
Electrical Response
of
Intact
Human
Lip
Responsive
current
from epithelial to
epidermal surface.
in this
lies
spot of light
is
apt to be erratic in
in
its
movements.
Much
overcome, however, by holding the protruded tongue lightly clamped between the teeth. The upper
of this difficulty
is
may
already described.
From
the teeth
matters
it
in
with a
is,
such
little
way
that
practice,
the
clip
and
possible to arrange
galvanometer
spot
is
COMPARATIVE ELECTRO-PHYSIOLOGY
322
The
practically stable.
tongue
is
human
as in the frog.
when we
subject the
is
This
is
now found
very
The tongue
is
its
characteristic
unfavourable direction.
Thus,
if
circuit,
is
induced
in
response.
This, as
in direction
The
its
a very
in series
already said,
the upper
always
with
is
in
is
in ailother,
by
direct,
and not by
the secondary.
when
the primary
shall
now proceed
is
to
pushed
again
occurs
may
is
3:^3
excessively strong.
We
the tissue.
have
seen
also
is
galvanometric
excitation
positivity
intensity
we may expect
this
being
positivity,
Hence,
effect.
of
sign
the
that
in
dif-
to find instances
falls
will
become galvanometrically
possibility
of
negative.
We
very
feeble
stimulus.
It
the contacts
is
of both
excitability
been overlooked.
two
different surfaces.
the
curve
which
By
shows
of stimulus and
response.
curves
of
the difference
is
the
tissues
characteristic curve
the
relation
is
will
at
the
here meant
between
This difference
which
factor,
intensity
be better
below
(fig.
197).
This exhibits
all
exist.
of the surface A.
surface B.
Of these two
under moderate stimulation, nornially the more
excitable.
In the middle portion of the curve, representing
response under moderate intensity of stimulus, the induced
is
galvanometric negativity of B
is
Under moderate
in the direction
from B to
A.
is
A.
b'->a'
But below
COMPARATIVE ELECTRO-PHYSIOLOGY
324
negative
relatively
to
it.
Hence
there
would
be
here
a^b through
now from A
to B.
This current
will
Fig. 197.
A, a,
a',
Under moderate
y^
b'
that
U'
is
is
seen to be very
much
steeper than
A a
a'
a"
We
a"->b'',
325
under
same specimen
but
it
may be
possible
to
find different
do not cross.
effect,
In
and
in
their higher
there
to
this will
is
the most
find
common.
We
normal
to
tensities of stimulus.
b'->a'
in
higher
in-
such an instance,
is
sub minimal.
An example
presently.
stimulation.
I
have not yet been able to find a single specimen exboth reversals, but it is not impossible that this
hibiting
exists.
The type
in
wide range of
COMPARATIVE ELECTRO-PHYSIOLOGY
326
may be expected
excitable tissue
fatigue,
other
than,
things
to
show
earlier or greater
being equal, a
less
excitable
depressed, so
as to
fall
reversal of response.
below that of
This
A,
with consequent
be seen clearly
will
in
a typical
Fig.
(a)
198.
electric
shocks.
intensity, the
the primary.
normal, the
the
upper.
The
The
responses
(fig.
198, a) are
They
also
show
signs
seen
the
to
be
lower to
In this matter, the nature of the tissue must be taken into consideration,
a record
same
to stimuli of the
(fig.
to
its
original position
intensity as before.
response
is
responses
successive
its
327
be seen
It will
B.
have
obtained
response
other
the next
interesting
induced by fatigue.
making
be shown
will
chapter
variations
surface
in the
208).
(fig.
normal
of
the secondary
of the
the tissue
Moderate
fatigue
was
thus
secondary was
induced.
to
When
the
was found
to
be reversed.
sufficient
to bring
when the
testing
stimulus.
normal response
given in the
record of
its
first
series of records in
fig.
199.
is
After taking
COMPARATIVE ELECTRO-PHYSIOLOGY
328
the
out
responses were
now found
The
series.
We
have
last to
Such
stimulation.
normal
the
responses,
true
whereas
due
reversal,
to
Such an
beginning.
I
have
observed
response
Photographic Record showing
Fig. 199.
Reversal of Response in Carpel of
Dillenia indica, under Sub-minimal
Stimulation
The
first
series
show normal
electrical
of
The
lip.
very
effect
in
the
human
the
direction
of
normally,
under
stimulus,
from
thelial
to
surfaces.
stimulus,
the
moderate
the
epi-
epidermal
this
may be supposed
intensity.
liable to
and
it
much
With
to be localised only
be involved
then becomes a
in
difficult
situated.
CHAPTER XXIV
RESPONSE OF DIGESTIVE ORGANS
Consideration of the functional peculiarities of the digestive organs
Relation
Alternating
responses.
tentacle of
Having now
and glands,
up
skin, epithelium,
alike in plant
mucosa.
And
is,
is
first,
the
functional
specialisation
accentuating certain of
its
of the tissue
has acted
in
responsive peculiarities.
it
consists, briefly, of
two
different processes
is
those,
rendered
soluble,
solved
foodstuffs
are absorbed.
its
In the
then, the
most inclusive
sense,
membrane of
epithelial
lining,
dis-
the
using
COMPARATIVE ELECTRO-PHYSIOLOGY
330
circumstances.
The
itself,
what are
more or
or absorption, must be
to,
whether of secretion
less long-continued.
Thus
The
understood to be by secretion.
different
traction
is
responsive
phenomenon
we
to regard con-
it
will
be well here to
and
ways
first,
that, in
is
or,
secondly,
The two
theories
may
Mimosa
pulvinus
it
secretion.
is
In
the
thin
that
is
noticeable,
1;
is
exhibited in a marked
33
The
is
invested
Thus a
is
may
The occurrence of contraction is thus most easily demonstrable when it is accompanied by conspicuous movement.
This, however, demands considerable physiological anisotropy, the differential contraction then giving rise to a very
marked
lateral
movement, as
in
Mimosa.
Ih radial organs
in
is
fluid-transference
is
manifested by mechanical
movement
is
directly
visible as secretion.
From
in the tentacles
of
Drosera,
summit,
is
cell-
fests itself
by bending.
It
COMPARATIVE ELECTRO-PHYSIOLOGY
332
may
excitatory reaction
movement and
secretion
portion offers.
citatory expulsion
is
would
a vegetable organ,
The
electrical response.
two
The
first
of these
galvanometric
The second
is
is
with positive
positivity,
the
wave of
turgidity-variation.
istic
the
in
The
vanometric negativity.
first,
character-
its
concomitant gal-
consisting, as
this
does,
it
could not, as
shock which
to
take
is
hydrostatic
transmitted to a distance.
place
an
active
expulsion
requisite.
In view of these
of discontinuity,
we
or,
facts, is it
when
there
is
evidence in
its
favour, are
reactions
The
have hitherto
tissues
been
the
first,
case
of the nerve,
visible
it
manifestation
however,
classes.
In this
this
supposition
is in
is
fact
by contraction. The electrical indication of galvanometric negativity which is concomitant with contraction
attended
in contractile tissues,
is
The
glands.
333
visible
only
in degree.
We may
now
turn
more
As
current of
natural
rest,
regards the
the
negative, as
relatively galvanometrically
The
positive.
opposite direc-
snail,
metrically positive.
That the
fact of cutting
experimental preparation,
making
in
electrical
persisted
for
subsided
no
fresh
experiment
and
could
be
until
in the habit
many
these
undertaken,
had
to
to external
COMPARATIVE ELECTRO-PHYSlOLOGY
334
mechanical
irritation
which would
of
mucous
negativity of the
contribute
the
to
lining.
tentacles.
it
made
here
electrical connections
respectively.
was seen
of that surface.
From
it round, an
galvanometer as negativity
may be gauged the difficulties
in the
this
a strong presumption in
difficulties
layer
As
is
this tube,
however,
is
closed,
it
rest.
cannot be regarded as
They have a
(fig.
200).
These, as
is
well
known,
of
their
lining
it
dissolved or decomposed.
The products
supplying
me
for
335
first
disturbance.
connections, by
next made
electrical
means of non-polaris-
and
dular)
outer
surfaces
tory reaction
of
this
As some excita-
pitcher respectively.
may be
induced
in
with
natural
the
pitcher itself
On
secretion
of
the
found, as
had expected,
negative,
obtained
diphasic,
it is
and
Before
advisable
and
We
Mimosa
tissue
that
is
is
COMPARATIVE ELECTRO-PHYSIOLOGY
336
we
accompaniment of a
positive turgidity-variation.
sive
posi-
tivity.
Again
when
excited,
chanical
Fig. 20I. Glandular Surface of a Portion
of the Living Membrane of the Pitcher
of Nepenthe.
the
fresh
Mimosa
vinus of
fall,
by a mea
negative
turgidity-variation,
galvanometric
But
after
pul-
responds,
and by
negativity.
long-continued
and the
leaf
is re- erected.
tivity is
now
reversed to positivity.
It will
(^)
J-Zl
the
&
electrical indi-
of
cation
this
process
in
a tissue which
hand,
been
under
con-
be a tendency towards
sponse to galvanometric
positivity, indicative of the
Transverse Section of Tissue of
Fig. 202.
Pitcher of Nepenthe.
u, outer surface
L, inner surface
g, glands
process of absorption.
these
excitatory
337
But in glandular
which they occasion.
become possessed of much greater significance,
they constitute the main function of such structures.
responses
motile
organs, they
since
Electrical
responses, in
way analogous
every
to
those
That
surface
tivity
when
is
on excitation.
conditions secretion
is
phase of response,
Mimosa^
asso-
is
After continuous
Mimosa^ is
is
every
From a consideration of
organ, we should be prepared,
already pointed
out, to
alternating processes.
acting
as
a stimulus, would
naturally
induce
excitatory
secretion
later
We
is
and glandular
that, as in the
one
tissues.
of
fluids
we have
secretion
We
In the
and
and negative.
As
far as
it
have found
it
has invariably
been the case that the negative electrical phase was associated
with secretion
good.
the
holds equally
COMPARATIVE ELECTRO-PHYSIOLOGY
33^
We
now
the pitcher in
youngest,
known
no
its
as 'fresh.'
insects.
excited,
Of
flies
are
present
these, in the
such a specimen
will
be
Still
another class
is
found, in
been
subjected
continued
The
to long-
stimulation.
responses of
these
shall
first
experiments
on
of
shocks
of
intensity,
at
alternating
Photographic Record of Series
of Normal Negative Responses of Glandular Surface of Nepenthe in Fresh ConElectric
Equi-alternating
to
dition
Shocks given at Intervals of Two Minutes
made
were
responses to equi-
their
Fig. 203.
out
specimens.
fresh
Records
describe
carried
moderate
intervals of
The
two minutes.
responsive
current
the
internal
glan-
glandular
surface.
Fig.
But under
is
later.
fig.
203,
is
in
the
series of re-
339
trends upwards.
Had
it
exceptional
Nepenthe^
it
naturally gal-
is
vanometrically positive.
I
may
point out
here
the
which occurs
tion of response
in the
due to fatigue
so-called,
makes
appearance
its
Some
stituent
show a
tendency
first
to
and
to positive,
more
become
or
in
dimi-
of the con-
stimulus
are
less
completely
reversed.
It is also interesting to find
make
that the
same
same modifications
action
of such
stimulation,
the
characteristic
response
fig.
205 that
effect is
which are
positive.
is
in
is
It
such a case
a series of mul-
COMPARATIVE ELECTRO-PHYSIOLOGY
340
downwards.
glandular surface.
The
results
in
the case
it
will
be observed,
galvanometric negativity
is
associated
in
which
with a predominant
secretion,
Fig. 205.
Photographic Record of Responses of Pitcher in Third Stage, the
whole Glandular
Surface
thickly
Coated with Insects. Stimuli applied
at Intervals of two Minutes
Photographic Record
Fig. 204.
of Responses of Pitcher in
having
Stage,
Intermediate
Attracted a Few Insects
The
It
is
condition
also
the
surface
fresh condition.
is
is
positive,
in the fresh
and
in
the
is
apt to be
341
if
strong,
-"'
1
'
'
o'
Fig. 206.
somewhat
different con-
ditions.
The
stimulus
found to
persist, in both,
for quite
the
first
an hour.
In
of these figures,
and
positive,
Thermal Shock
The
record
positive
207) the
phase is pre-
(fig.
constituent
dominantly
responses
are
here
pre-
positive.
dominant in the responses, and the trend of the base-line downwards shows increasing positivity of the glandular surface.
COMPARATIVE ELECTRO-PHYSIOLOGY
342
appeared to
me
that
thrown, by the
difficulties
connected
experiments which
in
justified,
the latter
seen in
it
be well
will
seen
plant
the
in
Drosera.
took for
my
experiment a specimen
of the Indian Drosera
longifolia,
upper
the
are covered, as
Photographic Record of Responses
Fresh Leaf of /);w^ra to Equi-alternating
Fig. 208.
in
Electrical
The
Shocks
show normal
is
well
Current
from upper glandular to lower non-glandular
surface.
In the second series normal response
first series
is
responses.
Oi
,
^j
I\epent/ie,
tnC
re-
of leaveS which
sDOnse
^
are fresh and have not
leaf,
to the
interesting
note here, as
to
343
the
in
case of
when
the response
is
the reversed
down when
it
As
Nepenthe^ so also in
the
in
leaf of
the
that
is
Drosera,
to say, previously
responses
(fig.
The
leaf,
therefore illustrates
the
fact that
plant, intact.
petiole
On now
cutting the
a long
across,
series
of
rapid,
at first
down
pulsation
being about
30
direction,
the action
these,
it
strongly
or
responsive
As
rest,
and that of
regards the
first
of
will
'
current.
'
COMPARATIVE ELECTRO-PHYSIOLOGY
344
From
we have
as
We
same electro-motive
similarity
was supposed,
is
as,
in
outer skin.
its
no such
reality
reaction
effect
it
reaction as
shall find,
this
positive, its
we have seen
further,
is
to
is
from
The ingoing
is
to isolation.
With
'
negative variation
current
that
mucous
Biedermann
by rapidly alternating
usually preceded by a
is
to
of
'
say,
it
evidently means an
galvanometric
ingoing,
is
outgoing
positivity
of
the
we must be
And
first,
citation
effect,
on the mucous
Method
demonstration of the
surface,
contact,
tissue.
of ex-
effect
uncomplicated by changes
I
employed the
Rotary
The
The
following record
(fig.
The
345
responsive variation
even
and
to
reversal
from
normal
the
to
positive.
In
order to show
relatively
the inner
that
mucous
surface
coat,
is
next
subjected the two to simultaneous excitation by equi-alternating electrical shocks. And for the sake of establishing a
Fig. 2IO.
Photographic
Record
of
Negative
FiG. 2 1 1
Photographic Record
of Normal
Negative Responses of Stomach of Tortoise to Stimulus of Equi.
Normal
Responses
of Frog's Stomach to
Mechanical Stimula-
Minute
a different specimen
responses in the
now took
namely, the
figure
(fig.
sponding
fig.
208).
effect in the
shall
COMPARATIVE ELECTRO-PHYSIOLOGV
346
external surface.
The next
record
has been
212).
(fig.
more than an
hour.
Of
series
this
produce
portions
here
four
(fig.
re-
different
213).
The
first
sists
of pulses of gal-
vanometric
negativity
The
incomplete,
and
the
showing an increasing
negativity of that
The
face.
sur-
negative
Photographic
F.G. 212.
Response
in
(b).
of
positive
response
{c)
the
base line
is
In
this
the
first
part
pronouncedly
horizontal.
It
then
or
increasing positivity
of
we have
we have already
(figs.
single
347
sustained.
rise to
There then
Fig. 213.
Four parts are given of this long series, {a) Negative series ; (If) Alternating negative and positive constituent responses
{c) Positive series ;
(d) Positive series.
Note trend of base-line upwards in negative a,
and downwards in positive d.
;
sorptive process.
the ab-
positivity,
may
or
we have seen
itself,
that
long-continued
stimulation
was apt of
response.
mum,
empty mucous
the
cells in
And,
in
its
maxi-
lastly,
COMPARATIVE ELECTRO-PHYSIOLOGY
348
tends to give
to a
rise
Now
to be the
expansion
is
instead of con-
dissolved food.
is
to be
remembered
in
that periodically-acting
which
persists for
well-known
tropic
illustration of this
movements,
is
met with
so-called, of plants,
in the nycti-
by the periodic variation of night and day. These movements persist for a certain length of time, even when the
plant
is
kept
in
continuous darknes.s.
Similarly,
animals
CHAPTER XXV
ABSORPTION OF FOOD BY PLANT AND ASCENT OF SAP
Parallelism between responsive reactions of root
sap Electrical
reactions,
Association
Alternating
as in pitcher of Nepenthe
of sap
Demonstration of excitatory
experiments on
effect of
Assumption
that
electrical response of
tified.
We
have seen
cess as a
in
and absorption.
The
we found
of absorption, by
by which
to take
by a process
though
seen
characteristically
in
the digestive
some
organs of
Here, situated
externally,
it
is
same sequence of solution by dissolving fluids, and subsequent absorption, must be gone through. And the organ by
which this takes place must evidently be the root. In this
regard
the
well-known experiments
on
the
corrosion
sufficient to
of
show
COMPARATIVE ELECTRO-PHYSIOLOGY
3 so
made
substances are
soluble.
by means of which
It is
insoluble
Thus
it
will
cesses of secretion
distinction
between the
main organic, and in the plant inorwe meet with connecting links
in the form of insectivorous plants, in whose case the organic
supply is obtained by means of the digesting leaf, and the
supply of food
Even
ganic.
is
in the
here, however,
roots.
We may
regard
digestion,
its
or inorganic.
Apparently, then,
other.
which
is
is
maintained continuously
useless water,
must be con-
which
is
it
must be possible
accordingly passed
off,
may
be
The
from the
absorption of
shall, in
the
which
(i)
any
Whether
and
(2)
35
fundamentally due to
is
With regard
ascent of sap
is
nature
of the efficient
it
may
The
as constituting
be
cause of the
will
As
been
urged
could
(a)
take no part
moreover
{b)
it
has
that
in
the
ascent
of sap.
It
is
known
And, lastly (^), in the wellknown experiments of Strasburger it was found that strongly
not prevent the ascent of sap.
it
trees.
From
But
if,
it
movement
was
of water in
The attempt
to
do
this will
next chapter.
The
first
question
investigation, then,
is
that
as to
falls
within
the
scope of our
We
have seen,
two
positive, the
35^'
COMPARATIVE ELECTRO-PHYSIOLOGY
and the
it
will
be interesting
young or
excitation.
when
subjected
to
The fact
fresh specimens,
that young rootlets secrete, on excitation by contact, has
to demonstrate here the occurrence of secretion in
already been
seen in
corrosion of marble,
hering.
One
of the very
young
roots
precipitate, or
in
the
silver
electrical
mode
of investigation,
we
galvanometric
becomes reversed
to positivity under
by galvanometric
is
recorded
Among
are dead
interesting
to
find
that
of
responsive
phases.
For
responses
its
is
it
alternation
similar
demonstration
this
One
and decaying.
The
root.
naturally
some which
and vigorous
positivity.
353
responsive
however,
normal
opposite,
be
to
reversed
the
to
this
to
its
this
we
consideration
of
From
positivity.
pass
found
by galvanometric
response
negativity
may
have often
Now we
typically positive.
is
two
under which the
are
previous
remembered, that
different
The
first
is
we
which
is
Photographic
214.
conditions
may
positive
negative response.
Fig.
have seen
chapters,
as will be
^
there
falls
And
below the
the
critical
we saw
less excitable.
positivity, in
a tissue that
am
significant.
A A
COMPARATIVE ELECTRO-PHYSIOLOGY
354
We
gives
the
Taking next an older root of the same plant, and employing the same intensity of stimulus as before, I found the
responses to take place, generally speaking, by galvanometric
This would appear to suggest a tenpositivity (fig. 215).
dency towards specialisation of function, galvanometric negativity being associated, as we have seen, with secretion, and
in
positivity,
specialisation
similar
and others
secretion
probability,
all
absorption.
for
absorption
unmistakably
in
namely,
of certain
cells
for
more
manifested
is
In the older
roots,
to
Fig.
Phoiographic
215.
the
to
activity
is
proof of
rise
the
to
so-called
further,
likely to
that
-^
'
root-pressure
have
physiological
activity
^ in virtue of which
^
water is taken up by the root, thus
into
we
indications,
electrical
would seem
giving
We
pressure.'
be absorptive.
from
we can
see
'
is
'
root-
activities
kept approximately
this
physiological
fact
which
is
soil,
and
is
I
undertook to
test
The
induced by these.
specimens which
have
355
young
when subjected
-5
Thus,
in
to the
The continued
induced.
action
the greater
cent,
number of the
much
and
it
per
'5
required a
to
We
and that
it
is
absorption as food.
We
reactions in
trical
young
root, as in the
characteristic
that
rendered soluble,
for subsequent
have seen moreover that the electhe two cases are similar
that in the
inorganic substances
We
saw
further that older roots, like the glands in the older pitchers
is predominantly
now, having thus completed our first line of
inquiry, we shall turn to the second
the question, namely,
positive.
And
is
or
is
physiological reaction.
from leaves.
Under the latter,
movement of water is regarded as
physiological, the
mainly due,
actions
AA
COMPARATIVE ELECtRO-PHYStOLOGY
356
manner,
this
factors.
Among
pounded
which have
physical theories
the
been
pro-
sap those of
inadequate.
transpiration,
put
The
mesophyll
fluid in the
cells
thus
difficulties in the
is
way
set
up by the
of this theory
leaves,
lie
(i) in explaining
how
and
(2),
in the
that,
under
have
tensile
sufficient
strength.
column of
It
is
energetic water-
absence of
transpiration.
a tree which
these
may
liquid
metres
in
height
is
an independent activity
01
is
in water.
makes a place
when
their cut
stems
'
vital
then
it,
it
is
357
obvious that
we
a theory
it
On
such
sponsive
movement of
They
water.
The
objections
made
to the
physiological
movement
of water
is
known
to take
rapidly,
shall
now
We
the
to
in trees
woody
dead.
It is not,
the presence of
asmuch
tissue in
it
wood
that
Pfefifer,
this
is
woody
tissues.
before the
In seedlings of Gramince
COMPARATIVE ELECTRO-PHYSIOLOGY
358
here at
its
is
ex-
Transpiration
leaf.
fibro-vascular elements at
cannot be regarded as
'
dead wood.'
Finally,
Assuming,
is
for the
wood
in the interior
in the case
were dead,
it
of
might
conceivably be of use
still
central reservoir.
rapidity of transit.
At
wood
and
at the
leaves.
According to
might be supposed that it
it
As
a matter of
fact, this
system
is,
as
reservoir.
an hydraulic machine.
The
indicated.
we have
also to
remember
conducting, system.
wood
is
cortical
tissues
we may
Here we have
insignificant,
is
no
In
woody
trees, then,
is,
that in this
case there
is
also an
The
added and
which
channel available,
requisition
In a
where quick
woody
transit
trunk, then,
359
naturally
will
come
into
required.
is
we have
woody
tissue,
it,
but
is
The
innumerable
cortex.
lateral
When
reservoirs,
demand
communi-
it
cells
of the
wood.
As
with the wood act in a manner not very unlike that of the
roots towards the
soil.
That
is
to say,
it,
into
it,
these
alternating
oi
makes
There
is
here, in
we
may
is
find access.
On
re-absorbed by the
Such
movements of inflow and outflow evidently take place in the
trunk of the tree itself
Under the stimulus of sunlight,
organ, causing expansion and re-erection of the leaf
the excited cortical tissue will squeeze water inwards into the
central reservoir.
in
is
a diametric contraction.
At
the time
when
be seen
transpiration
is
thus
COMPARATIVE ELECTRO-PHYSIOLOGY
360
the
besides
coming
water
from the
lateral
roots,
an additional
The
reservoirs.
loss
of
We
central reservoir.
Mimosa
water
expelling
absorbing
it
the
into
wood
on
of these reactions
made by Kraus
the cortex
in
and
excitation,
explains
The
re-
occurrence
the observation
We
excitatory
is
supposition that
passive
The
agent.
has
question
still
to
be
attacked,
wood used
conduction of water
for
is
is
dead.
that
possible
is
it
for
woody channels
it
is
conducting
it.
For
my own
part,
and
useful
perfectly
for
purpose
the
colourless,
of
this
investigation.
It
is
appropriate development.
in this (Jilute solution
and
left
for
some
time.
Transverse
361
made and
now nothing
placed in the
There
of a microscope.
field
is
up by the
The
now
crimson colour.
By
this
means
easy to
it is
conduction
of
is
concerned
water,
the
in
older,
purpose.
If,
see, as
the work
or
'
already
the younger, or
is
it
of the rapid
heart- wood,'
part in the
it is
being
difficult to
as a
woody
living,
It
tissues.
occurred to me,
finally,
is
an
rest.
The
first
them a period
local
then subjected
then soaked
in water, it entirely
COMPARATIVE ELECTRO-PHYSIOLOGY
362
response.
response.
stem,
With sap-wood
was
dication of galvanometric
negativity in response to
Hot
thermal stimulation.
distance
of
mm.
was next
to
records
Photographic Record of
Electrical Response of Sap-wood
Fig. 216.
desirous
photographic
obtain
of
the
normal
response of living
and
The normal
its
wood
variations under
For this
employed both
the electrical and vibrational modes of stimulation.
For the
first of these, the strip of sap-wood was cut in the form
of a two-pronged fork, of which one prong was killed by
chemical agents.
purpose
was kept
now
by immersion
alive
in water.
air,
After
half
usual
men.
this,
electrical
On
next
killed
subjecting
connections were
made
in
to
equi-alternating
the
speci-
shocks'
first series
of records to the
left.
in
fig.
216
in
and we observe the consequent depression of rewhen the chloroform was blown off the responses
applied,
sponse
363
have thus been able to establish the fact that the woodysap-wood are not dead but living, and hence
vessels of the
This
will, I
think,
physiological theory of
the
We
have
been
by
killed
boiling^
'
These observers
burger.
J
ends
of trees
solutions, such as
if
it
in spite
It is clear
they must
this
^ a1^"o1
,.
s^ri,*-,"
set cut
/.
^,,
Photographic Record
217.
showing Normal Responses ot
Fig.
kill
was
all
elements
And
from
stem could
Strasburger was thus
in
transpiration current
is
in
the
ascent
of
the
absolutely precluded.'^
It
part of Strasburger
fact that
to exhibit suctional
cerned
'
if
activity.
the process
is
Hence the
active
to be regarded as
cells
con-
due to such
COMPARATIVE ELECTRO-PHYSIOLOGY
364
zone, then,
Thus
a poisonous
it
maintained above
again show in
when
suctional activity
Under the
show that
water.
able to
is
by boiling
killed
have been
tissues
in
It is
we have
areas
whose
activity
That copper
sulphate solution arrests rhythmic activity, and induces
death, is seen by the rapid stoppage of pulsation when
the pulvini
wc apply
When
of the
it
it is
inserted
lateral
leaflets.
end of the
petiole, the
to
one after another, and that the death of a point below does
From this experiment it is evident that the application of poison, at the root, or the cut end
not stop the suction above.
experiment
come
to a stop
We
it is
killed,
when
movement of water
did
tree.
shall therefore
proceed
in the
next chapter
this process.
CHAPTER XXVI
THE EXCITATORY CHARACTER OF SUCTIONAL RESPONSE
Propagation of excitatory wave in plant attended by progressive movement of
to stimulus The Shoshungraph Direct and photo Responsive variations of suction under physiological
by various agents Effects of lowewng and raising of
Explanation
the over-balanced
by action of stimulus
standstill,
stimulus
of maintenance of suction,
influenced
of poison
at
by
after-effect
of stimulus Response
In the
chapter
last
it
shall
the plant
justified.
is
in
Instead of
we
moreover,
proceeding
shall
attempt,
of other excitatory
reactions,
from the
basis
to see
We
it
immersed
salt,
for
until the
specimen
water.
is
some time
tissue has
in a
common
this.
The
It is
now
COMPARATIVE ELECTRO-PHYSIOLOGY
366
in
up
tissue higher
The
now
is
portion of the
is
wave
at the lower
The
tissue.
end
is
attended by an expulsion of
This expulsion
instantly
is
made
visible
by the formation of
From
experiment
is
it
in
this
is
at-
the direction of
propagation.
The next
point to be realised
continued stimulus
is
If the tip of
we
see successive
that
excitation
proceeds in
all
directions
from the
be terminal, it
is evident that the direction of propagation, being away from
But if the tip of the root be highly exthis, will be upwards.
If then the point of excitation
excited point.
owing to local
amount of secretion
citable, then,
certain
into
the
Even highly
show
soil.
a tendency, as
istic
we have
response.
become changed
into absorption.
Again,
tion than
these,
if
for
which
the secretion
of
fluid.
And
in
absorp-
addition
to
evaporation
particular
electrical
young
help
to
the
are
conclusions
tend
will
We
uni-directioned flow.
roots give at
We
there
first
that
further,
there
with
the
regarding
for
secretion,
and
Thus the
cell,
serial
be upwards.
movement
In this connection
it is
cell
to
is
may
by
produced
be
And
of the tissue.
the
multiple
excitatory
activity
instance, the
be gauged by their
in
movement of water
of Desmodium^
leaflets
multiple
may
mechanical movements, so
we have
means of
by a determination of the
the heart,
circulating blood.
of
rate
of flow
of the
movement might be measured, either by means of the proby the suction exerted behind.
In order to demonstrate the fact that the water movement
in
the
ascent of sap
reactions,
it
is
is
pose
was successful
in
devising
the
Response.'
'^^.
364-371.
'
COMPARATIVE ELECTRO-PHYSIOLOGY
368
"
may
the specimen
be subjected
to the
rapidly
(2)
action
of
a potometric
The
in
is
means
of
reservoir,
fig.
which
may
be
india-rubber
filled
roots,
cork.
is
by
the
By
the plant-vessel, in
is
watertight
218.
appropriate manipula-
in the plant-
may
vessel
ment observed.
For
figure.
this
purpose, a
pen,
P,
is
fitted
collar,
over the
which has a
The
collar,
is
attached to a thread,
follow
the
water- index
pulleys.
movements of the
is
followed in the
water-column.
way
When
the
is
obtained.
curve
is
the
suction
is
As long
If,
as
however,
Fig. 218.
plant-vessel
r, reservoir
The Shoshungraph
c,
B ^
COMPARATIVE ELECTRO-PHVSIOLOGY
370
the plant
is
in
no way disturbed.
by allowing water
The balance
obtained
is
its
com-
withdrawal
declination downwards.
by me
suctional response
and
logical modifications.
variations,
fuller
Plant Response.'
As
it
will
a simple
manner of
its
be found
my
in
book on
is, how-
which
it is
shall
is
trans-
move
in
it.
This consists of a
water-column.
is
is
placed
the
in
field
of
on a moving
focussed
The
sensitive-
two ways.
the bore of the capillary tube may be made finer and
But this cannot be carried to an extreme, as the
ness of this
First,
finer.
in
movement of
the
may
be increased to any
extent by the employment of a highly-magnifying and
Secondly, the sensitiveness
index.
short- focus
devices,
we
objective.
By
of both these
the combination
most
difficult
it
is
ease.
For our
this sensitiveness.
The photo-
The
which casts an image without any magnification.
Method of Balance again, as we have seen, affords us another
opportunity of arriving at an experimental adjustment of
great sensitiveness.
The experimental
delicacy obtained
tion
greatest
We
when
vessel
by suction
is
its
inflow
(fig.
218).
final
adjustment
the narrow bore of the thick india-rubber tubing, which connects the compensator
gradual constriction.
is
capable of
compression on
We
shall
now proceed
it,
vibrating
leaflet
of
Desmodium has
Thus an automatically
its
vibration-frequency
BBS
COMPARATIVE ELECTRO-rHYSIOLOGY
372
enhanced by a
rise,
A similar
temperature.
response of plants.
effect is seen to
Thus,
in a
fall,
of
This arrest
normal rate of suction reappeared on the return of the water to a normal temperature.
On applying water of raised temperature to the root, on the
other hand, the rate of suction was immediately found to
be enhanced. This is illustrated in the following record
(fig. 219), in which the normal rate of suction at 23 C. was
arrest the suction in the course of 8 minutes.
for the
7 cubic
in
mm.
in
volume, or 7 mg.
The
plication of water at 35 C.
induced a steep
1
n
on Return
Normal Temperature
internal
to
beginning.
to 14
low as at
so
fall
It
now
fell
returning to the
is
due
to
original 8
the
fact
water, the
in
mg.
that
the
enhancement
this.
water over
tained.
taken,
vessel
On
from 58
in
the
We
of
This
the
rate
not
to
indicating an enhanced
ap-
now
it,
is
nevertheless main-
it
at
show that
appear to
first
protoplasmic
had
activity
described,
unkilled
is
contiguity with,
above
The
by the
plained
which
that
tissues
unabated.
however, in
killed,
is
is
immediate
The only
which
part
immersed
actually
the
experiment
the
water
boiling
continue
their
and
is
in
the
activity
suctional
just
or
in,
to be ex-
fact that
now
the root, acting virtually as a wet rag tied round the base
of the
possible
stem
in
to
suck
evidently
is
The mass
stem.
living
through the
up
of water which
directly through
much
the
it
is
thus
broad-sectioned
resistant, organically-conducting
channels of
the rootlets.
The
fact thus
above
top of a
it,
tree.
explains
It is
why
a poison
all
may be
when
it
carried to the
And Strasburger himadmits that arrest under these conditions does occur.
With
its
it
is
im-
pulsatory
movement
shake
COMPARATIVE ELECTRO-PHYSIOLOGY
374
absorbed, by
growth
is
have shown
on the response of
The experiments
in question
optimum
tonic condition.
The
have shown, an
application of a
per cent,
of these was
first
arrest of growth,
finial
set,
followed
optimum
arrest of growth,
The
was
still
by the application of a
This was
induce a depression of growth, which was ultimately
more
smaller dose
found to
strikingly exhibited
namely,
of a
rate of growth.
Thus
plant
is
seen
of the plant.
plants
suction
to
In accordance with
this,
in
others.
All alike,
however,
Excitatory character
The
may
sap
of"
The
record
is
taken
first
down
to the
solution.
The
cubic
of suction
rate
mm.
is
per minute.
replaced
had
in
by ether
After
in the
went
preliminary
This short
arrest
arrest.
was
suc-
which lasted
for fourteen
to
by
induced
ether.
This, however,
liminary,
is
is
induced.
only pre-
an
after
renewed at a
suction
interval being
very slow
rate.
When
Action of Ancesthetics in
Fig. 220.
Abolition of Suction
this
the
process
gradual and
It
undergoes
is
final arrest.
reaction
said
that
COMPARATIVE ELECTRO-PHYSIOLOGY
376
evaporation,
in
the
That
leaves.
osmotic effect
this
is
is,
chiefly
The
physiological action
sodium
Thus the
citatory.
action of
physiologically
is
the
at
common
hand,
is
same
more
osmotic
time,
is
A strong solution
pronounced.
of
its
salt,
on the other
osmotic.
we apply
then
If
KNO3
end of a
plant, in
Effect of Strong
KNO3
will thus
be a
Solution
The
There
of sap.
which
branch of Croton.
If,
however,
in
/r
(fig.
22
l),
it,
plant.
'
It
In
fig.
222
is
shown the
effect of
NaCl
solution in
salts, in
strong solution,
UJ
We have here a very great enhancement of the ascensional movement caused by this solution, which would, acting osmotically, have retarded the
normal rate. The physiologically excitatory action of strong
sodium chloride, however, is not permanent, and the enhanced
excitability is followed by depression. The effect on suctional
response in such cases, then, is modified by the factor ol time,
the
first
enhancement being
followed by a fall below the
original normal rate of suction.
The experiments which I
have here described show how
condition of excitability.
connected
is
action.
of suction,
only remains to
it
demonstrate
stimulation
is
responsive
in
attended
movement
a tissue.
by a
of water
In order to do
we should have
some
means
stimulus which
at our disposal
of
is
Fig. 222.
applying
capable
Effect of Strong
NaCl
Solution
this,
The
of
in
itself
of
normal
.the
effect
rate
of suction,
immediately induced
The
possible
COMPARATIVE ELECTRO-PHYSIOLOGV
378
I.
The water
in the plant-vessel
decomposition by electrolysis.
evolution
of heat in the
plant-vessel.
In
2.
a sluggish
response.
borne
but
little
first
of these objections,
be
to
is
it
polarisation effect.
The
also likely
be very
to
The extent
slight.
of
from
disturbance
the
be determined by
blank
The
experiment.
by
whose proper applicaelectrodes,
allowed
are
hang down
Record of Blank Experiment showing
Absence of any Disturbance of Record from
Fig. 223.
Induction-shocks as such
vessel
to
in the plant-
without
direct
The alternating
current will now pass
itself.
ducing
effective
excitation,
The Shoshungraph
given as we have
that
in
ex-
is
intensity
subsequent
is
passing
incapable
through
of
the
proplant.
seen,
a horizontal
line
of induction-current which
experiments
is
of record, and
is
to
be
used
the
plant-vessel
for
disturbing effects,
if
is
In
to be studied,
seldom exceeds
It
would,
if
anything, be towards
We
and
is
minutes.
five
is
shall
movement
however,
see,
in the opposite
an enhancement of suction.
With
tion
may
or
may
not be
made
by means of induction-shocks,
tible
For
form of stimulation.
this
this
purpose
it
is
This necessity
coil.
much
further increased
is
of the induction-current
is
uselessly
two
either of
The
Mode of
different ways.
described as the
Terininal
first
This
connection
is
less
at
may
its
Or two
be
object
the lower
of a thread.
its
in
may
of these
Application,
to
by means
This
circumference.
across
transverse
effective
as
longitudinal
stem,
is
stimulation
mode
not,
of stimulation,
theoretically, so
would
be, but
under
be devised
(fig.
electrical stimulus
224).
I
of applying this
Here,
(fig.
225).
COMPARATIVE ELECTRO-PHYSIOLOGY
38o
The
record
of normal suction
is then taken by means either of the ordinary recorder or of photography. The variation induced in
many ways
as possible,
The
first
\y^
Fig. 224.
Terminal
Fig. 225.
Mode
Mode
of Application of Stimulus
Sub-terminal
of Application
of Stimulus
effect,
means
rate be
is,
of record
is
horizontal
line,
a depression
downward movement.
balance, where,
rises
owing
The
If this
method
is
that of Over-
com-
movement
of the index
to that of suction.
is
induced
down, and should the stimulus cause any enhancedown-curve must be replaced by a
balance,
is
ment of
as
in a direction opposite
have
said, the
most
in
striking manner.
As
made
movement of
to previous results,
in
studied
roots,
we
by galvanometric
negativity,
Less excitable
about to describe.
I
shall
deal
first
with
the
results
which
invariably
month of February.
The Indian
winter was just over, and the spring had not yet fully set
The
in.
still
cold,
We may here
once more to the initiation of multiple activity by
this plant
is
sat standstill.
deprived of
its
COMPARATIVE ELECTRO-PPIYSIOLOGY
382
multiple
movement
for instance,
cease
lets
of
its
leaflets is arrested.
to pulsate.
But
if
If the plant,
in a
now an
electrical
leaf-
shock of
as the imparted
case,
quent
(cf
energy
is
exhausted.
fig.
light.
Or we may,
in
such a
effect
141).
latent period
owing
and
from the impinging stimulus, the activity of the leaflet continues for some time, even on the cessation of stimulus itself
It is, in fact, by that enhancement of the tonic condition,
mode
of response, as
activity
a latent period
after
of two
autonomous
response
of
Desmodium
leaflet
referred
to
above./
We
suction
Fig. 227.
Record
226.
viously
of
Effect
of
Stimulus in Enhancing
Rate of Suction
FiG.
Photographic
Stimulus
The half-shaded
2i,.
Vertical line
ditions.
represents moment of
application of stimulus
for 30 seconds.
a condition which
is
The normal
represented
in
rate
is
by
exactly balanced,
by
the record
'
Stimulus of
If a cut
(fig.
227).
is
now
in
its
suctional
known, disappears.
This is commonly attributed to the
blocking of the cut end by mucilage and bacterial growths, since the making of
a fresh section is found to renew the activity.
This making of a fresh section,
activity,
as
is
well
COMPARATIVE ELECTRO-PHYSIOLOGY
384
firs^
This
is generally
somewhat
due to the persistent after-
is
stimulus
there
applied
be
will
once
more,
further
still
of latent energy,
is
enhanced
to be induced
There
Fig, 228.
Variation of Latent
Period as After-effect of Stimulus
teresting
observed,
rate
of suction
stimulus
responsive
of
was
thirty
in
is
and
in-
often
is
,.
This
record
case balanced,
duration
seen
to
question;
for
and
as usual,
place
seconds'
it,
is
The
forty-five
The enhanced
making
mitial
was applied.
take
thirty
in
The
228).
(fig.
which
effect
stimulus.
another
in
COnsequence of
energy absorbed from previous
Stimulation. This is the diminu-
this
seconds'
acceleration
by
is
rate
duration
involved
the
doubt, obstruct
the passage of water, and yet the total abolition of suction not be due to this
The more effective cause is, in fact, the run-down of energy, as
cause alone.
It
is
The
this, I
Fig. 229.
initial
It will
(fig.
minutes.
applied,
lasting,
long, being as
much
before,
for
is
five
as twenty-
once more
The
seconds.
latent
minutes.
will
Method of Over-balance.
by a down-curve, and
acceleration of suction
is
by diminution
manner the
Here,
indicated
first
In
fig.
down
C C
230
part
COMPARATIVE ELECTRO-PHYSIOLOGY
386
now
Stimulus of hah
and it will be noticed
that on account of the re-
applied,
sponsive
acceleration
becomes
slope
diminished,
the
increasingly
an
after
till,
interval
the
half,
curve becomes
After this
horizontal.
it
is
It will
thus
be seen
Suctional Response
under Over-balance
Fig. 230.
I
of suction which
sufficient
(fig.
231)
it.
Fig. 231.
is not merely
compensate the
ceeds
graphic record
to
showing
In order to do
in
this
within
the
limited
range of a photographic
plate,
The
first
over-balance.
owing
becomes
record
horizontal.
now
further
enhancement of the
The
rate,
two
second stimulus of
This
applied.
curve.
and afterwards,
of stimulus, the
which
is
is
seen to induce a
But on
is
induced a more or
less persistent
after-effect,
first
was seen
effect
to
make
renders
and other
the
facts
From
previously enumerated
it
these
be under-
will
is
to
increase
suctional
up
activity
to
certain
limit.
tissue,
under
we
effects
induced
in
mode
of
the
simplest
shall
proceed to inquire
what are the effects induced under a somewhat more complex mode of stimulation. This is the case with sub-terminal
stimulus, where the point stimulated is not on the external
extremity, but within the tissue, though near the lower end.
Under these conditions the excitatory wave will proceed in
two opposite directions, upwards and downwards.
The
as to
short
terminal
to
the
directly
than the
stimulation,
natural
upward
suction, probably
the
more
fatigued.
The
however,
may become
COMPARATIVE ELECTRO-PHYSIOLOGY
388
stimulus,
may now
be expected to reassert
itself,
thus con-
of this
of stimulus
followed, under
expulsion,
renewal
mode
of
the
are
seen
the
up-curve
in
upward
the
continued
suction.
following
represents
first,
is,
the
movement
intense stimulus,
These
various
of
by
effects
photographic
record, where
unbalanced suction
normal
Continuous
232).
(fig.
now
stimulation was
at the point
applied
marked with
vertical line.
It will
be seen
is
here
diminished, and
afterwards
reversed
expulsion.
This
into
movement
knew
expulsive
continues, as
already
time,
is brought
about by fatigue of the lower
versal to suction
Photographic Record of
Response to Continuous Sub-terminal
Stimulation
Fig.
232.
reversal
intensity.
In order to expedite
zone.
the
so
reversal,
curve
that
the plate,
applied
expulsion, converting
We have
It
it
still
vertical line.
the
will
be seen
into
renewed
how
this
from
reversed the
suction.
positive
young
roots
give
We
negative.
saw,
that
further,
was
reason
to
tion,
or
there
secretion.
positive
sub-tonic,
intensity
feeble,
tissues,
slightly, excitable,
tissue
on the
characteristic
when
the
roots
other
is
by overhand,
Under
subjected
are
tissue
stimulus-
fatigued
is
Negative response,
then,
general rule
when the
or when the
either
stimulation.
conditions,
moreover, we
as a
further, that
obtained,
is
and very
is
see
shall
is
natural
to
the
tion
is
we
of stimulus induces, as
effect,
either
by
initiating
saw, the
suction,
or
absorpto-positive
by enhancing that
branches,
cut
we should expect
meet
with
two
In one, where
the tissue
is
terminal
stimulation
tissues,
to
will
be
negative
or
expulsive.
In
whose characteristic
might be possible to
find cases in
390
COMPARATIVE ELECTRO-PHYSIOLOGY
limit, after
no
little
or
effect.
If
expulsive.
it,
may
to say, zero.
somewhat strong
These
results
Thus the same strong stimulus which in the one season will
induce absorption, might be expected in the other to provoke
expulsion.
of seasonal variation,
inferences.
These experiments, as
in
the
will
were
circumstances,
in
we saw
sub-tonic
that
Under these
condition.
terminal
the
application
ot
positive
response,
when
expulsion.
From
these
investigations,
it
The
shown
I have also
have shown to be
and to exhibit the normal response of
The sap-wood
to be untenable.
The long
when
the
fact
persistence of suction,
killed
the cut
specimen placed
by the
are
roots
poison,
in
Those which,
like cold
or anaesthetics, act, on
depress
finally,
the
excitatory
and
fact
arrest
that
its
suctional
activity
the water-movement
response has
ascent of sap
may
The
physiological
And,
a form of
also.
is
in the
by the
course
theory of the
CHAPTER XXVII
RESPONSE TO STIMULUS OF LIGHT
Heliotropic plant movements reducible to fundamental reaction
or expansion
organs
Various
mechanical
Electrical response
effects of light in
specific,
of plant to
of contraction
but concomitant to
light not
determined by
on transversely
distal
Mimosa
to light applied
to indirect effect
Mechanical
movement, followed by
and
response of leaf of
Mechanical
response
ment
Electrical response
of leaf of
Mimosa
fall
or negative
to light applied
response to light
Periodic variation
Direct and
under
light,
effects
Multiple
of excitability
after
effect
of(
Multiple
i-)or(+
-i-
mechanical
electrical
either plus or
response
+
minus
After
Three
types of after-effects.
The
first
arises, in
such response
is
peculiar in
its
is
character, or fundamentally
The
mechanical
is
movements
are
stimulus.
induced
These
growing as well
different
in
393
by
cases
as
pulvinated
in
This
organs.
by
incon-
sistency of effects
reaction, but
same
the
effects,
is
own power
its
any given
what
to decide
for
is
its
individual advantage.
I
bave
shown
however,^
elsewhere,
that
regards
as
light
is
extremely
definite.
This,
like
other
is
The
intensity
thus dependent, as
distant
point,
there
to
induce a
positive
turgidity
sponsive expansion.
I have also shown that
the various mechanical movements induced by the unilateral action of light, depend (i)
upon whether the stimulus remains localised on the proximal
side of the organ or is conducted to the distal
and (2) on
the relative excitabilities of proximal and distal.
I have
shown, moreover, that all the diverse effects induced by light
;
are
And,
finally, certain
highly
from
'
COMPARATIVE ELECTRO-PHYSIOLOGY
394
and
positive,
or
vice
oscillatory response.
to
summarise
the
constituting
versa,
On
multi-phasic
these considerations
principal
by
caused
effects
or
possible
is
it
light,
as
follows
Description
of tissue
Action
I.
Tissue sub-tonic.
observed
Eflfect
Expansion or enhanced
rate of growth, e.g.
of internal energy.
Coprinus
of
Piletis
drooping
made
ness,
dark-
in
re-turgid
by light.
Renewed
growth
darkof
rigored
plant
posed to
n.
Normally
ex-
I.
Moderate
light,
unilateral
light.
tive
subjected
A.
organ
causing
excitatory
contraction of proximal and hydro-posi-
citable
Organ
to
radial.
mitted to
dium
of
2.
Strong
3.
Ex-
light.
distal,
I.
Excitatory
con-
1.
traction of proximal
predominant,
owing
proximal
Transmission
of
through
highly
conducting
2.
excitation
tissue
to
more
ex-
lower
or
distal.
Greater contraction of distal.
citable
upward
indica^ and
ternatea.
or feeble transverse
conductivity of tissue.
response,
folding
of leaflets in so-called
' diurnal
of
sleep '
Robinia, Erythrina
Positive
e.g.
(Oltmanns)
strong
and
long-continued light.
nega3. Reversed or
tive
response, e.g.
Sinapis and Lepidium (Oltmanns).
under
Fatigue of proximal
excitatory contraction of distal.
aniso-
2.
neu-
first.
and
Organ
positive
expansion
continued light.
tropic or dorsiventral.
towards
e.g.
curvature
of seedlings
of
Sinapis ;
positive curvature of
Lepidium seedlings
(Oltmanns).
Neutral effect, e.g.
Sinapis and Lepi-
tralising
Curvature
light,
distal.
B.
1.
ex-
light.
2.
Negative
e.g.
Clitoria
response,
downward
fold-
called
of
'
in.
Description
of tissue
exhibit
multiple or auto-
nomous
Effect observed
Action
which
Tissues
395
absorpConsiderable
tion of energy, im-mediate or prior.
of multiple
response in Desmo-
Initiation
dium
re-
sponse.
under
action
continuous
of
in
light
photo
Biophytum
movements
tactic
of swarming spores
;
cf.
electrical responses
induced by
light,
is
which
we have
already
mechanical response.
I
been
able
to
through
analyse
'
From
the leaves of
Iris,
for
example, during
With
its after-
and Mathiola, on the contrary, he obtained positive response during illumination and
subsequently negative. Beyond the suggestion that negativity
may be associated with dissimilation and positivity with
assimilation, Dr. Waller offers no explanation of this opposition of effects observed by him.
He states, however, that he
effect.
leaves of TropcBolum
gave no response.
ordinary garden
'
under
Even
chloroplasts
light,
'
as essential to these
inasmuch as
shrubs and
trees,'
petals,
he found,
green leaves
'
of
moreover, he found no
Ixvii.
pp. 129-137.
'
COMPARATIVE ELECTRO-PHYSIOLOGY
396
response.
It will
'
is
offered
mutual opposition of the electrical effects observed or of the way in which the presence of chloroplasts
acts as a determining factor.
either of the
'
To
turn
now
to the subject of
my
proper inquiry,
it
has
may
be observed
in vegetable
tissues
by means of mechanical
is
vini,
is
And
it
may
be as
chloroplasts,' that
light
from pul-
chloroplasts
in
excita-
For
'
it
same
response.
'
chloroplasts
'
same
plant, characterised
by
less
'
chloro-
would often give much stronger response. The etiolated stem of celery, moreover, gives strong electrical response.
And, finally, it is an error to suppose that petals of flowers
are irresponsive to light, for I have obtained strong response
from petals of Sesbania coccineum and from Eucharis lily.
Animal nerve, again, in which there is no chlorophyll, gives
plasts,'
response to
As
light.
same organ,
shall
now proceed
to
show
exhibit
As
the electrical
wish, moreover,
action of light
is
will
397
side will
by
therefore
its
hydro-positive
be positive
but
if
suffi-
sponding
first
This
will
the corre-
Fig. 233.
Experimental Arrangement for Detection of Electrical Change
induced at the Point transversely Distal to Point stimulated
stimulator,
positive
into
the
movement, or
excitatory negative.
be observed
if
it
if
Similar
electrical
be so old as to have
lost
power of
its
motility.
nome caused
applied a
(fig.
233).
by means of a metro-
upper or proximal
side.
first
By adjusting the
made moderate.
heating current
It will
be seen
COMPARATIVE ELECTRO-PHYSIOLOGY
398
(fig.
an increasing positive
opposite point.
rise
under-
latter
variation,
upper
side.
It is at this
consequence of the
in
being transmitted to
is
By
experiment
this
it
B.
is
by the
tion of
jmm
joint
unilateral applica-
any stimulus
of the
effect
excitatory
is
the
direct
of
contraction
expansion of the
Another
Fig. 234. Record of Response to Moderate
Unilateral Stimulation under the Experimental Arrangement described
phenomenon
after the
has
ple response.
first,
after the
is
that,
been
reached,
effect
there
of oscillatory
may be
its
a gradual percolation
opposite reaction, the
ob-
tions
be
maximum
a series
is
factors in operation
to
Response
multiple responses.
distal.'
interesting
and, secondly,
we know
itself in oscilla-
that increased
hydrostatic
but to pseudo-conduction.
on
its
itself to
is,
wave
its
effect.
Thus
initiation
(/e
this
in
under a
heart,
snail's
399
high degree of
sufficiently
As
conduction
regards
strong stimulus
weak.
is
in
At
(fig.
first
(d)
235) this
now
may
be seen
in
an
The
maximum
positive varia-
the
reach-
causes a reversal of
owing to induced
the curve,
galvanometric negativity.
at the beginning
excitatory effect,
that
interesting manner.
ing
we know
general,
If
we had used
intensity the
first effect
would
and afterwards
reversing the
now
first.
shall
response,
positive effect
due to hydro-
this is
converted to
due to transmission of
excitatory effect under stronger stimunegative in
{b)
lation.
one of
its
effects thus
upper
half,
positive effect
and consequent initiation of excitatory conand (3) the continued action of excitation
and increasing contraction on the more excitable lower half.
traction there
COMPARATIVE ELECTRO-PHYSIOLOGY
400
shown
in
fig.
236,
where the
first
seen in
is
the up-curve,
is
here
that
an
light.
interesting
it
is
It is
observe
to
possible to obtain
after-effect
tive
increased
effect, this
due
positivity being
to
the
energy
internal
Fig. 236.
Mechanical Response of
Pulvinus of Mimosa to Continuous
Action of Light from Above applied
at
In
Moment marked ^
fig.
electrical
on
effects
stimulation
by
light
another
specimen
pulvinus
of
electrical
contacts are
in
this
case,
of
Mimosa.
in
the
The
made
point.
case,
is
more important
theoretically
further
of temperature.
lies in rise
401
Though
in
of light
by the
tissue
must
In con-
gradual
tion.
however,
periment,
contacts
electrical
the
as
not
are
on by
directly
acted
we
results
uncomby such disturbing
light,
obtain
plicated
factors.
The
first
electrical effect
brought about
half
application
distal
in the
lower
pulvinus
by
of light on
the
the
of
upper half
is
seen
Electrical
Response
in the
in
is
FiG. 237.
positive
half,
effect
at
the lower
its
true excita-
tory reaction, galvanometric negativity, the indirect or hydropositive effect being one of galvanometric positivity.
mission
of the
true
we obtained a
excitatory
effect.
transverse trans-
Similar effects
DD
are
COMPARATIVE ELECTRO-PHYSIOLOGY
402
also obtainable
by longitudinal transmission.
In order to
do
this
mens
at a distance of 5
gave
rise to
mm. from
metric negativity.
which
in
The
applied directly.
much
larger,
enfeeblement
was
ob-
is
naturally
there
as
by
238 shows a
Fig.
light
effect
no
is
transmission.
series
of
such
sunlight
through
during
case,
Photographic ReFig. 238.
cord of Series of Negative
Responses of Petiole of
Bryophyllum to Stimuli of
Sunlight of Five Seconds'
Duration applied at Intervals of Two Minutes
ward trend of
base-line.
a
five
previously
stratum
of
seconds only
passed
water
in
each
leaf of Bryophyllum.
noticed
that after
It
will
be
each response
an after-effect of positivity, in
consequence of which the base
is
line of
the
series,
instead of re-
wards.
see
to
multiple response.
The
we
shall
initiation
of
is
Mimosa
403
where the leaf, after its excitatory fall, was reon cessation of stimulus, above its original height.
This, as we saw, was due to a certain portion of thQ incident
stimulus becoming latent, and thus increasing the internal
236),
(fig.
erected,
energy.
We
shall
positive response, as
sometimes induced by
light.
This
Thus,
This
is
responsive
its
depression
Mimosa^
fall will
light, usually
is
for instance, to
the action of
or
mechanical stimulation.
We
we
fact,
it
caused by thermal
because
if
may
Now,
stimulus
is
it
is
also incapable
it is
necessary
even
(p.
moribund.
83) that
it
is
Under such
possible
conditions
have
shown
under stronger,
will
normal negative.
The
more
is
very moderate,
finely
D D 2
COMPARATIVE ELECTRO-PHYSIOLOGY
404
more or
less per-
It is
of
raised, by long-continued
once more give normal response.
nerve
the
stimulation,
Fatigue
is
has
that
it
again
will
been
is
The
it, also, a
mechanical stimulus, say vibrational,
throws into activity the whole mass of tissue, not only in its
further
limitation.
superficial,
but also in
its
deeper lying
Now we
strata.
may
is
have
less excitable
even, in fact, on
It is to
excitatory
influence
affecthig
gradual percolation
the outmost
tissue
first,
its
be
and only by
Owing
to
and the
highly excitable,
response.
We
effect in
varying
We
is
all
alike liable to
undergo periodic
variations.
may
Under
be ex-
to
positive,
make
is
3een
fact that
an
Again,
usually give
positive,
though the
first
by
to light,
Even
405
in
The same
sponse of growth.
If the
growing organ be
in
a normally
tion,
of growth.
retardation
If the tissue,
however, be in
tremely
an
sub-tonic
ex-
con-
by
increasing
the
internal
is
to
hancement, of the
of growth.
rate
grow-
If the
we
sub-tonic,
have a preliminary
positive, or
enhancement,
of
the
This
is
lowing
or
rate
retardation
of
growth.
seen in the
record,
fol-
which
Fig. 239.
Crinum
Light
Continuous
growth.
lily, in
This record was made with the Balanced Crescograph, where the normal rate of growth is recorded as a
(fig.
239).
horizontal
rate
line,
enhancement or
negative variations,
by down-curves.
It will
be noticed that
COMPARATIVE ELECTRO-PHYSIOLOGY
4o6
the
first effect
a positive
response, followed
negative.
Continuous stimulation
is
to oscillatory responses.
exhibits galvanometric
Each
positivity.
by exposure for
five seconds,
was in this case
applied after an interval of two
stimulus,
minutes.
should be mentioned
It
Fig. 240.
Photographic Record of Positive Response
of the Petiole of Cauliflower
to Light of Five Seconds'
Duration applied at Intervals of Two Minutes
multiple
of
response, as
growth
the
in
phenomenon
is
For example,
The
effects studied
upon
consisted
up
to the
of single
light.
Crimim
observed
lily
in
the
in
fig.
case
239.
The same
of motile response.
in
which the
leaf-
lets are
now
the
to
present have
responses induced by
response
positive
it
direct sunlight.
record of which
is
represent excitatory
given in
fig.
241.
was
of stimulus, a
The up-curves
here
leaflets, as
the result of a
leaflet
is
half,
and
in
is
407
also a relatively
The
drives the water inwards to the interior of the plant, and, the
loss of turgor in the
we
But
leaflets.
complete
in
The water
pulvini,
Under
returns
is
to
set
the
turgid.
more turgid
is
Ex-
brought about.
posure to light
in
this
condition
Fig. 241.
Multiple Mechanical
of
Leaflet
of
Biophytum under the Continuous Action of Light
Response
leaflets
by a
series ot
And we
in
consequence
impressed upon
acted alone
its
after-effect
it
is
COMPARATIVE ELECTRO-PHYSIOLOGY
408
but
it is
maxima
We
excitations
electrically.
Fig. 242
is
Bryophyllum sub-
leaf of
jected
the
to
continuous
We
action of light.
see here
an alternation of phase
in
This
by
positive in each
throughout the
constitutes
series.
parallel
Fig.
is
24 1, masmuch
as, Owing
tO
incomplete
rc
^
covery from each negative
m
.
response
fig.
But
this
equal,
and
for the
We
have seen, in the chapter on Multiple and Autonomous Response, that these effects are due to the absorption
of an excess of energy. When this absorption is great the
energy
may
find
an outward
expression,
An example
Desmodium
leaflet (fig.
141)-
its
record taken.
It will
be noticed that
under
which
persisted for a
sation of light.
time as an
after-effect,
were
initiated,
And
after-effects.
in this
409
connection
which
light
The
electrical
after-effect of
tissue,
tion
The
first
sues
response of galvanometric
nega-
tivity
under
short
light.
When
exposure
to
in
sequence
minus-plus
of
niinus-plus-
Confining
h).
pairs
243),
first
two
of
stimulation
Type
light
wc observe
that, in the
first
attains
its
maximum
at
b^
base
may
line,
arrive at the
is
here
after
mum
original
positive.
direction, a!
this phase,
The maximum of
a\ we shall designate
',
Withdrawa
go beyond
{- -^ - ^)
maximum
the
I.
the
action of light,
phasic alternations are a h,
During
c.
it.
as
maximum
positivity.
Under
now changes once more
It
to negative,
reactions
as
becoming
out, in
and so
on.
effectively
(p.
COMPARATIVE ELECTRO-PHYSIOLOGY
4IO
may
be so increased
as to
Hence,
such cases,
in
negativity
of
maximum
positivity,
of
gradually
is
moment.
If the
stimulus be stopped at
b,
the increasing
internal energy
and natural tendency to recovery will conspire with each other, and the after-effect will, generally
speaking, be b a'.
That is to say, this effect is the same as
that of natural recovery.
When
the second or positive phase has reached its maxia\ the response under the continued action of stimulation is once more reversed to negative a' b\ as we have
mum
At
seen.
this point
by the negative.
If
is
lus
unopposed,
itself
will
an
after-effect of
it,
I.)
seen
(fig.
in
244),
the
should thus be
following
(fig.
243).
balanced
series
That
of
-H
),
the dotted
photographic
records
the after-effect.
The
after-effect
The
We
(</).
II.),
exem-
here
at the
by specimens which
( +
...)
end of the
first
negative
is
The sequence
Insomeinstances,
if
411
phase,
we
after-effect.
obtain a small
The sequence
...).
curve
is
Turning to Type
teristic
response
to
III.,
short-lived
stimulus of
light
characis
by
continuous action,
quence represented
As
may be expected
to
be in the
se)
COMPARATIVE ELECTRO-PHYSIOLOGY
412
Type
tive
L,
we may expect
element
The
demonstrate
its
maximum
in this case,
will
by
presence
seen
is
over-shooting
a"
of
nation
under
1/
Diagrammatic Repre-
During action of
a' b' , b'
a"
are
alternations
(
-f
a\
b,
)
is
maximum positive,
the maximum nega-
fig.
maximum
is
246.
at
nega-
overshooting
direction.
(fig.
stimulation,
part of
now stopped
Stimulus was
tivity,
Type
first
Fig. 245.
cauli-
plus-minus-plus-niinus^
continuous
seen in the
specimen
of
This
245).
the
in
the
in
(fig.
The
(246 and 247).
taken was the petiole
i"
minus.
opposing positive,
its
negative direction
on the with-
drawal of stimulus.
at the
to obtain the
247),
in
In
the
negative
next
the
figure
is
here the
and
tive.
a'
Withdrawal of stimulus
reversal
a'
at
point
of
causes unmasking of negative
element, which is exhibited
by the overshooting of the
curve in the negative direc-
its
after-effect
lus
was
The impinging
that effect.
in this
at the exact
stimu-
of
maximum
Type
III.,
we
negativity,
formula of
(-f-
...).
classified as intermediate, or
Type
II.
plants to light
is
that
the electrical
413
response of
The
various
effects
observed
in
we
to be exhibited
response.
In
Fig.
Continuous
Type III.
The stimulus
of light was in
these cases withdrawn exactly
on the attainment of the
negative maximum.
Dotted
portions of the record exhibit the after-effect.
highly excitable tissue the direct effect gives rise to galvanometric negativity.
The
COMPARATIVE ELECTRO-PHYSIOLOGY
414
citable or in
sub-tonic
As
in electrical response.
in
is
the
The sequence
in the electrical,
observed
the
first
formula
is
may be(
may
be
classified
( +
its
direct or after-
-I-
-f-)or(-f-
).
results
is
continuous stimu-
continuous stimulation,
Taking
pre-
The
condition or fatigued.
-f
!).
...)
or
the formula
4-
tissues, the
-...).
And
in the third
it
type we have
CHAPTER XXVIII
RESPONSE OF RETINA TO STIMULUS OF LIGHT
Response of retina
Determination
of true
of optic nerve
differential excitabilities
current of rest
Determination
Oi
those
vegetable
in
excitations in
tissues Three
human
retina
The
types
Binocular
human
of
after-effect
Alternation of Vision
many
investigators,
chief
and
earliest
among
these being
results obtained
the later researches of others; but while their general observations are fairly concordant, the
way
in
Observed
results, in fact,
is
tissues.
nerve or
is
For whereas
by
excitation
The
subject
much
is
very
positive
and negative.
tions are so
named
in
varia-
COMPARATIVE ELECTRO-PHYSIOLOGY
41
we have already
which, as
very definite or
found to be reversed.
The
(i) What
rest, and do
which have been observed really fall
head or not? (2) What is the galvanometric
ing questions
is
under
this
the retina
that
May we
the excitatory
phenomenon
is
to say,
to,
is
in
?
induced by light
in
vegetable tissues
.'*
Of these we
In an eyeball which
is
is
isolated entire
through
In an isolated nerve-retina
retina to nerve.
These observed
be true currents of
due to injury
rest,
currents, however,
in preparation.
may
we have
In such cases
not
after-effects,
seen
We
tive.
from the
less
we have
first
to deter-
We
excitability
of a
under
preparation can
such conditions the resulting responsive current flows from
be determined.
saw
also that
The
ment used
in this investigation is
417
experimental arrange-
shown
in
fig.
In such
248.
an experiment, equi-alternating shocks are g'iven to the preparation by means of a secondary coil included in the circuit.
The
previously by a potentiometer.
may
be balanced
is
^.
the two.
of
such responses.
be
noticed
current
Fig.
more excitable of
249 gives a
Fig.
is
the
in
also
will
It
excitatory
the
that
series
same
direction
Experimental
248.
Arrangement
current of rest ;
sponsive current.
R,
re-
varia-
it.
From
cornea,
it is
is
it
its
excita-
tory negativity, which will persist for a time and slowly disappear.
Owing
That
more
this current
E E
COMPARATIVE ELECTRO-PHYSIOLOGY
4i8
it
conclusion
was able
to verify
by
away
carefully dissecting
making
connections
with
the
Under
the
these
was
resting-current
true
and was,
detected,
conditions
ideal
as expected,
Fig. 250.
ment
Experimental ArrangeDemonstration of
tion
true
resting-current, or merely
an excitatory
is
pro-
in
the
after-effect,
for
Differential
Excitability as be-
R, re-
sponsive current.
ceeded
to
determine,
was found to
flow
to
the
nerve,
thus
proving that the retina was the more excitable (fig. 250).
The first series of responses in fig. 251 gives a record of these
effects with a moderate intensity of stimulation, while the
twice as great.
nerve-retina preparation
after-effect
of
is
preparation
which
is
inseparable
tissue.
from the
// zvill also be
injury-current^
is in
and thus
example, the
retina
is
electrical, the
the
419
same
constitutes
light,
such
by induced galvanometric
negativity.
This at once
lation in general
is
we have
we might
it
tissues,
tissues
is
not justified.
E E 2
COMPARATIVE ELECTRO-PHYSIOLOGY
420
As,
in
urged that
the
retina
is
it
may
exceptionally sensitive to
be
light,
may
it
insensitive as
is
It
distance from
them of
The
12 feet
have myself
its vicinity,
was thrown
movement.
shall, in
unmistakable manner, that, when exposed to light, it underIt is now theregoes a change of galvanometric negativity.
fore necessary to
show
change of the
tive
then,
is
the
way
in
negativity comes
is
What
retina.
which
due
really
this
to appear
opposite-directioned
clearly that
currents
has to be demonstrated,
as a positive
of
rest
in
variation
the
of the
eyeball
and
As
we have seen that the existing injuryfrom nerve to cornea, and that this undergoes a
positive variation when the nerve is stimulated in any way.
This is because the added negativity induced in the nerve
gives rise to an increase, or positive variation of, the existing
Now, when light falls on the eye it acts
current (fig. 248).
on both the cornea and the retina. The former, however,
nerve and cornea,
current
is
is
moderate a stimulus
and its excitatory
the optic nerve, which
as that of light.
condition
is
rapidly transmitted
is
excited,
to
thus
galvanometrically
arrangement, then,
The
negative.
under such
observed
variation
positive
42
so-called
an experimental
is
In
contacts
current
are
is
we
shocks,
When
have seen
that,
and retina
both uqvwq
by equi-alternating
simultaneously,
excited
electrical
differential, is
of stimulation by
light,
however,
it is
excited.
In the case
is
directly
in the retina
injury-current
Hence,
250).
(fig.
negative
supposition
positive,
effect.
the response
that
normal excitatory
involved
of the retina to
in
the
light
was
and thus of opposite character to that of other exis thus seen to be due to a misinterpretation
citable tissues,
of observed results.
It is
be
understood
quoting these
as
'
negative,'
results, therefore,
and
shall
by
different
'
vice
versa.
When
and in parentheses.
Another observation which lends independent support
to the view that the retina exhibits the true excitatory reaction,
is
found
and Engelmann
retraction under
the fact
in
that
light.
noted
by Van Genderen-Stort
the optic nerve contains not merely sensory, but also retino-
motor
But
fibres.
shall
show that
it is
itself, which
under stimulation (cf
will be shown in a later
excitatory contraction
324,404).
COMPARATIVE ELECTRO-PHYSIOLOGY
422
variations
shall
be able
by
anticipation,
of nerve and
which
may
refer here,
be given in detail
concerning the mechanical response
results
XXXV.
Chapter
in
to
will
variations.
In
here,
We
reversed
to
excitability
But
positive.
is
Thus we
obtain,
in
a somewhat
sub-tonic
tissue, the
following results
(i)
Under a
level,
whose
response
positive (expansion).
(2)
value
is
Under
at
efficiency.
first
In consequence of this
we
obtain a
first
phase of
is
positive, followed
423
by a second which
is
The second
(3)
quence of
The
eye
will
cases.
in conse-
(expansion)
fatigue.
all
I
experiments which
will
jecting
249).
(fig.
preparation
have already
by sub-
responses, obtained
to
had
becoming
It
it
occurred to
me
is
due to
In dealing
it
ought to
in its
I
say, dilute
depressed condition.
As
NagCOg
to the nerve
to
nerve
The normal
(fig.
238)
responses
under
current from
equi-alternating
shocks,
These abnormal
responses, then, rectifiable to normal, are those due to subtonicity.
And under fatigue finally, induced by longcontinued, or over stimulation, I find the normal response
COMPARATIVE ELECTRO-PHYSIOLOGY
424
effects
factory explanations of
already demonstrated.
'
have
sum-
In light, frogs
hours to the
which
results
full effect
i.e.
of daylight
the
for
positive fore-swing
is
Since
means, as
have already shown, the true excitatory negative,' and the
negative
variation
exposed
for a long
This, then,
the
conversely,
means
time to
'
The
of
positive
is
occurrence
fatigue,
which
have already
With regard
positive phase
to this
it
may
here referred
to,
positive response
which
'
is
by the excitable
The former
is
translation), vol.
ii.
pp. 474-477.
is
425
(- +
-)
Now
in
stimulation,
highly excitable
we observe
tissues,
under instantaneous
similar, are
is,
as case (i).
in
(p.
422)
in fig. 240.
We
come next
to the question
whether or not we
may
table
hitherto detected lay in the inevitable depression of excitability in the isolated retina or eyeball.
fishes,
COMPARATIVE ELECTRO-PHYSIOLOGY
gave as
after-effect multiple
its
it
We
252.
fig.
is
end of
As we have
stimulus of
is
light.
the expression
and
for a time,
since, as
on the cessation of
show
light, will
In
after-effects.
my experiments
on the
itself
is
it
already been
has
The
effects
found,
three
said,
and
after-
of these related to
first
normal responses
^^"
^i^^A^^T^?^'''
cord
of Multiple Response
of Retina of Frog under
Continuous Action of Light
^
^''''
was thus
the
cases,
'
\-
the
the
if
h).
In
stimulus
was
attainment
the
.^-
maximum
In
*').
on
stopped
was an increase of
after-effect
tissues,
such
.v
positivity the
positivity.
third
type,
of
^j-^*.^
immediate
The formula
with
sub-tonic
stimulation,
was
( -h
effects
I
),
was
shall
either
now
discuss in
or
some
.)
after-effects
427
and not
one which has undergone depression in conI was fortunate enough, however, to
in
sequence of
isolation.
is
When
out of water.
many
it
lives for
is
pithed,
hours.
its
heart continues
made
a preparation
of the eye
the fish
In
fish,
so exceptional that
254
is
it
light.
its
cessation.
Fig. 254.
It will
Fig. 255.
Fig. 253.
oft'
completing two
was even slightly
reversed at its maximum positive phase the light was withdrawn. The immediate effect was a sudden increase of positivity, followed by a series of after-effect oscillations.
In the
(
f-).
oscillations,
It
and
will
be noticed
that, after
maximum
the last
positivity,
namely,
COMPARATIVE ELECTRO-PHYSIOLOGY
428
under light
When
may
(fig.
obtain after-effects of
(__{_...)
or
tissues (cf
fig.
245
d\
when
the specimen
result
also,
(fig.
256).
shown
in the
formula
exemplified
In Ophiocephalus,
this
256,
was able
was
257,
its
to obtain
slightly fatigued
which
seen to be
is
).
Fig. 257.
Type
II.
1-
Fig. 256.
Response of retina of Ophiocephalus when
fatigued.
Fig. 257.
Response of frog's eye (Kiihne and Steiner).
sub-case of
Type
II.
is
represented again, by
reversing force.
may
light, there
is
-
)
slightly
),
a transient
we
of the antagonistic
influence of a
is
vegetable
in
Fig. 256.
we
in fig.
fig.
not so high,
II., in
was
This
).
Type
is
Phis, as
its
been taken as a proof of the existence of antagonistic processes of assimilation and dissimilation, rather than as due to
molecular. derangement b)^ external stimulus and
its after-effect.
however,
explanation,
its
it is
429
not necessary to
similar
substances,
in inorganic
effect
is
is
seen in
258.
fig.
We
owing
sequence of responses,
depressed
condition
reversed,
the
(H
effects
),
the
to
of the tissue,
formula
is
being
here
are
by
represented
-(4--^
).
certain fishes
(fig.
retinae
of
261).
the
question
in
isolation
of the
be modified
unknown extent by
an
of
and
responses
Bromide
the
The
the effects
preparation
of
Fig. 258.
After-effect
of Light on Silver
prepared speci-
'J
to
intact
that
eye,
itself
Multiple
But
its
exhibition
'
COMPARATIVE ELECTRO-PHYSIOLOGY
430
experimental
It is
object for
some
many
repeated
time,
times.
if,
sensation
will
is
owing
to
maximum some
exposure
its
Fig. 259.
with
after-effect,
Fig. 260.
Fig. 261.
(+ -...)
Response of petiole of cauliflower. Light was here cut
on attainment of maximum negativity.
260.
Response of retina of Ophiocephalus fish when de-
Fig. 259.
off
Fig.
pressed.
Fig. 261.
Response
of isolated
retina
of fish as observed
by
The
true
excitatory
its
'
the eye
is
was
by these
positive variation.
when
response
negative
Th^s
is
not
so,
however,
considerable time.
is
a positive rebound
with
is
;
con-
negative,
and these
more prominent.
noticeable.
If,
now
will
43
will
be noticed alter-
It
be due to some obscure form of fatigue, but the regular alternations observed clearly demonstrate them to be a case of
oscillatory
The demonstration
an
retina, as
after-effect, is
human
But
their
is
more difficult to prove. This arises from the fact that the
waxing and waning of the effect are so gradual as not to be
noticeable, unless against some definite standard of comparison.
constant
now prove
shall
light,
that,
These
want
sum
total of
I
the two remains always approximately constant.
have
been able to provide the necessary comparison-standards by
this fluctuation,
had
i.e.
if
maximum
the
is
But
moment
excitation
as the
have found
in
maximum
that, as
the one
regards excitation
moment
retinae, so that
the
had
excitation in
maximum
period,
effect at a
given
the other.
in
COMPARATIVE ELECTRO-PHYSIOLOGY
432
slit
left
When
left.
The
slits.
The
is
On now
262).
(fig.
closing the
is
here perceived in a very interesting manner, for the afterimage is not now the complete inclined cross. Instead of this,
Fig.
262.
Inclined
is
That
is
to say,
Composite Indecipherable
Fig. 263.
Slits
Word,
is
at
its
The impression in
Some very
two inclined
pose a word.
other.
open,
The result of
we obtain an
this is that, as
indecipherable
(fig.
due
to
super-
263).
closed, then,
owing
to binocular
is
RO ME RO ME,
of dark vision, as
and so
on.
In
433
this
curious instance one sees better with eyes closed than open
is
that of
demon-
The
stereoscope,
in
its
tinuous action
distinctness.
It
may be
greatest
is,
generally speaking,
Even
shorter
in
the
same
individual, again,
and
is
it
observer
Time
of
8 A.M.
12 noon
3 P.M.
day
.
3 seconds
In this connection
the period
measured
Time
Period
II P.M.
may
day
.
Period
5-4 seconds
5-6
6-5
of a single oscillation in
in frog's retina,
ten seconds.
6 P.M.
9 P.M.
it
of
a multiple response,
CHAPTER XXIX
GEO-ELECTRIC RESPONSE
Question regarding
pansion Crucial experiment by local application of cold Reasons
delay
in initiation of true geotropic response Geo-electric response of shoot
Statoliths
Due
to active contraction of
negativity
upper
side,
it
lateral
stimulus,
possible,
is
to predict the
of incident uni-
nature of
the responsive
in my work on Plant
movements of a plant organ
can be deduced from the simple law that it is the more
excited side that becomes concave.
But though this law is
sufficient guide in dealing with the action of a known external stimulus, yet the problem becomes much more obscure
when we have to account for any movement which occurs in
response to a stimulus whose seat is internal. An example
movement.
Response, that
of this class
is
wards
till
Thus, a shoot
curve up-
In connection with
First, is the
question as to the
cises stimulation
in
the actual
with gravity.
this
all
answer to
this stimulus, a
As
takes place.
regards the
stimulation
is
by some
the responding
ferential
cells.
weight-effect
way
definite
first
responsive curvature
of these,
it
in whicli gravity
differential effect of
According to
may be due
this,
to
may
be said
could produce
weight acting on
the necessary dif-
GEO-ELECTRIC RESPONSE
contents, whether of the sap itself or of those
like starch-grains,
in
it.
435
heavy particles
The
former, or
In
the
264),
(fig.
lation
is
the particles.
obvious
is
weight of
It
case of a
the
we
antago-
are
yet,
nistic;
this,
horizontally
which stimu-
the
that
produced on the
effects
in
spite
in
of
1
obtain a resultant
curvature upwards.
This
shows that the excitation
of one side must be
greater than that of the
On
inner,
outer,
tioned mechanical
Darwin).
effect
fact.
It is clear, then,
horizontally-laid shoot
is
due
We
shoot.
effective,
traction
if it is
is
the more
the curvature
is
effective,
to
That
it
is,
is
have
to 511.
F F 2
COMPARATIVE ELECTRO-PHYSIOLOGY
436
cold.
known
It is
diminishes or abolishes
it is
the" excitability
induced,
it
will
of a tissue.
If,
then,
is
Effect on Apogeotropic
Fig. 265.
of Cold on
mm
its
application
(fig.
265).
This
fact,
of response by contraction,
effect
This question
is
fully
side.
concordant with
GEO-ELECTRIC RESPONSE
437
by means of
tion
examina
electrical response.
ticles
is
way
such a
the
that
side
sure
the
opposite
The magnetic
side
functioned
case
(fig.
266).
particles in this
as
Diagrammatic Repre266.
Fig.
sentation of Experiment showing
Curvatui;e Induced by Unilateral
Pressure Exerted by Particles
F,
virtual
attracted
by
electro-
statoliths,
general
organ
sides
ever,
is
the
of
symmetrical.
is
Asymmetry
bution
of
distri-
induced, how-
when
the shoot
is
and
horizontally,
laid
on
distribution
two
the
,,
Fig.
267.
Induced
in
long.
this,
we
is
at first
But, instead
downwards
and
COMPARATIVE ELECTRO-PHYSIOLOGY
438
it
not
is
till
which
sometimes
is
ment.
Fig.
this prelimi-
minutes before
its
ture upwards.
Thus,
in a
Fig. 268.
The up
its
own
curvature due
to apogeotropic action
in this
The
way
is
a ten-
in
Scape of Uriclis
Lily*
laid horizontally.
effect of these is
downward
curvature.
The
fact
own
its
appearance.
We
will
may
crucial
account
its
for
experiment
be given presently.
have seen
in
internal excitation
is
GEO-ELECTRIC RESPONSE
439
The apogeotropic
that side.
directly
curvature
is
indirectly
by the
so,
we may
effects
that
And
it
is
that
is
side
excited
will
found that it is in fact the upper side of the horiwhich exhibits galvanometric negativity
zontally-laid shoot
the
positive turgidity-effect
had been
show galvanometric
posi-
tivity.
But
this result is
may
occur.
We
And, before
take an erect
As
are
now
exhibited
sides,
COMPARATIVE ELECTRO-PHYSIOLOGY
440
be galvanometrically positive
relation
in
And
below to above.
finally, in
we have
the
that
is
now
Thus, the
side
electrical
ultimate
this
since
it
is
that
that
find
in
And
downward movement.
this
becomes
galvanometrically
indication, like
the
stage
we
negative.
mechanical, gives
the
characteristic
response of the
Fig. 269.
In the first phase of response, the current is from the upper surface to the
After fifty minutes,
lower, the upper being galvanometrically positive.
.the excitatory geotropic effect reverses the current, which is now
-' ascending, of from below t6 above, the excited upper-surface being
(Compare corresponding mechanical
galvanometrically negative.
.
record,
plant-tissue
effect
and
is
268.)
fig.
now
exhibited
and
concavity,
The
stimulus.
gravitational
to
excitatory
electrically
by
the
galvanometric
These
269).
similarities
268.
We
be seen on comparing
have thus
seen
that,
owing
this figure
to
with
secondary
how
GEO-ELECTRIC RESPONSE
441
have
1
which at first appears
It has been shown that
very great, in the following way.
the state of excitation, even when all mechanical expression
of it is restrained, may be detected by galvanometric negaThose secondary effects, due to mechanical distivity.
turbance, which mask for a time the excitatory effect of
gravitational stimulus, may thus be eliminated completely by
restraining all movement of the shoot.
The problem thus
placement of the hypothetical
been able to overcome
statolithic particles.
this difficulty,
Fig. 270.
response
lily
in
that mechanical
The next
completely restrained.
is
moment,
shall
way
such a
now
point
is
to
to the stimulus of
electric response.
method by which
that when
acted on symmetrically by the force of
It is clear
two
lateral
points
are
When
again
the shoot
is
laid
acted on sym-
by the force of gravity, and there is thus no differbetween the two. But if the shoot be now
rotated on itself, so that one of these points is diametrically
metrically
ential action as
COMPARATIVE ELECTRO-PHYSIOLOGY
442
above, and
the
other below,
differential action
periment
In
first
Fig. 271.
lateral.
Owing
to
The
270).
A and B to
was now no
(fig.
be at
be
upper
will
sides, the
symmetry there
now
a horizontal
The galvanometric
record was
The specimen, on
through 90. The statolithic
its
line.
support,
was
particles
next quickly rotated
were thus displaced, falling on the inner tangential wall of
An
the upper side, and outer tangential wall of the lower.
electrical response was perceived in about one minute, which
went on augmenting with time, the upper side being in-
(fig.
271).
The
fact
GEO-ELECTRIC RESPONSE
443
It will also
CHAPTER XXX
DETERMINATION OF VELOCITY OP^ TRANSMISSION OF
EXCITATION IN PLANT TISSUES
Transmission of excitation in plants not due to hydromechanical disturbance, but
instance of transmission of protoplasmic changes
Difficulties in accurate
determination of velocity of transmission A perfect method Diminution of
conductivity by fatigue
Increased
in
rise
trans-
velocities in
When
tion
is
and negative.
is
is
leaflets.
found to give
is
This
rise to
motile responses
we have
it
actually nothing in
is
For
in the animal.
to be
certain
common between
the two.
if
in
movements of many
vital
manifestations of
is
merely
and the
exceptional cases
superficial.*
it
is
easy to
ii.
p. 32.
445
by
theless found
Pfeffer to
was natural to
infer
from
of the pulvinus
It
anaesthetic, so
must
was therefore
inferred that
It
is known to take
by the propagation of protoplasmic changes the apparent conduction of excitation in a plant was purely hydro-
in
animal
tissues,
place
mechanical.
But
in
shall
demonstrate again
bility
and conductivity are related phenomena, yet the variaone is not necessarily identical with that of the
tion of the
other.
Thus a
sufficient to
certain degree
of anaesthetisation
may be
may
is
Mimosa
is
In
been
killed
by
scalding.
But
it
is
extremely
difficult
to
It is
is
totally abolished.
Only
after
COMPARATIVE ELECTRO-PHYSIOLOGY
446
interior tissue
really killed
is
thoroughly done
by
scalding,
and unless
is
it
this
cells
is
may
There
is,
conduction^
when transmitted
dead
area, of imparting a
across
the
up excitation de novo
in that
shall,
brought about
in
is
same transmission of
The
responses,
electrical
again,
afford
us a crucial
in plants
by means
disturbance,
its
electrical sign
galvanometric positivity
the
contrary, that
sign
The
is
distinct characters
by the application of a
selective
animal, that
are
(p. 66).
So
propagated
true
in
indeed
is
it
that
isolate certain
whose responsive
from those
peculiarities
447
are indistinguishable
The determination
nerves.
of the
stimulus
at
velocity
of transmission
may be made by
be
of
applying a
interval
which
is,
and consistent
results until
successfully eliminated.
These are
(i) indefinite
changes of
the
point of application
(3)
unknown
the
of varying
on
stimulus
effects
intensities
of
velocity
of
transmission.
As
a result of investi-
can only be
regarded as a determinate
quantity
Diagrammatic Representation
Fig. 272.
of Electrical Connections for Determination of Velocities of Centrifugal
and Centripetal Transmissions
are exciting' electrodes, and L the
indicating leaflet.
A and B
is
found,
by causing
to
of
rest,
varying
in
COMPARATIVE ELECTRO-PHYSIOLOGY
448
In the
ten minutes.
exact
results,
temperature
it
possible to
various factors
which made
and, having
such
in
obtain
effects of
and
modifying
the
of
velocity
transmission.
Before proceeding to describe these results in detail, howit should be mentioned that, though the velocity of
ever,
transmission of excitation
is
constant in
the
same plant
Time
Distance
Direction
Centripetal
Centrifugal
22-5
45
mm.
mm.
Velocity
11-2 seconds
15-2
Specimen II
Time
Distance
Direction
28
Centripetal
Centrifugal
39-5
mm.
mm.
Velocity
15-2 seconds
17-5
2-2
mm.
,,
we may gradually
fresh
was found
direction.
to be i'88
rest.
velocity of transmission in
mm.
The
when
being
now
i*86
mm.
per second.
by a
The
series of five
experiments on the
449
same specimen. The distance through which the transmission was observed was 27 mm.
Table Showing Variations of Velocity of Transmission and of
Amplitude of Response with Increasing Fatigue
The
14-5
minute
^ minute
i6-4
17-5
half a
i4'5
2-5
i-o
minute's
-86 mm.
172 mm.
'64 mm.
'54
,,
,,
mm.
effect
rest
cent.
The
,,
specimen by 18 per
The
divisions
34
20
157
Velocity
response
14-3 seconds
plant fresh
3 minutes
2 minutes
It will
Amplitude of
Time
Intervals of rest
that
with
increased
velocity
In
way
this
increasing
it
was
intensity
of stimulus.
This fact is shown in the following table,
which gives the results of an experiment on a petiole of
Biophytum.
Table Showing Increase of Velocity with Increasing Stimulus
Specimen
The
I.
Centripetal Transmission
Time
Stimulus
01
j>
12
i6
Specimen
The
,,
11.
-,,
14-9 seconds
I4'4
,,
12-8
1-8
1-9
2-1
mm,
mm.
mm.
16
24
32
per second
,,
,,
Velocity
Centrifugal Transmission
distance traversed
Stimulus
01
mm.
,,
11-6 seconds
IO-2
,,
lO-I
9-9
Velocity
3-27
3*72
3-76
3-83
mm.
mm.
mm.
mm.
per second
,,
G G
COMPARATIVE ELECTRO-PHYSIOLOGY
450
With regard
cold
further,
still
it
found that
Thus,
to one-third,
one
in
and when
it
of temperature from
rise
doubled
was
that
case
this
in
that
at
the
30 C.
37
at
first
determined
C.
the
to 35
rate
The
temperature.
in
the
C.
was
centrifugal
direction.
Time
Temperature
30 c.
35 C.
37 C.
show
shall
again,
We
of anaesthetics.
7 "4
9-1
it is
sensitive
'
mm.
mm.
per second
,,
,,
shall
also
see, further,
ing as
'
is
37 mm.
effects of currents
same
Velocity
II seconds
5-5
4-5
Transmission of excitation, as
and
mm.
by taking advantage of the motile indicaby the leaf or leaflet, it is possible to determine
of transmission of excitation and its modifica-
plant,
tions afforded
the velocity
tions.
With ordinary
plants, however,
no such indications
we must
being available,
it
One such
is
obvious that
have described elsewhere as the Electrofound that the passage of the excitatory
tactile Method.
wave, even through an ordinary tissue, brings about minute
These give rise to pressure-variations as
form-changes.
transit.
It is
is,
excitatory
wave during
In this
tissue.
the
way
45
of detecting the
its
Electro-motive Method
the
galvano-
has been
leaflet.
is
motive
the
metrically negative.
is
state of excitation
variation,
tissue
It
at the given
response,
as
base being
is
on the leaf
its
The two
spots
is
lie
On now
it
will
-a distant point,
may
COMPARATIVE ELECTRO-PHYSIOLOGY
452
from
observed
the
between
interval
the
application
In this
trans-
for
manner
of
have
determined by
and of ordinary
me
and
plants,
in the case
for the
of sensitive plants,
purpose of comparison,
more
or less of the
same
it
will
be seen that
(a)
Velocity
....
Nerve of Anodon
Nerve of Eledone (observed by Uexkiill)
[b)
mm.
mm.
ID
5 to
,,
Velocity
14
petiole
:
per second
Sensitive Plants.
Subject
Wave
Animal.
Subject
Mimosa pudica
these
order.
some of
all
mm.
mm.
per second
,,
...
(e*)
mm.
mm.
37 mm.
2'i
3*8
Ordinary Plants.
Velocity
Subject
isolated nerve of
Ficus religiosa stem
Cucurbita tendril
stem
Jute
Artocarpus petiole
Fern
50
9*4
5
3*5
54
mm.
mm.
mm.
mm.
mm.
per second
,,
,,
,,
,,
453
occur most
protoplasmic
elements
it is
continuity.
It
evident that
which there
then,
clear,
is
is
it
must
greatest
that
certain
in
conducting medium.
leaf,
on the other
conductors of excitation,
remaining more or
stimulus
the
contain
less
localised
them.
in
fibro-vascular elements,
such as the stem, peduncle, and petiole, are for that reason
good conductors.
relatively
again,
is,
we should
as
expect,
much
than across.
now
shall
describe an important
its
may
means the
in
a thick peduncle of
nections, of
Mum,
B,
transversely
The second
point. A,
distance from
(fig-
it
C,
and
its
273).
If
we now take
CA
is
making
that point
direction of the
from A->B
first
is
earlier
than that
galvanometrically negative.
responsive
in the tissue.
A contact
If,
current,
The
therefore, will be
first,
COMPARATIVE ELECTRO-PHYSIOLOGY
454
point of transition,
may now
or of balance,
A',
that in the
transverse
direction
will
is
to
con-
then be, to
The
responsive currents.
CB.
the peduncle
longitudinal balancing-distance
cm.
Fig. 273.
Experimental Arrangement for Comparing the Relative
Conductivities in Transverse and Longitudinal Directions
c, point of application
A,
a',
of stimulus
B,
for
obtaining
balance.
It
may
In
animals there
known
are
channels of conduction
specialised
and
in
plants also
It
may
velocity of transmission
nervous channels
is,
of true
in
anticipation,
excitation
that
the
through these
mm.
This, for
fern.
that there
is,
different velocities
namely, the
Of
excitatory negative.
greater.
of trans-
In connection with
two
vegetable
455
is
also
is
great,
is
it
not
their succession.
even
in
animal
But
tissues,
a subsequent chapter.
shall
be able to demonstrate
in
is
wave
it
generally
(p. 59).
Burdon
Sanderson, in his determination of the velocity of transmission of excitation in Dioncea^ arrived at the exceptionally
it
has been
my
object to
demon-
The examples
shall,
however,
in
delicate
may
given
be
book on Plant
'
my
plants
will
means by which
be studied
in
plants
CHAPTER XXXI
ON A NEW METHOD FOR THE QUANTITATIVE
STIMULATION OF NERVE
Drawbacks
Modification
Response
of response by decline of
Positive
after-effect
Stimulation of nerve by thermal shocks
Enhancement of normal response after tetanisation Untenability of theory of
evolution of carbonic acid Abnormal positive response converted into normal
injury
negative after
Gradual
tetanisation
of stimulus-intensity
tion.
electrical effects of
excitation on the
such investigations
it
is
usual to
marked drawback
to
its use,
made
facilities
however,
lies
It
quantitative.
employ the
electrical
which
it
In
form
offers.
it is
liable to lead
of
sensitive
lating current.
In
some
cases
it
is
possible
to
series.
Under
we have
shocks,
If,
for instance,
we wish
457
is
due to
A and
B, it
is
This
latter.
negativity
.series
manner.
at
of
In
A.
records
The
274
is
seen
obtained
in
this
fig.
nerve.
They
after-
exhibit
also
the positive
effect.
With
reference
to
the
method of
commonly employed,
it
may
be
'^w^y..
said that the assumption that the excitability of the injured point is totally
abolished
is
not justified
for
have
after a lapse of
tends to recover
its
excitability
to
In such a case
different effects to
Response of
Fig. 274.
Frog's Nerve under
Simultaneous Excitation of both Contacts,
by Equi - alternating
Electrical Shocks, one
Contact being Injured
in the responses.
positive
after-
the gradual
appearance of
fatigue.
Under these
This appears to
fall
in the
me
conditions,
may
it
and
almost disappear.
explains why,
in
the
after
section,
is
It also
causing
COMPARATIVE ELECTRO-PHYSIOLOGY
458
renewed depression of
excitability,
necessary in order to
is
The second
due
effect
of the excitability of
The
the responses.
shown
in
fig.
274
B, is
may
when
is
The
B.
apparently liable
in
electrical re-
way
this
to great
variations,
object,
the
have succeeded
new mode
in devising a
of observing
uniform reponses.
by equi-alternating shocks,
not
is
by crushing.
rise to disturbing
when the
physiological
destroyed
is
as, say,
by
Thus
across
two
coil,
'
If the
nerve of a
leg
frog's
is
laid
applied
observed
it
to the
myopolar
tetanus
tract,
may
still
off
These unipolar
be
from
effects
may
459
in vivisection and also in experiments with the galvanometer, if not avoided by due
precautions. Hering has pointed out that in experiments
ductive of fallacies
in
which
most
no guarantee
against the overflow of induced electricity through the
interpolar part of the nerve into the galvanometer
This kind of unipolar stimulation is an
circuit.
obvious danger in all experiments on action-currents and
negative variation in nerve, while it shows what narrow
bounds restrict the intensities of current that may be
by a long
of the two
complete insulation
circuits is
our
is
Heidenhain em-
electrical.
was subjected
The employment
of this
mode
escape of current
which
electrical stimulus.
Though
it
inseparable
is
this
is
impossible to say
structure
so
how
far the
delicate
as
223.
is
ii.
pp. 222-
COMPARATIVE ELECTRO-PHYSIOLOGY
460
As
already shown
phenomena
platinum wire
mental
its
vegetable tissues.
in
may be made
tissue,
regards the
Thus a
latter, I
have
on excitatory
single
loop of
now
nome. The intensity of stimulus may be graduated in a predetermined manner by the adjustment of the heating-current.
Excitation may then be caused either by one or by a
summated
series of
thermal shocks.
As
regards
its
of stimulus
are
made
pliability
and
How many
quite unique.
is
by
possible of attack
its
means
difficult
will
form
problems
be realised
in
variations
nerve
that of
frog, for
and appropriate
meter.
The heating
amount of
example
the
electrical connections
current
excitation.
is
responses of animal
distal contact is killed
made with
the galvano-
The thermal
variation,
it
must be
any way.
The platinum
loop
is
is
the
mode
generally employed.
The
tissue.
nerve, as usual,
must be enclosed
in
be seen
will
and
in
fig.
a moist
in practice,
291.
shall
46
which
efficiency with
this
mode
of stimulus
may
be
When making
applied.
ditions.
These records
under
frog's nerve,
will
electrical
The
first
of these was
was diphasic
last
the second
positive.
This
From
It
must be
said,
accompany-
however, that no
such cases.
shall
effects are in
shall
now
a series of
give, in
fig.
In
275,
normal
negativity, given
condition.
COMPARATIVE ELECTRO-PHVSIOLOCV
462
its
The subsequent
is
amplitude.
The next
series of responses, in
galvanometric positivity.
It
276, exhibits
fig.
abnormal
may
supposed
as
not,
by
Dr.
For employing
and
more delicate
methods of record I have
the
nerve.
other
found
even
fresh
nerves,
under certain conditions, to
exhibit
is
in
this
effect.
this positive
general
to
degradation.
as
we
Neither
response due
any chemical
Instead of
this,
may
be
attributed
run-down of the
Enhancement of Amplitude
Fig. 275.
After-effect
of Response, as
of
Thermal Tetanisation, in Frog's
first
Bjrief
thermal tetanisation
applied, and
quently
the
are
here
responses subse-
under
obtained
stimulation
is
seen
to
original
be
en-
hanced.
the
specimen, a process
When
pressed specimen
again
with
energy,
it
or
it
latent energy
Nerve
The
of the
to
even
citable.
the
in
such a deis
supplied
requisite
becomes normally,
supernormally, ex-
The
first
part of the
following record
was
in a
(fig.
somewhat sub-tonic
condition.
responses in
the
it
will
QUANTITATIVE STIMULATION OF NERVE
463
diphasic.
be exhibited
the
in
positive, diphasic,
same specimen,
in
and negative
may
series
then
passes
sation.
first
The response
by negative
phase,
this
by
succeeded
where the
diphasic
positive followed
and
277,
positive.
by a gradual
into
transition
fig.
lastly,
is
of
series
We
come next
to the ex-
We
will
On
and conductivity
below par. We have
citability
will fall
also
seen
that
in
this
de-
be reversed to
With
regard
to
positivity.
the
duction of excitation
be said that
it
con-
may
this condition of
Conversion of Abnormal
276.
Positive into Normal Negative Re-
Fig.
Abnormal
positive
converted
on
into
response
normal
to
left
negative
tion.
when
same
the tissue
distance.
is
thus depressed,
It will
make
And
effect of
its
appea,rance alone at
COMPARATIVE ELECTRO-PHYSIOLOGY
464
We
may have
its
is
not in the
by
tonicity increased
ment of
its
excitability.
shall
demonstrate,
also
in
very sub-tonic
first
failed to
negative
in
is
positive alone.
Under
is
excitability
into play.
465
to
rise
an
amplitude of
increased
In a
275.
fig.
and the
effect is positive,
But under a
in fig. 276.
series
and the
(fig.
length
of time, and
Or,
277).
if
transi-
certain
record
seen in
is
is
fig.
276.
say that
positive,
which the
in
and
is
converted
After
increasing
this,
making an increasing transmission of true excipossible, gives rise to a diphasic, and ultimately to
conductivity,
tation
This result
is
analogous to
positive, diphasic,
and negative
gave
ever,
rise to
When
was not so
great,
we obtained
this,
the diphasic.
it
how-
F'inally,
when
only
the
abnormal
which
gradation by
fully,
partially,
positive
response
the transmitted
That
was made
appeared.
stimulus
or non-effective, to induce
true
excitation,
In the cases
HH
COMPARATIVE ELECTRO-PHYSIOLOGY
466
by
conductivity.
If,
is
no necessity there
to
also,
it
in
sion of
Fig.
to
shall
now
describe an
278.
Stimulus,
experiment which
the
negative.
Thus with an
identical
diphasic,
specimen we
or
positive
may
obtain
response,
by
467
distance
sive stimulations
gradually, and
thus
give
rise
enhance conductivity
to
The
in
Under varying
lie
cir-
different
method,
principal
object
will
in
be given
this
in
chapter
a subsequent chapter.
My
been to
the
has
nerve.
Its
wider applicability,
mode
in the case
prove
of stimulation of
of other related
II
CHAPTER XXXII
ELECTRICAL RESPONSE OF ISOLATED VEGETAL NERVE
conducting tissues Isolated
Specialised
nerve
vegetal
Method
of obtaining
electrical
animal nerve
{a) action
of vapour of alcohol
{d)
of ether
(d)
action of
ammonia
of carbonic acid
action
{e)
{c)
action
Effect
of modified tissue.
is
in the previous
the
is
not due to
It is
of animal
the case
Hence
fibro-vascular
stems and
tissues like
petioles,
to
which contain
be good
conductors
somewhat
localised.
ground
tissue
elements.
was
left
The former
excitation to
of
these
was
found
to
transmit
in the latter
the
469
away from
and ground
tissues respectively,
means of transverse
cut, or
by
The
of a hot plate
now
In
this
subject of conduction,
I
effect
of excitation to
of
stimulation,
being
perfect
and uniform.
this I was led
From
the
disposition of the
conductors
be
must here
particularly
Fig. 279.
well
COMPARATIVE ELECTRO-PHYSIOLOGY
470
or more.
Now
in
what
On
its
is
protoplasmic
continuity,
we
find
enclosed within
it
mainly consists
few vessels
in
in
itself
It
the centre.
may
external resemblances
sclerenchyma.
sheath-like
structure
to
those
tests
of electrical
It
may
be said that
and NepJirodiuin
for the
common maiden-
niolle
suitable.
is
possible to dissect
injury.
It is
half an hour, so as to
When
handling.
it
then placed
remove
is
about
due to
for
traces of excitation
found to remain
is
weather
which
specimen
it
in
ice-cold.
The
the same
nerve
that
is
moist chamber.
is
be placed
in
found to induce
47
form of stimulus.
have seen
nerve
in
of the most
may
is
perfect forms
as
we
same manner,
the
One
it is
that
is
to say,
be applied in precisely
by means of a platinum
the
nome
manner previously
closing
described,
an electric
circuit.
summated
the
still
greater amplitude
lies
is
sufficient, as
be obtained by
this
One
of the
and
For
plant nerve
may
such stimuli.
effects of several
less
A still
further characteristic
is its
of nerve
indefatigability.
A long
series of responses to
induce
little
or
marked
none.
fatigue, will in
Rapidly succeeding
an
effect
little
we
of such
little
fatigue.
long
After
re-
COMPARATIVE ELECTRO-PHYSIOLOGY
472
parallelism
be seen
is
The completeness
to be observed.
next chapter.
may
of this
the
shall, at
Et/ier, for
is
followed under
On
blowing off
In
restored.
fig.
is
effects of this
the
Fig. 280.
(a)
(c)
application of ether
Normal response
Enhanced response in first stage of
(S)
quent depression
of ether.
subsequent
effect of depression,
after
original condition
on the blowing-off of
the ether.
Carbonic acid
have the
is
effect, in
known,
the
first
in the case
of animal nerve, to
its
prolonged action,
is
seen in
fig.
281,
is
twenty minutes
chamber.
in
seen to increase
473
Fig. 281.
a,
(c).
normal responses
fi
and c, enhanced response during
action ; d and e, subsequent growing depression.
;
Alcohol vapour
in strong, or
first
long-continued applications,
The
of
effect
istically different,
tissue.
is
same
tissues
is
characteris
j^
the
Parallel
in fig. 282.
nervous or ordinary
that of nerve
nerve
ammonia on animal
WUUl
Fig. 282.
stage of
is
but
While the
little
affected
quickly abolished.
characteristic
difference
excitability of the
by
its
application,
would be exhibited, as
4H
COMPARATIVE ELECTRO-PHYSIOLOGY
Ammonia on
Fig. 283.
Ordinary
Note
walnut.
It will
ammonia
be seen from
here
is
fig.
practically negligible.
the
But when
same reagent was applied
to
the
in the excitability.
nerve
isolated
of
underwent
depression, followed by total
fern the response
abolition,
One
the
in
minutes
five
(fig.
teristic
course of
284).
the
electrical
the
in
three
Ammonia on
The
Plant-nerve
is
response, the
give a
mixed
same
nerve,
when
Thus,
been
highly-excitable
hibits
it
the
to
as
said,
its
has
while
nerve
ex-
normal negative
or diphasic response
of
types of re-
according
condition.
already
rapidly diminished
finally abolished.
response here
and
of
of
is
referred
chapter,
last
distinct
sponses,
Photographic Record
284.
Application
of Similar
Effect
Fig.
as
will
is
still
same
conditions.
records given in
the
fig.
be seen
will
The normal
285.
in
abnormal
is
475
represented as
more remarkable
down.'
'
is
In
modified.
are,
we have
as
case of
the
fresh
frog's
Fig. 285.
Photographic Record of Exhibition of Three Types of
Response, Normal Negative, Diphasic, and A])normal Positive, in
Nerve of Fern under Different Conditions
a period
of
The
tetanisation.
vegetable nerve
is
effect
precisely similar, as
of
is
on
tetanisation
seen in
fig.
it is
286.
found
become normal.
This
is
effects
in
287 we see
the abnormal diphasic response of vegetable nerve converted,
the case of modified vegetable nerve.
Thus, as
in the
In
fig.
normal negative.
is
converting
the
effect of
abnormal
into
COMPARATIVE ELECTRO-PHYSIOLOGY
476
normal response.
of nerve
we found
diphasic
of stimulation.
\}^]^WWVW\.\
UUJ^'
Fig. 286.
WV^-
The
first
to
sation, T.
responses
now brought
nearer,
the
distance
the
then
The
being
477
Fig. 287.
Photographic Record
Conversion of the Abnormal
of
Diphasic into Normal Negative, after Tetanisation, t, in Nerve of
Fern
Fig. 288.
Response
responses
seen
that there
is
negative
(fig.
288).
continuity of response
in
It is
the
thus
same
COMPARATIVE ELECTRO-PHYSIOLOGY
478
tissue,
as
intermediate diphasic.
From
the various
it
will
nerve
is
many
And we
shall also
we
We
shall in the
next
induced
in
excitability of vegetable
will
strictest
parallel with
in the animal.
CHAPTER XXXIII
THE CONDUCTIVITY BALANCE
Receptivity, conductivity, and responsivity
Necessity for
distinguishing these
responsivity.
We
know
that
when any
external stimulus,
it
point in a tissue
This excitation
at
some
is
distant point
may
is
thrown
then conducted
is
be made outwardly
by means of a
acted on by
and
suitable in-
There
response.
the excita-
first
or
Conductivity
and
thirdly,
the
Though
all
alike
of excita-
excitatory
which
effect
I
shall
it
is
nevertheless
480
COMPARATIVE ELECTRO-PHYSIOLOGY
have seen,
for
example, that a
rise
by
of temperature,
enhances
Thus, while a
may
rise
The term
responsivity.
'
'
So much
commonly used
is
But
indifferently.
investigation,
represented by R,
it
excitability
and responsivity
receptivity
this,
be decreased.
in
may
the
in-
But
conductivity.
is
it
In
differently.^
shall
for
show
important to
same external
the
'
following
excitability,
or
receptivity, will be
by
C,
conductivity
citability, or responsivity,
by
E.
series of
animal nerve,
on responsive exis
it
usual to take a
responses
after
of the reagent.
the application
By com-
this
each chemical
method
lies, first,
may be
The drawback to
by the addition of the
inferred.
of
the
resistance
undergoes an unknown
is
may
circuit
in
electrical
se.
It
is
in the circuit,
but
by
this,
by reducing the deflection, necessarily reduces the sensitiveDifferent specimens again cannot
ness of the method also.
but be characterised by slight individual peculiarities, and the
experimental arrangements therefore can only be considered
to be perfect when we are able to compare the effects of two
'
48
identical
observe the
effects of various
receptivity
and
still
when we attempt
agents on
to
conductivity and
difficulties to
be overcome when we have to study the property of conductivity versus responsivity or receptivity, or of receptivity
same external
The methods
agent.
by
their
connection.
shall
now, however,
describe
arise in
very
as
regards
citatory
reaction,
namely,
receptivity,
conductivity,
responsivity.
the result
un-
may
It
also enables us to
changes induced
in
and
is
comparing the
two
last in respect to
fig.
289
is
The thermal
specimen.
The
excitatory
wave
travels along
both arms of
c',
and
E'.
The
II
COMPARATIVE ELECTRO-PHYSIOLOGY
482
E and
when these
When
other.
contact
galvanometric
negativity
induced there
is
is
is
effect
of
which
is
left
greater
relatively
The balance
E.
is
the stimulator
the excitatory
e',
than at
are equal,
galvanometer indication
other, the
thus dis-
When
is
then,
say,
stimulator
the
is
downwards.
placed, on
metric
deflection
suitable
movement
will
be
up.^
By
of the stimulator,
and
to
fro
tremes, a point
give here
290) a record
(fig.
taken
The
E and
e'
recorded by galvano-
meter, G.
e'
is
near
too
being
excited.
indicate
left,
more
The up-responses
by
caused
the
relatively
over - balance
s
being too
The
the right.
record shows
of
exact
attainment
balance.
much
to
horizontal
It is to
condition of conductivity.
is
the
More
right.
movement, as seen
when
when
adjustment reduced
careful
far
up-
this
in
null, as
it
483
was
effect
Any
If the reagent
stimulatory character
but
if it
we
be of a
be of a depressing nature, E
consequently be down.
will
is
variation of excit-
two
It will
will
be rendered
rela-
points,
is
due simply
to
the
relatively excitatory or
is
completely inde-
trical
made
is
The
again,
it is
the influence of
E' are
if
the effect of
c,
E, will
conductivity.
in
COMPARATIVE ELECTRO-PHYSIOLOGY
484
conductivity-variation induced
by two
different agents
on
which
C'.
possible again
It is
to
changes
In
will
fig.
C'.
E,
The mode
of investigation of receptivity
be described presently.
we have the complete apparatus.
291
or vegetal nerve,
Fig. 291.
N, rests
The animal
on non-polarisable electrodes of
U -shape.
sliding-rod
S.
by balance,
these experiments
all
is
it
always made
is
for
485
to
be borne
in
perfect balance
at the
more or
In order to
show the
of excitability,
in
known
is
enhances excitability.
out on
to
the nerve
of frog.
be an agent which
FiG. 292.
Effect of
Na^COg
When
a dilute
Solution
Nerve
In this and following records the horizontal line at the beginning indicates
exact balance. The upsetting of the
balance in the up-direction represents either the enhanced responsivity of the right-hand responding
point E, or the increased conductivity of the right-hand arm c.
Down -curves represent corresponding absolute or relative depressions.
NaXOg applied to e is seen to exalt
the responsivity of that point.
plied at the
Fig. 293.
Effect of
CuSO^ on
Frog's
Nerve
The down
excitability.
point E on the right side, the upbalance upwards immediately indicated the
greater excitability induced by the reagent.
The longcontinued action of this reagent, however, showed that the
responsive
setting of the
enhanced
(fig.
excitability
was
undergoing a gradual
decline
292).
The
horizontal
previous
line
at
COMPARATIVE ELECTRO-PHYSIOLOGY
486
salts.
For
Fig.
shows the
this
294
CaClg on
effect of
being applied
at
right-hand side.
E on the
It
will
be
an
of
increase
excitability
that
becomes
after
considerable
minutes.
pression
is
FiG. 294.
ing
re-
to say, a de-
seen in
record
gives
way,
diphasic,
and
after-
balanced
first
by Application of CaClg
is
to
wards
to
down-response,
an
indicating
effect
of
de-
These two
of the basic moiety in inpression
at
E.
shall
agent
is
may
administered
shown by
at
E'.
In
the case
For
be compared.
this purpose,
one
it
was
487
FiG. 295.
record given in
Fig. 296.
fig.
NaCl was applied on e' and NaBr on e, the formula being E'^aciENaBrThe record shows the greater and earlier effect of NaBr at E in
causing relative excitation followed by relative depression.
right-hand side
which
is
E,
E',
a process
E
NaOl-L-JfaBr-
The
greater
and
earlier,
excitatory
effect
of
COMPARATIVE ELECTRO-PHYSIOLOGY
488
iUU^%ttttiW|tW[^^
Fig. 297.
c.
by a depression
variation
in
(p. 136).
certain
ductivity, but a
-5
of
excitability,
followed
isolated
followed,
conductivity,
489
long-continued action, by
after
depression.
C,
An
solution
was
-5,
and
in the
first
experi-
second 2 per
ticularly
inspection of
Fig. 298.
first
four minutes.
This increased
The
fig.
NagCOs
Solution on Conductivity.
c.
Note grow-
conductivity
inasmuch as
it
is
usually
associated
with
suspected.
increased
It
COMPARATIVE ELECTRO-PHYSIOLOGY
490
In the
excitation transmitted.
first
series the
The
fifth
is
shown by
how-
record,
ever,
direction, followed
greater.
The continued
became converted
also, so that
into a purely
this
diphasic ultimately
monophasic down-response,
298 we observe
the depression of conductivity by a stronger dose of 2 per
gradually increasing to a
cent, solution of
Here, again,
maximum.
In
fig.
we can
elements of conductivity
effect transmitted
that
is
we
two
at C.
In the
first
And
the up-curve
shows the later arrival of the less intense effect at the righthand contact E. It is thus clearly seen that conductivity
includes two different elements of speed and intensity which
may
I
not in
shall
all
cases be coincident.
will
demonstrate
ability
responsive excitability
than
depression
speaking,
generally
salts,
affect
salts.
conductivity
There
is,
491
But neutral
to
much
however, a very
where
to a
6-i
find a
much
greater extent
is
of
capable
striking
by the
comparative method of
demonstration
simultaneous variations
of
and
conductivity
order
In
scribed.
to
KI
of
effects
an
give
account
ments.
In
the
first,
liminary
was
KI
balance,
applied
at
c,
on
same
the
cess
left
end
e'
of
Responsivity versus
Conductivity under KI
Fig. 299.
KI on
effect of
being represented
COMPARATIVE ELECTRO-PHYSIOLOGY
492
In respect of conduc-
This
is
seen
more
301) to induce
(fig.
depression
of
responsivity.
We
said
chapter
vapour
in
was
the
previous
when
that
passed
alcohol
into
the
Fig. 300.
This
result,
however.
Responsivity verms
E'xaiCNai>
record
shows an
opposite to that of KI as
previously described, there being
now a relatively greater depression of conductivity than of
effect
responsivity.
was due
downward
direction
shows
been
in
the
positive
direction.
we must,
Nerve
Upsetting of the balance in the
depression.
have
Fig. 301.
Effect of Alcohol on
the Responsivity of Frog's
Such a
deter-
The
area.
The thermal
first
taken
killed
were
on a
distal
first
receptivity.
493
was
strictly
circumscribed
Normal responses
platinum loop.
The
were recorded.
Fig. 302.
fig.
302,
which gives
The
three normal responses to the left are seen to be exalted after application of ether on* the receptive point.
by
dilute alcohol.
The
trast
to
effect
this.
on conductivity, however,
On
applying
is
in curious con-
per cent,
solution in the
conducting region between the stimulator and the proximal
contact, a very great diminution of the conducting power
i
in fig. 303.
It may be stated here
a similar enhancement of receptive excitability, and
depression of conductivity, are found to be the result of the
action of alcohol in animal nerve also.
In the next experiment, it is the variation of responsivity under the action
is
observed, as seen
that
is
tested.
COMPARATIVE ELECTRO-PHYSIOLOGY
494
usual, a
responses as
fig.
Fig. 303.
The
It will
of alcohol was
was a depression
effect
excitation.
depression of conductivity.
Fig. 304.
a,
and
finally the
and
is
effect.
tive excitability,
it
495
and responsivity.
For the purposes of such a comparison, a new balancing
arrangement has to be employed (fig. 305). Here, two
tivity
electro-thermic stimulators
ex-
may
two
at
be produced
different
points
The
simultaneously.
vanometer contacts
gal-
E'
and
Diagrammatic Repre305.
sentation of Experimental Ar-
Fig.
rangement
for
Demonstration
of
and
Fig. 306.
shows
The photographic
the
relative
record
of
enhancement
receptivity.
E are made with two points intermediate between the stimuThe distance of one of the two stimulators is kept
lators.
constant,
at,
moved nearer
to,
receptive
formula
point, R',
now being
is
The
fact
that
is
obtained.
is
left
area, c, the
the receptive
excitability
is
downward
response.
COMPARATIVE ELECTRO-PHYSIOLOGY
496
Alcohol
R'aic.Ejiic..
is
variations
consists of a
of
receptivity
And we
find
E, the
here
in
comparison of the
and
responsivity.
CHAPTER XXXIV
EFFECT OF TEMPERATURE AND AFTER-EFFECTS
OF STIMULUS ON CONDUCTIVITY
Effect of temperature in inducing variations of conductivity
{a)
by Method of
the
first
effect of
we may
For the
generally
is
a distance.
of
the
arrival
afford
the
it is
sensitive
necesplant,
'
conspicuous motile
excitatory
wave
from
leaflet at
duction
and
if
we
different temperatures
experiments at
we have a means of determining the
carry out
successive
of temperature on conductivity.
effect
method,
effect
of
of these
as a
application of stimulus
known
first
known
Employing
this
Biophytum sensitivum.
It
lower-
ficiently intense.
With moderate cooling the velocity was
found to be decreased to about one-third. The effect of
rise of temperature was, on the contrary, an increase of
KK
COMPARATIVE ELECTRO-PHYSIOLOGY
498
velocity.
When
it
rose from 30 C. to 35
C,
for
example,
By employing
we
the electrical
Thus
in
30 C.
After
first
obtaining
of the
This one-sided
right
arm of the
Stimuli
On
rise.
by
was confronted
that a sudden
temperature per
se^
it
of temperature
variation
Hence,
act as a stimulus.
is
essential
These
in
will,
in
difficulties
were overcome by
499
Fig.
The
307.
will
be understood from
small chamber
fig.
308.
it
.^
COMPARATIVE ELECTRO-PHYSIOLOGY
Soo
intensity of which
cork chamber
is
The
is
within.
spiral, is
warmed, and
raises the
perature
is
trolling
regards
the
processes,
t~^
|1
LJ^J
'^'^^
interposed,
these
rheostat
air,
controlled
by
of the
adjusting
joint manipulation
rate
of
be
rate
may
The movement
current of
two
carbon
by which the
temperature
of
regulated.
is
of
heating-
has
circuit
rise
first
the electrical
of rise
By
the
the
of both these
of
temperature
perfectly
Fig. 308.
uniform,
for
A and
mately
As
1 C.
per minute.
already said,
selected a
B, the
a balanced record.
temperature of the
thermal
cell
At 48 C,
$01
was now being depressed. And at still higher temperatures it was found to undergo a very great depression,
as is seen by the abrupt downward movement of the curve.
It is thus seen that,' by means of the Method of Balance, this
very difficult problem of the variation of conductivity under
tivity
variation of temperature
I
shall
Fig. 309.
a subject of
much
importance.
It
difficulty
has been
in
Animal Physiology
its
length.
is
When
is not equally
such a nerve, with
From
is
subjected
to
it
is
stimulation.
the point excited from the motor organ, Pfliiger was led to
COMPARATIVE ELECTRO-PHYSIOLOGY
502
the
excitation
is
is
have depended
all
enhancement of
It
follows
nerve which is distal from the muscle, and in the neighbourhood of a section, must be ascribed to some other cause. In
reference to this Heidenhain, indeed, explained the greater
excitability of higher tracts of divided nerve
by the proximity
section.
artificial
distance travelled
by the
may
be.
The
For so
effect.
from increasing
causes diminution.
It
to
is
it,
this, as
a matter of
be remembered that
fact,
far
is
any response by
now
arises.
Why
point of section
It
cal
The
question
electro-tonic
variations
of excitability.
in turn
We
electri-
gave
rise to
shall
electrical current
see,
in
through
And
this
phenomenon
Now
any
'
injury,'
is
known
is
known
excitatory.
And
But
it is
is
section
That
explanation, however,
this
ments which
shall
opposite of
this,
the
nerve-current.
meet
does not
all
the
of
short-circuiting
internal
will
due to
that
section,
to say, of depression,
is
is
seen to be
induced.
All these various facts will be found fully reconcilable,
and
opposite effect
mental
overstrain,
would
result in
demonstration of
this
hypothesis,
we
shall
first
nerve.
In
the
And
stimulus.
first
since
we know
we found that
place
that
a cut acts
as a
excitability
section,
which
has
itself
the
nerve
is
relatively
inexcitable.
The
and
in the
enhance-
now
more
in
new and
interesting manner.
it
once
COMPARATIVE ELECTRO-PHYSIOLOGY
504
of
(fig.
310).
The
effect of this
stimulation
induce
were to
jOJOJ^
Fig. 311.
Fig.
310.
Excitability
The
obtain
Stimulus
to
upward
ductivity
Effect of Moderate
Enhancing
Stimulation in
Conductivity
and Excitability
stimulator adjusted
direction,
Photographic Record
Showing
depression,
and
of
excitability.
con-
The
fact
side,
and balanced.
however, there
application,
this
after
are
resultant
said,
that
responses
is,
then, in the
a positive variation of
it
were,
it.
In
same direc-
greater
and consequent
injury
same point
In this
wire.
case resultant
that
the
and
excitability
been
depressed by over-stimulation.
I
was
next
desirous
of
of the over-stimulated or
For
this
purpose
took
Photographic Record
Effect
of Excessive
Stimulation in Depressing Excitability and Conductivity
Fig. 312.
showing
first
ob-
The
(fig.
312).
and
E'
are
COMPARATIVE ELECTRO-PHYSIOLOGY
506
3een
to
to injury.
It also
by negative
For,
variation.
in
the
by
experiment,
was
is
further
seen
from these
experiments
that
the
It is
the effective
intensity
CHAPTER XXXV
MECHANICAL RESPONSE OF NERVE
Current assumption of non-motility of nerve
Shortcomings
of galvanometric
of
stimulation
nerve
Mechanical
Determination
Similar
effects in
response due
recorder
Kunchangraphic
Mechanical response of
mechanical response
transmitted
of velocity of transmission
eftects
of
Indeterminateness
records on
to
afferent
smoked
nerve
glass
Record
of
HAVE
fatigue.
monly
insisted
animal
tissues.
as
on,
Certain
view, however,
it is
fast distinction.
It
regarded as motile
of these are
From an
difficult to
would be
evolutionary point of
of the
simplest
some
in
tissues,
requirements.
Thus the belief
by physiologists that nerves exhibit no
motile response whatsoever^ becomes questionable, and is
held
so implicitly
After submitting
to understand
'
'
Nerves are
this change
them
there
is
difficult
it
to this,
how such an
irritable
is
no change of form
microscope.'
it
(Kirke's
in
edition, p. 105.)
COMPARATIVE ELECTRO-PHYSIOLOGY
508
cations themselves.
we
shall first
at present available
Since ex-
no
visible
it
certainty in the
become
results.
possible
to
If,
should
it
obtain
employed
without misgiving.
The second
limitation
character of the
which the
is
mode
electrical
of
response which
it
indicates.
For
proximal and
distal
the record
made being
But
it
is
is
artificially
effective in
how
It
is
finally
due to
depressed by injury.
has been
point
will
by
positive
is
due to
this or
some other
cause.
509
For
that case
in
tatory change,
it
this
arise.
uncomplicated
the galvanometer
is
change
posed
effects of tetanic
high sensitiveness
is
necessary.
In
a galvanometer of very
effects,
That of a
fairly delicate
it
can
and
in such a degree of galvanometric sensitiveness we approach
a limit which cannot be very much exceeded.
Returning now to our original question, we have first to
determine whether excitation causes any motile effect in
give efficient indications of excitatory effects in nerves
Under
nerve.
nerve
is
observation,
is
it
in
every
it
in
when the
increases in
of
the
contractile
stimulation
by
fairly
is
We
length.
way analogous
to
by no means microscopic.
the
The
of an order
(fig.
313)
under continuous
strong tetanising electric shocks.
This
response
of
nerve
the
cent.
at
magnification
It will also
which
state of
be seen that
maximum
this contraction
reached a
limit,
After this
we observe
a tendency
5IO
COMPARATIVE ELECTRO-PHYSIOLOGY
to decline,
owing
In
to fatigue.
some other
cases, moreover,
however,
it
desirable
is
avoid
to
over-stimulation
of the
tissue.
moderate stimulation, we have to employ a higher magniThis magnification, if made about 200 times, is
more than sufificient for all practical purposes, and the photofication.
a magnification of
Fig. 313.
fifty
nerve, however,
IS erve
under
apparatus used
the
for
purpose
glass
on
this
The
Kunchangraph
surface.
was the
I had already devised
employed
tissues.
This
apparatus,
recording mechanical
a
nerve-chamber, N
and
thirdly, a
as
adapted
response
in
for
the
purpose of
nerves, consists
of,
first,
photographic recorder,
(fig.
314).
closed
by a semi-cylindrical
Fig. 314.
Optical
51
COMPARATIVE ELECTRO-PHYSIOLOGY
512
its
When, on
subjected to
is
we wish
to study
clamped
the nerve
is
(fig.
Excitation
315).
lightly
is
after transmission
vening
in
the
nerve B
One
tract,
at
then induced
a,
through the
and
inter-
responding
portion
of the
c.
precaution which
very necessary
is
find to
be
the maintenance of
nerve-chamber.
Fig. 315.
Diagrammatic
Representation of Arrangement for Obtaining
Transmitted Effect of
Stimulus.
important
in
characterises
the
specially
is
year in India.
L, indicating
This
usual
means of
lever
and the
air,
in the vessel
w,
is
made
to escape
is
kept
in
chamber.
the
513
have been
much
as
study the
effect
is
The
the nerve-chamber.
show
will
either
will.
For
reagent at
it is
easy,
and
obtained
after-effect of the
re-establish the
first
or normal
arm of the
light
mirror
is
attached
lever
is
The
downwards.
give
responsive relaxation
of the
rise,
light in
it
nerve would
on the other hand, to a deflection of the spot of
The long arm of the lever,
the opposite direction.
the mirror.
will
be noticed,
movement
is
light.
The
responsive
recorded on a moving
photographic plate vertically below the mirror, and whose
of the
spot of
light
is
light.
is
in a direction at right
The photographic
plate,
or the film
slit.
The length of this is parallel to the
movement of light, and at right'angles to that
or film.
The advantage of having the plate
a narrow incised
direction of the
of the plate
COMPARATIVE ELECTRO-PHYSIOLOGY
514
vertically
candle
may be
in the
lies
fact that
lighted
such diffuse light could now find access to the plate would
But if the ceiling of the
be by reflection from the ceiling.
experimental room is blackened, or a black cover placed over
the nerve-chamber at a certain height, even this possibility
eliminated.
is
room
is
placing the
work in a fairly
arm of the
attached sufficiently short, and by
By making
obvious.
is
the
hundreds to several
be desirable
to subject the nerve to a certain amount of tension, and
this is secured by placing a small weight on the arm of
With high magnification, due adjustment, which
the lever.
thousands,
is
may
be obtained.
very troublesome,
lies
in
several
It
may sometimes
This
difficulty is obviated,
The adjustment
Optical Lever.
relation
to
the
light in
one direction,
moves it to the
This movement can be made very fine, and the spot
right.
adjusted to any part of the photographic field.
say to the
left,
while
its
left-handed rotation
If the
nerve-chamber, for
would
result
in
relative
Under
515
rod
have been able to take records, without the least disturbance from the adjacent electric tram line. As a matter of
fact, when the magnification required is only of a few hundred
I
of disturbance to remain.
It is
any source
unusual
One
care
necessary to avoid
is
precaution
from
arising
currents of
of disturbance.
errors
disturbance
The remedy
air.
for
this
is
obvious, namely*
This
is
is
necessary for
usual in
next chapter.
now
how
easy
shall
explain
logical variations
induced
in the
citability
well-known reagent
of the nerve is ammonia.
Its
response
is
nerve of frog.
seen in
fig.
316.
by
In
all
effect
on mechanical
of
records
In
fig.
Two
responses being
sciatic
after the
COMPARATIVE ELECTRO-PHYSIOLOGY
5i6
first
series are
found to be abolished
When
highly diluted
with
temporary exaltation.
In the next figure
be abolished.
seen to
is
The
strength
of application which
brings
about
find to
this abolition
Photographic Record
FlG. 316.
of Effect of Ammonia on
Mechanical Response of Frog's
Nerve
Photograph ic
317.
Record showing Abolition
of Mechanical Response
of Frog's Nerve by Action
of Solution of Morphia
Fig.
And
it is
this
to
strong application of
shall
plays
many
tissue
is
effect
preliminary effect
318.
We
interesting features.
its
fig.
51:7
are induced
one of these
is
of stimulus, and
absorption
other
the
The
the
is
expenditure
former, as
we have
The
and expansion.
latter,
the
In
tion.
excitatory
of
generally
We
have
may
negative or contractile
be unmasked
first,
in
effect.
two ways
either of
or,
second,
altogether.
first
say
diphasic,
converted into
positive
followed
by
the response
In the latter,
negative.
becomes
is
of these cases,
by the suppression
of the negative.
An example of this
unmasking of the positive element, by
positive,
P'iG.
318.
Photographic
Mechanical Response
of Frog's Nerve after
Application of Alcohol
am now
about to describe
is
interesting from the fact that effects parallel to those there seen
in
specialised
effect,
animal nerve.
moreover,
is
it
shown
The unmasked
electro-positive
COMPARATIVE ELECTRO-PHYSIOLOGY
5i8
found
in fig. 319.
It will
first effect
citability,
hour.
this,
Fig. 319.
application
interesting
reaction
made
its
appearance.
The impinging
response.
effect
519
tion of stimulus, as
and
this followed
Each of the
successive
stimuli
sequence.
It
action of chloroform,
how
regularly in
We
shall
abnormal
positive, diphasic,
and
nerve.
The
excitability.
will
from
its
result of this
depression
of excitability
efficient,
to
evoke the true excitatory reaction of galvanoThe absorbed stimulus will now induce
metric negativity.
The
depression of conductivity
the
requisite
'
'
As
COMPARATIVE ELECTRO-PHYSIOLOGY
520
a result of
negative
this,
response, through an
The
still
ment
of
tetanisation,
normal
the
about
bring
to
negative
response.
tissue,
jecting
we
it
typical stages
(i)
thus,
and sub-
obtain
four
enhance-
the
Starting
which
finally
becomes
(4) the
enhanced negative.
In studying electrical response, both of animal and vege-
tal
shall
simpler
still
is
applied
directly
its
opposite
expansion.
With regard
that this
is
to
abnormal response,
condition, with
its
staleness,' or the
with
its
On
finds
inde-
On taking
pendent support of the strongest character.
even the freshest specimen, I generally find that its
responses at first are the abnormal positive. These gradually
pass into moderate negative through diphasic. This is due
to the raising of the tonic condition
by the absorption of
52I
so
much
its
tonic condition
unprecedented magnitude.
specimen, in
may
fact,
which was
be brought to
any state of exalted excitability desired, with concomitant
increase in amplitude of response, by merely subjecting it for
at first almost
irresponsive,
generally
'\
^\^
fY\
\\1
Trrn
av
\\
Fig. 320.
abnormal positive
First series,
second
shall
now
experiments.
describe
in
some of the
detail
principal
took
intervals, in
testing stimulus
The
a visual
demonstration
The
variations seen
record of these
is
in
of the
given in
the
responses
variations
fig.
320.
in
The
thus
give
excitabihty.
responses in
COMPARATIVE ELECTRO-PHYSIOLOGY
522
the
first series
to say,
The
tissue
therefore,
exhibited the
still
shocks
longer
of
This gave
to a short-
rise
down-
wards,
succeeded by large
After the
second
of the
cessation
the
tetanisation,
absorbed energy
seen to
is
staircase increase.
seen in
where the
first
diphasic,
but
responses are
the
positive
ing
also
responses.
be
This
should
It
noticed
that
is
the
tnuch
fact is important,
phenomena
The
effect
response,
condition
is
illustrated in a
record given in
fig.
is
still
not yet
more
in
normal
maximum
tonic
manner
in the
striking
of frog's nerve.
523
responses
series of
first
was purely
positive.
have
shown how
already
That
in
is
under
On
tetanisation.
every respec
in
of the vegetal
nerve to
similar
characteristics
its
arriving at this
successive
illus-
mechanical response.
second
stage of
undergo enhance-
responses
diphasic
response,
ment
in
This
in
321, the
the
first
two
records
given
side
is
is
sufficiently
by
side
in
of frog's nerve,
in
that
of fern
intensity of response,
we must,
in
We
have seen that in consequence of the absorption of stimulus, not only does the
abnormal positive phase disappear, giving place to the
normal negative, but the subsequent negative responses
themselves also show an enhancement in a staircase manner.
I
give here
(fig.
ment.
From
it is
COMPARATIVE ELECtRO-PHYSfOLOGV
524
We
that,
by the
direct
mode
of investiga-
tion afforded in
the experimental
In
method
appropriate
condition
are
the
to
not only
is
tonic
superposed on parallel
Thus
responsive change of
the
after tetanisation,
The power
of
is
howis
transmitting
Fig. 322.
Photographic
Record showing
Stair-
case
Effect in Mechanical Response of
Frog's Nerve
transformation,
from
the
abnormal
which latter
on previous excitation.
These
effects,
seen in electrical
in
by the very
fact of its
That
is
to
say,
an isolated
its normal
525
be gradually restored.
This leads me to what is theoretically a very interesting
mode of determining the velocity of transmission in nerve,
by the mechanical response of the nerve itself, which will be
understood from the diagram already given
that figure,
A B C
any mechanical
(fig.
315).
In
is
slip,
transmission of excitation.
The
is
when brought to a
A <^ by a pair
connection with an induction coil. The
of electrodes in
nerve,
first
excited at
induces a contractile
c,
obtained
The
in this
manner
effect of stimulus
be given later
will
in
the chapter.
and the
/,
initiation of response
The
determined.
difference
(tf)
is
b.
By
this
I
found the velocity of transmission in a certain
specimen of nerve of fern to be 50 mm. per second.
means,
It
is
It is to
ultimately
to
initiated as
As
be
small,
we can
by the nerve
is
relatively
it
this loss of
time
is
much
lessened.
COMPARATIVE ELECTRO-PHYSIOLOGY
526
The
organism,
in the intact
tion
we have
understand that
will
it
conducting tissue
the
become
is
conductivity, even
isolated.
and
fatigue,
more
still
The
easy to
is
it
fluctuating
when
inevitable changes
will
then be,
By
the absorption
in,
conducting
feebly,
The
And
highest conductivity.
has been
even, further,
transmission
will
the
velocity
its
tonic
of
condition
Over-stimulation, again,
conferred by previous stimulation.
by inducing fatigue, diminishes the power of conduction of
This fact I shall be able to demonstrate by
true excitation.
special experiments.
owing
to
is
mal-nutrition,
or
falls
for
other
reasons,
the tonic
effect.
And
here
it
may
may
in general
employed
in
527
made by
recording-point, as
it
in
is
thus unmodified.
mode
very-
is
movement of
smoked surface, that it
The difficulties
most purposes.
of record by the
would be adequate
for
lie, first,
obtaining of a
in the
in the
overcoming
is
Even
always necessary, as
this
degree of magnification
have already
said.
The
is
not
records which
The
to be overcome.
smoked paper
is
friction offered
by a writing-surface of
surface of
the construction
of
my
Oscillating
Recorder.
The
glass
plate,
means of spiral
COMPARATIVE ELECTRO-PHYSIOLOGY
528
in
a state of to-
this,
recording point
surface.
is
Thus the
its
oscillation
adjusted,
plane,
and the
right angles to
being about
barely to touch
mm.
the
it
The
smoked
periodi-
speed
of the
recording-surface
low,
the
curve of record
appears as continuous.
In
determination of time-relations
is
important, a high
speed
From
these,
(fig.
323).
529
smoked
surface.
The
latter.
following
response
is
a photographic
is
converts
stimulation
negative, in a
observed
324)
(fig.
Owing
this,
through
diphasic,
normal
Another interesting
in
Successive
into
nerve
given later
is
(fig.
404).
show
in
of the conductivity of a
nerve.
It is
But
shall
not only
customary to suppose
nerve
the
that
is
indefatigable.
Record of Mechanical
Responses to Electrical Stimulus obtained on Smoked Glass,
and given by the Optic Nerve
of Fish Ophiocephalus
P'lG. 324.
is
but
its
positive
re-
The demon-
also.
a succeeding
in
effect
of
chapter.
fatigue on
conductivity
for
experiment
to
the
Kunchangraph,
diagrammatically represented
in fig.
a length of
selected
315.
the
The
manner
length B c,
cm.
At
depression
undergone, from
M M
COMPARATIVE ELECTRO-PHYSIOLOGY
530
isolation, its
obtained were
After a series
positive.
of
stimulations,
demonstrate
the
effect
of
on
fatigue
commenced only
we can
periodic fatigue.
In order to
conductivity,
responses
see
that
accentuation of fatigue
the
many
the
series
In
The
at
the
after
first
is
exhibit
then mani-
fested
Fig. 325.
effect
effect of fatigue,
;
01
Transmitted Effect
Gecko
seen
first
as periodic fatigue
positive.
and the
positive response
by excessive
is
completely abolished,
series.
as
is
disappears.
We may
is
excellence^ will
sometimes
53
Conduction, therefore,
non-conductor.
is
not
molecular condition.
a certain
remains uninterrupted,
and cease
to
may
conduct.
Recovery
by
nerve,
nevertheless
may
whose continuity
undergo paralysis
then, in
many
in-
tetanisation.
fifty
times,
we have been
may
able to
be observed
in
its
lowest terms.
When
and important phenomena regarding the excitatory changes in nerve are revealed, and some of these will
be described in the next chapter.
still
further
MM
CHAPTER XXXVI
MULTIPLE RESPONSE OF NERVE
Great sensitiveness of the high magnification Kunchangraph
shown
Individual
con-
Sudden
enhancement of
mechanical response of nerve on cessation of tetanisation Secondary excitation
Multiple mechanical excitation of nerve by single strong stimulation
Multiple mechanical excitation of nerve by drying.
tractile
twitches
in
tetanisation of nerve
We
changes
in
more than a
million times.
as
we have
occurrence of differential
the Kunchangraphic
effects at
method of
In
is
no
We
also
saw
that,
whose
aid
hitherto
many
It
will easily
be seen that
this places at
533
may be
times,
ways.
may,
By
magnified
the
in
may
The
series.
of
either
two
different
image of the
be thrown on a distant
place,
first
may employ
the
Or, in the
of these gives
first
screen.
sensitiveness of
second place, we
five
in
this
record
accomplished
magnification, say,
in
of
lever,
times
is
easily obtained.
It is
however, not
difficult to
it
is,
all
disturbing elements.
conditions.
from
isolated nerve in
natural
its
sources
becomes increasingly
attended by an abnormal relaxa-
of energy,
This process is
which causes a steady movement of the spot of light
one direction. When the nerve has become very sub-
sub-tonic.
tion,
in
of the
spot
of
light
is
retarded
beyond
is
the
this
or
is
to
downward
arrested.
In
no further response,
this
arrest.
pronounced, stimulus
response
is
by recovery
followed
in
the opposite
downward
'
tonic
drift.
The abnormal
condition,
feeble stimulus.
it
By
and
is
direction.
The
further effective in
final
arrest
of the
positive response
is
COMPARATIVE ELECTRO-PHYSIOLOGY
534
condition
raised,
is
From
enhflnced.
this point
At
normal negative.
tractile or
showing
contraction.
successive
will, after
bring on fatigue.
may
responses
its
This stage
is
characterised, again,
a whole, and
this,
itself
The
maximum, and the
Long and intense
tonic
stimulation
nerve
increase, the
amount of
certain
amplitude, second,
of a
become,
diphasic
first,
diminished
in
and,
thirdly,
amount
We may
character,
stages
four
response of
the
in
nerve.
characteristic
more
as
its after-effect.
first
tion
is
allowed to
it
its
ever, brings
stage.
UNIFORM
transition
responsiveness, the
FATIGUE
stage, there
is
a tendency, as already
show
to relaxation
the
diphasic
or
Further
characteristics
relations
each
to
Chapter XLI.
A few words
the nerve,
when
of
other,
may be
it is
the
in
abnormal
these
will
four
be
positive
stages,
treated
in
character.
and
their
detail
in
We
in
535
of nerve
lation
even
obtained with
is
so
short-lived
a stimu-
as
The
of a second.
stimulus,
and
its
due
contractions
quick.
this response
the
to
become
It is in
conse-
fused, but
intermittent
show themselves
Owing
twitches,
to the high
up or down,
it
difficult in
is
effects.
But
it is
The
a revolving
screen
at
mirror,
some
and
distance.
This
may
after-effect
extremes
is
it
made
incident on
to a large
white
trend either in a
continuously, according
sub-tonic
from
reflected
relaxation
of
may
as
there
is
or a growing
induced a continuous
contraction,
Somewhere
between
due
to the
these
two
less
stimulus.
COMPARATIVE ELECTRO-PHYSIOLOGY
536
as a horizontal line.
movement
of the re-
Under
tetani-
is
observed as serrations.
will
it
is
the
in
stimulation
by
be remembered
tetanic
shocks.
(p.
shows on
its
cessation a sudden
transient
But
mechanical
response of nerve.
This
increase.
maximum
there
is
contraction, which
by the usual
recovery.
the latter
is
is
be seen,
in
is
may
its
it is
a factor in re-
by the
results
and
It will also
covery.
effect of negativity,
converse positivity.
be exhibited.
many
differ-
diverse
dominant.
recovery
is
for
if
its
a second
is
com-
a similar stimulation
will
once complete,
response.
is
537
be a brief period of
its initiation.
used
the
animal nerve.
for the
vegetal
secondary contraction.
of frog
shocks,
excitatory electrical
variation
found
is
known
as
is
with one end lying upon the end of the other nerve.
laid,
On
is
my own
first
by
electric
to
cause
preparation.
employed, instead
of nerve of
such repeated
responses.
in
It
appeared
that
the
COMPARATIVE ELECTRO-PHYSIOLOGY
538
nerve-and-muscle preparation.
Here
muscle.
at
it must
be
one end of the
its
excita-
The
of the frog-preparation.
power
is
much
rence of the
its
conducting
first
of these responses.
These considerations
for the
It
delay
in the occur-
when subjected
to a single
In
after a longer or
were
in
the series a
number of
There
by
is
susceptible of multiple
excitation.
If
the
nerve
in
nerve-and-muscle
is
preparation be
of spasmodic contractions.
the correctness
by
verify
the
supposition
may
regard
the nerve.
as
And
As
experiment.
responses
individual
in
539
in
amplitude,
ex-
such
of
record
series
by an ordinary magnification
on smoked glass. In order
obtain this record under
normal conditions a stream
of
air,
bubbling
through
water, was passed through
to
the
chamber
at
Owing
to
rate.
down
in
of the
the
the
latent
we
nerve,
observe
to
a uniform
run-
energy
are
able
consequent
By
growing relaxation.
the
manipulation of a stop-cock
the
air
is
through
passed
In this
way
the nerve
air instead of
This
moist vapour.
substitution
sented in
is
repre-
the record
by an
is
The
nerve,
tonicity,
traction
(fig.
contractile
of
irregular periodicity.
CHAPTER XXXVII
RESPONSE BY VARIATION OF ELECTRICAL RESISTIVITY
by
'
modification
'
Excitatory
Characteristic difficulties of
change,
its
investigation
Morographic record by variation of resistivity Inversion of curves at deathSimilarities between mechanical, electro-motive and resistivity curves
of death The true excitatory effect attended by diminution of resistance
Response of plant nerve by resistivity variation Independence of resistivity
and mechanical variations Responsive resistivity variation in frog's nerve,
and its modification under anaesthetics.
point
ance
'
modification
and
this
'
modification
'
Subsequent observers have also noticed a diminumany cases when a tissue has been
electric
shocks.
These diminutions of resistance
to
subjected
are observed as more or less permanent after-effects.
The
experiments in these cases depend on obtaining the galvano-
the tissue.
tion
of resistance in
undergone a diminution.
is
open to several objections.
The passage of constant or induction currents through the
tissue would not only give rise to polarisation effects, but
would also induce an unknown electromotive variation at
the two contacts on the surfaces of the tissue, to an extent
depending on their differential excitability. The observed
deflection by a small testing E.M.F. is thus affected, not
tissue has
This
method, however,
The
of resistance,
question
What
remains,
still
effects.
is
the
nature and
As
the
does
it
as
remote
Or
is
it
consequence
of
other
changes
excitatory
molecular
change,
induced
by
excitation,
which
of
the
independent expressions
showed
in
the
first
molecular change
identical
may
be detected
in
in
is
that one
different
Thus the
electromotive variation.
is
shown,
for
Desmodium
is
restrained,
may
similarly,
by means of the
It was, in fact,
first
latter
in
to obtain
it
should be possible
it
will
In order to demonstrate
'
Bose,
triciti stir
De
la Giniraliti des
la
matiire Inorganique
Paris, 1900.)
COMPARATIVE ELECTRO-PHYSIOLOGV
54^
In this investigation
have
first
many serious
These
to be overcome.
be
well,
however, to see
experimental
will
in
difficulties
be dealt with
in series
method to be employed.
what important respects
we employ an
it
The
the tissue.
differential
contacts.
Such
artificial
to give an electromotive
fails
by
variation
excitatory response
of
by
may
resistance.
resistivity
expect
it
response
to exhibit
The recording
variation
has
of
thus one
advantage over that of the electromotive variation, inasmuch as the record is not affected by complications due to
differential action,
of excitation.
mine, then,
is
living tissue
but
is
The question which we have next to deterwhether or not the excitatory variation of the
is
its
resistance,
such variation
if so,
is
or
is
according to the tonic condition of the tissue conIn subjecting this question to experimental investi-
cerned.
gation,
well to
is
it
negative,
resistivity
variation
by which the
true
We
When
positivity.
the
definite
there
is
the temperature,
critical
point
of
a reversal
of the
si^n
of
however,
has reached
we have
death,
induced,
This
response.
seen
that
attended by
expressed
is
change to galvanometric negativity. I have already explained in Chapter XVI. that, in mechanical and electrical
morographic curves, the abrupt point of inversion represents
the death-point.
I have also shown that this death-response
is a true physiological response
that the temperature at
which it takes place is definite in all phanerogamous plants,
being at, or very near, 60 C. in normal specimens and that
;
it
displays depression,
ture,
when the
tissue
by transposition
is
to a lower tempera-
physiologically depressed
by such
influences as fatigue.^
From
these facts
we might
response by variation of
there
expect,
resistivity, that
if
up
a tissue showed
to 60
C, or
so,
would
resistance,
In that case,
it
p.
177.
COMPARATIVE ELECTRO-PHYSIOLOGY
544
(fig.
ohms.
P and
hundreds of thousands
of
Q represented
the ratio-arms
n
y^
r\
^*
^
5'>^o -v-^
i'
by me,
R a standard megohm,
"^^s
X of
--/^h^
/"
-^--rJ
II
C\
'
^^^-U_
Fig. 327.
ment
or half-megohm
and
r, standard
the specimen
when
the bridge
The
is
balanced, S
=^
It
or
-5
megohm
whose variations of
is
now
evident that
R.
was only
'05 volt.
owing
to the
the circuit,
very low
in
order to avoid
result
strong current.
specimen
In
continuous
of temperature,
rise
it
to
and
gradual
was placed
in
the thermal
is first
for
an
specimen is subjected
have already seen that no
to
no variation of temperature.
We
of stimulus,
in
is
physiologically
induced, in consequence
isotropic
tissue.
Any
movement of
diminution.
time at every 2 C. of
Is
of temperature.
rise
of 2 C.
The method
now so
described, proved
of
delicate that
it
was impossible
to
It
thus
point
commences
of inversion.
at 56
C,
it
to increase,
till
N N
the
COMFARATIVE ELECTRO-PHYSIOLOGY
546
critical
still
point
is
reached.
And
here,
we
see a
Fig. 329.
Fig. 328.
Fig. 328.
Fig. 330.
is
is
astonishing
to find
different
curves
methods so
as the mechanical, the electro-motive, and that of
as
by
is
side
The
(figs.
excitatory effect
may
thus
ment
is
not a consequence
is
The
restrained.
is
of the
mechanical re-
when
physical move-
exhibited independently,
we
is,
as
shall see,
already familiar.
employed the
As
before.
it
same
order to determine
was important,
The thermal
able.
this
question
of stimulus,
previously given,
I
employed those
of platinum
in
as
to be so reli-
wire enclosed
contact with
the
it.
by a metro-
But as, on
sudden thermal variation acts as a
it
the
should
ments,
The
400,000 ohms.
was
in
to be
It
COMPARATIVE ELECTRO-PHYSIOLOGY
548
minutes.
The consequent
The
The normal
resistance
The
Fig. 331.
resistance of the
That of the
tissue
of Fern
Response to stimulation
resistance.
The
The
In
order, next
to
determine
of resistance
variation
in
the
present
whether the
case,
resistance
variation
form, or
to
include
carrying
certain
length
in
of
this
the
specimen.
manner
way
On
obtained
It
variation,
is
is
thus seen
that
the
by
resistivity
of the
excitatory
response,
an independent expression
variation.
by the methods of mechanical and electrowe saw that, while the normal response
was negative, this was liable to become reversed to positive,
under two different conditions namely, sub-tonicity and
table nerves,
motive variations,
Similar reversals
of resistivity variation
is
employed.
We
gradually converted
normal
into
to
response.
Parallel
resistivity
variation.
exalt
results
The
is
inter-
feeble
this
increase of resistance,
negative, through
enhanced
are observed
initial
in
the
negative
case
of
abnormal response, by
an
increase
of
resistance
or
of diminution
of
resistance.
Further tetanisation
In the
first
fig.
332,
we
COMPARATIVE ELECTRO-PHYSIOLOGY
550
no response or an occasional
either
flutter, in
the positive
direction.
We
of response, there
that
by
results
It
is
also a third
Resistivity Variation.
We
mode
available
namely,
is
in
wiAilMy uwv
t'iG.
332.
The normal
all
may
be.
three methods,
we
find that
up
by these
at or near 60
tivity,
there
is
to negative, contraction,
crease of resistance.
CHAPTER XXXVIIi
FUNCTIONS OF VEGETAL NERVE
Excitability of
stimulus
tissue
Physiological
sensibility,
of
growth, ascent
of
and
sap,
of
motile
light
to remotest
that in
of stimulus.
we
And
it is
now
high degree.
seen
The
What
power
is
the
tissue
The
place
gravity, are of
part.
And
this
is
in
bringing
little
or
no
A vena,
observed, even
when
is
across.
If, indeed, the highly conducting nervous elements had
been concerned, these curvatures in response to unilateral
COMPARATIVE ELECTRO-PHYSIOLOGY
552
we
This
will
be clear
on by
unilaterally acted
light.
Here a
positive heliotropic
movement
in the
of
peculiarity
phenomenon
by
this
concavity and
curvature towards
movement continues
perfect there
proximal and
is
responsive
This heliotropic
light.
until the
become
the
in
lies
When
itself in
this orientation
the
has
now
equally stimulated.
Had
curvature
depended, been
as
highly conducting
as
the
movement would
distal sides,
Indeed
this neutral-
seen even
For
under
when
unilateral stimulus
circumstances
these
is
excessively strong.
stimulus
conducted
is
the
result
And
it
is
place, even
shown elsewhere
that, in
under feeble
stimulation.
I
have also
movements of
the
responsive
leaves, conduction
little or no part.
acted on by light without showing any responsive movement.
Hence the lamina is not to be regarded as the perceptive
plays
organ.
The organ by
movements of the
pulvinoid.
This
is
at
is
is
the pulvinus or
553
induced.
is
When
movement.
The
lamina,
it is
true, is
provided with a
fine fibro-vascular
and serve
ramification,
it
to stimulate the
plant as a whole.
When
petiole,
from
true,
it is
excitable on
its
two
movement of the
dorsiventral character,
its
But
sides.
is
a great
loss,
most
and
since,
no directive
Nor could
between
parison
any
there, in
the
The
unequally
elements there
leaf.
is
case,
effective
strength
an
external
nervous elements
by no means the
is
essential
factor in
subserve so
The
many important
functions in the
question, therefore, as to
what
is
life
which
of the plant.
still
economy
remains
to be answered.
One
Physiology
activities
is
that of Correlation.
may be
set
up
less
some excitatory
influence.
In
this
way every
COMPARATIVE ELECTRO-PHYSTOLOGY
554
now proceed
shall
stimulus,
and
its
the
essential
serving
factor
the
of
activities
sub-
in
various
the
life-
organism.
important,
is
main-
in
seen
easily
is
the
in
case of
Under
these
conditions,
motile excitability
to
And
disappear.
its
found
is
this
is
We
light.
again, in
the
have
seen
course of the
Fig.
333.
Effect of
Tetanisation
in
is,
of
normal
favourable
becomes sub-tonic
or moribund, and then its
ordinary responsive power is
conditions,
Photographic Record of
En-
abolished
Under
or
even
these
reversed.
circumstances
Abnormal
into
positive response
normal negative.
stimulation,
in fig. 333, to
when
is
become enhanced.
It is
is
excitability,
found,
as
less
its
normal
continuous supply of
555
is essential to
condition of a tissue.
It
known
is
tivity of a
that
nerve
is
in
animal,
the
when
the
conduc-
abolished
It is therefore
kept at
is
its
normal
level
shall
next
briefly
to
refer
all
fact
which
have
autonomous movement,
and growth are fundamentally excitatory
phenomena. Thus, for example, the autonomous rhythmic
movements of the lateral leaflets of Desmodium gyrans come
to a standstill when their store of latent energy is exhausted.
And it is only by the accession of fresh stimulus from
logical activities of the plant, such as
ascent of
sap,
when
sub-tonic, the
its
We
internal energy
is
normal tonic
condition, or health of the plant, being dependent on the
sum total of stimulation previously absorbed by it. Turning
next to the question of the ascent of sap, I have shown that
the most important factor in bringing this about is the
multiple rhythmic activity of certain interior tissues of the
plant.
is
fall
On now
it
owing
it
we
find
the excitability to
(p.
383).
COMPARATIVE ELECTRO-PHYSIOLOGY
556
But
is
if
stimulus be applied,
is
The importance
is
quantities
which, as
is
is
shown
may enhance
in the following
the ex-
photographic
The
three responses
to
The specimen
and
under the same stimulus as before. Light was now cut off,
and after an interval response once more taken. This is seen
to have been of the same amplitude as at the beginning
(fig.
334).
Thus
for the
factor
we have
just
namely
finally
enhance excitability.
been considering we have,
light is seen to
the
photo-tonic
among
the
many which
And we
this
source of stimulus,
activity,
and
its
its
motile
response of growth,
is
all
Now
it
is
interior tissues,
whose
its
suctional
disappear.
When
activity,
activity
is
many
of those
life.
How,
then,
is
557
This
fact,
more
still
sensitiveness oi
one branch of
rest of
it is
Mimosa when
this plant
exposed to
Fig. 334.
is
again
If
light, it is
First series,
first
undergo no
loss
of motile
sensitiveness.
It
is
thus
COMPARATIVE ELECTRO-PHYSIOLOGY
558
The
transmission of stimulus
One
the
is
life
rendered possible.
leaf,
now made
is
Among
external
perhaps
is
so
clear to us-
none
stimuli,
essential,
or
so
green
to
energy
as
And we now
light.
the
plants,
ramification
fine
see
ot
that
of fibro-
an area as possible
in the leaf,
provides
catchment-
virtual
Distribution of^Fibro335.
vascular
Elements in Single
Layer of Stem of Fapaya
Fig.
There are
at
least
lus.
thus
not
is
merely a specialised
structure,
for
purpose
the
of
20 such layers
sitive
area
for
the
absorption
of
transmission
And
even
downwards
in the interior
into the
body of the
plant.
these
is
cited
ments buried
in
them.
How
is
reticulated tlj^y
may
be,
ele-
even
Papaya
(fig.
335).
portion
is
out
as
its
many
thus seen
how
all
559
by means of
It
is,
upon any
other.
is
affected
CHAPTER XXXIX
ELECTROTONUS
Extra-polar
effects
of
variation
polarisation
of
electrotonic
currents
excitability Bernstein's
increment Investigation
on
vegetal
polarisation
nerve Electrotonic
decrement Hermann's
Conductivity en-
hanced when excitation travels from places of lower to higher electric potential,
and depressed in opposite direction When feeble, anode enhances and kathode
depresses excitability All
action of these factors
When
an
electrotonic
phenomena
reducible
to
combined
electrical
current
is
nerve entering,
say, at A,
e:
Kat-t
<
a
An
Fig. 336.
Fig. 337.
Fig.
Fig.
cally positive.
regions.
is
On
On
the
kathodic
found to be lowered
side, the
near
electric potential
away.
is
found to be raised.
feLlECTROTONUS
56
vegetal
nerve,
Taking a
fresh
have
exactly
obtained
In the
similar
results.
2*5
elec-
electrotonic
The
value of the
10-
i-0
=-
1-2
==-1-4
Fig. 338.
potentiometer arrangement.
E.M.F.
k, kat-electrotonic to right.
and kat-electrotonic
effects
in
The
circuit, gives
% 338
seen such a record of effects both an-electroand kat-electrotonic, taken while the acting E.M.F. was
increased from -6 to 14 volt, by steps of -2 volt at a time.
I"
is
tonic
'
Certain
considerations, which
this
I shall
assumption.
O O
COMPARATIVE ELECTRO-PHYSIOLOGY
562
An-electrotonic effect
Kat-electrotonic
50 divisions
65
107
126
150
i-o
1-2
1-4
In this particular
58
experiment,
it
will
eflfect
divisioiis
no
124
148
.-..
.,
an-electrotonic
equal.
slightly
constant electrical
current
We
next turn
is
its
the
to
question
of
the
variation
of
by the passage of a conOn this subject the most important constant current.
tributions have been made by Bernstein and Hermann.
Bernstein, experimenting on the sciatic nerve of frog, found
excitability induced in a tissue,
that
excitation
experiment
is
induced
polarisation
339, 340.
diagram
(figs.
339,
Fig. 340.
Fig. 339.
F'igs.
This
decrement.
Diagrams
Decrement
Fig. 339 shows decrement of kat-electrotonic, and fig. 340 of an-electroIn this and following figures
tonic currents, under stimulation at s.
the inside thin arrow indicates direction of polarising current, the
outside thick arrow the direction of responsive current.
340).
In
fig.
339
the
kathodal
by
When
effect
seen
is
the nerve
is
now
induced
excited
current
is
induced.
When
is
induced
in
ELECTROTONUS
the extra-polar circuit, by reversal
563
of polarising
current,
kat-electrotonic,
of the nerve
(fig.
undergoes
340).
It
diminution on excitation
Fig. 341.
^Figs. 341,
342.
Fig. 342.
secondary
coil
of an inductiorium.
to
left,
augmentation.
left
is
Hermann
to right or from
undergo an
to
to
alterations
is
polarised.
It
its
is,
'
Increment' of excitation).'
It
vol.
ii.
p. 315.
002
COMPARATIVE ELECTRO-PHYSIOLOGY
564
a
to
shall,
however, be able
to the combination of
and on
to
These separate
excitability.
circumstances,
Hence the
either
conspire
may, according
effects
or
act
antagonistically.
at
first
by which an electric
current induces a variation of conductivity and excitability,
we must first determine the pure effect of the current on
conductivity, apart from any excitatory variation
and,
secondly, its effect on excitability, uncomplicated by any
In order, then, to discover the laws
variation of conductivity.
To
take conductivity
first
to the region
or
an-
moreover, a
circuit,
by
side.
It
is,
influence
differential
is
upon
action,
In
it.
unless
led-off
proper pre-
therefore, desirable to
cell
took a petiole
its
ends through a
(E.M.F.
volt).
The
of the same
leaflet,
great
to
(figs.
343, 344).
thus seen that the led-off electrodes are at a relatively
it
is
is
ELECTROTONUS
reduced
to
The
nil.
565
excitation
is
to
is
negative
of
namely,
direction,
from
electrical
excitation,
current have an
this
fact
an
in
galvanometrically
the
Now,
galvanometrically positive.
an
of
conduction
manner,
uphill
negative to the
direction
against that
or
point,
electrically
will
effect
the
if
on the
by
be detected
The
determine
experiment
<
ill
^-
'^
Fig. 344.
"Fig. 343.
Figs. 343, 344.
be found to
will
this question.
shown
now
in fig.
'
potential
that
of the kathode.
is
manner, or
It will
retarded or abolished
trically
so,
we
anode to that
when
excitation
is
made
be
expect to detect the fact by the diminution of
If this
its
actual
is
to travel elec-
to a feeble or
is
re-
makes
moderate
its
appearance alone.
COMPARATIVE ELECTRO-PHYSIOLOGY
566
In
fig.
will
it
will
We
actual reversal.
conductivity,
of
direction
that
region.
For
this
passage
of
excitation
ends
effect
in
on
an uphill
in
to say, from
Fig. 345.
when
the
is
it
Excitation
Current
a,
Normal response
response.
direction.
that
polarising
current
is
in
same
now made
the kathode.
direction as normal
current in opposite
Before doing
this,
however,
of
the
previously
acting
left-to-right
slightly
current,
enhanced above
first
ELECTROTONUS
567
This
ing current.
is
shown
In a
sponses reduced
is
to say,
we have
the
current.
In
<^
normal response
we have the re-
which excitation
in fig. 346, in
that
'5
volt in
c,
"i
volt for
On
polarisation.
Fig. 346.
Photographic Record of Modification of Conduction during
Passage of Excitation from Anodic to Kathodic Region, under Increasing
Intensity of Polarising E.M.P\
a.
Normal response
I
volt
under
"5
<:,
now
being
by negative.
on the application of
Finally,
volt in
</,
we
specimen,
was made
when the
region.
It will
excitation
to
of a polarising E.M.F. of
"i
shown
in b, as
compared
COMPARATIVE ELECTRO-PHYSIOLOGY
568
in
i,
"5,
a.
and
The
1*5
application of higher
volts
now
respectively,
and
d,
e.
was unable
volts because
1*5
c,
the
unsteady.
these
be borne
It is to
potential gradient
experimenting on electrotonic
that the E.M.F.
employed
is,
effects,
it
cm.
Normal responses
b,
c,
d,
e,
In
must be remembered
Fig. 347.
in
from
From
we
A
tivity.
The conductivity
is
unmixed
effect
seen that
circuit are
there
is
effect of a constant
points in the
same
no resultant
response,
since
the two
excitatory
ELECTROTONUS
each
balance
effects
The
other.
569
galvatiometric
effect
is
But if one of the two have its excitability enhanced in any way, this balance will be disturbed, and a
resultant current will flow through the circuit, the more excitable contact becoming galvanometrically negative.
I now
took a long piece of isolated vegetal nerve and connected
the galvanometer at E and E\, a secondary coil,
it with
then zero.
The
longitudinal
con-
circuit
two
tacts
(fig.
and E
e'
more
or
no
to
in
the
being
was
at
resultant
sponse
being also
equally
less
excitable, there
first
shocks,
re-
stimulation.
meter
in
the direction
of e'GE shown
thin
inner
by the
arrow,
E'
FiG. 349.
Experimental Arrangement
Figs. 348, 349.
to Exhibit the Enhancement of Excitability at Anode, and its Depression at
being galvanometrically
On now
positive.
plying
ap-
equi-alternating
sponsive
ege', as
and
the
re-
current to
be
is
feeble
Fig, 348 shows enhancement of excitability
at anode ; Fig. 349, the depression ot
excitability at kathode.
Inside thin arrow indicates direction of polarising current. Outside thick arrow, direction
Note that in both
of excitatory current.
there is a so-called polarisation -decrement.
in the
E'
now undergoing an
citability
of
e'
variation.
has
in the direction of
become
ege'
(fig.
349).
e'
is
now
the kathode, E
COMPARATIVE ELECTRO-PHYSIOLOGY
570
in relation to
current was
current, in
kathode
it
in
being anode.
the opposite
On
direction
to
the permanent
E' or the
E.
Two
In
fig.
different
Photographic Records of Response, illustrating the EnhanceExcitability at Anode, and Depression at Kathode, under
Feeble Acting E.M.F. in two Specimens of Nerve of Fern a and b
Fig. 350.
ment of
the enhanced
ELECTROTONUS
But we
decrement.
termined as this
An
electrodes.
I
shall
is
57
is
by the
identical variation
two
of excitability may, as
a polarisation-increment or as a decrement.
I
have obtained
It
But,
if
much
this
be prolonged, there
is
From
variations of excitability
A feeble
tissue
currents,
we
This result
is
startling, contravening, as
it
does, Pfliiger's
currents
would be shown by
COMPARATIVE ELECTRO-PHYSIOLOGY
572
'
r<r<~rt^
it)
Fig. 352.
Fig. 351.
employed
case to the
Here we
left.
anode being
in
the
first
by the proximity
at e'
proximity
Of
of
kathode.
citation,
moreover, which
S,
uphill,
Photographic Record of Response
under the Arrangements given in Fig>. 351,
352 in Nerve of Fern
Fi^"-
a,
353'
moving
anode,
towards the
or
and
owing
to
conductivity
increased
in
electrically
that
direction
the
current.
I),
r,
Resultant response downwards when polarising current is from left to right, as shown
by arrow ->. This shows excitability of
e' and conductivity in direction Se' to be
relatively enhanced.
Resultant response upwards when polarising
current is from right to left <-.
This
shows excitability of E and conductivity in
direction SE to be relatively enhanced.
E'
is
excitation
is
ducted than
of
the
which
is
itself
in the case
second
is
wave,
proceeding
better con-
E,
but also
ELECTROTONDS
573
kathode.
first
of
pair
down
responses in
is
and we
When,
figure 353^.
reversed
now
(fig.
352), the
fig.
353
<:.
To
go back to the question of the relative directions of electrotonic and responsive currents, we find in that case, when
the anode is to the left, that E' is galvanometrically positive,
while the excitatory change makes it galvanometrically
negative.
This means that the excitatory response takes
place by the. so-called polarisation decrement.
When the
anode again is to the right, the galvanometrically positive E
tends by excitation to
This
will
become galvanometrically
figs.
351, 352.
negative.
which
The inner and thin
the
arrows
decrement.
In
example of the
so-called polarisation-
the
same
shall describe
another experiment.
The experimental
tissue
is
is
galvanometer
is
it
will
be noticed
(figs.
354,
The
left
electrode
COMPARATIVE ELECTRO-PHYSIOLOGY
574
it,
The
the thin inner represents the polarisation and the thick outer
the responsive current.
On now
take
excitable,
place
by the
a polarisation increment
Fig. 355.
Fig. 354.
Figs.
sets
fern
and frog
respectively.
Balance was
first
obtained at
When
made anode,
the
enhanced excitability of the anodic point. When the righthand end, however, was made kathode, the balance was
upset in the opposite direction, that is to say, down, showing
that the left-hand anodic point was now the more excitable.
Fig. 356 gives a record of these responses as obtained from
The
the nerve of fern, and fig. 357 from the nerve of frog.
responsive currents in these cases, it should be noted, are in
the same direction as the polarising current.
ELECTROTONUS
On
^71>
on polarisation increment
first
case, the
galvanometer.
position
This
of a shunt in the
was placed
in
we
polarisation-circuit, while
in
the
second
it
series.
From
now
shall
find ourselves in
Fig. 357.
Fig. 3s6.
Fig. 356.
Fig. 357.
experiments of
Hermann and
these, although
consistent.
referring
is
Bernstein,
First, then, to
back to
figs.
we
find this to
be an instance
in
COMPARATIVE I^LECTRO-PHYSIOLOGY
576
causes an increase of
it,
we
we have
here
to
deal
excitability at the
same
(fig-
339)
i"
We
time.
circuit
is
under kat-electrotonus.
E' is therefore relatively anodal,
and consequently more excitable. The excitation from the
stimulator S which reaches E' is in this case impeded in
reaching E by the fact that
that
it
has to travel
electrically
kathodal
E.
Thus,
which reaches E', and owing
also to the greater excitability induced in it by the fact that
downhill
owing
it
is
is,
E' to
The
thick arrow in
direction
indicated
the
to
by the
polarisation
thin arrow.
polarisation current
therefore relatively
(fig.
340),
opposite
is
which
reversed
is
current,
is
latter
is
when the
anodal, and
E' kathodal,
and there-
ability of E
stimulation, galvanometrically
by a thick
shown,
same
in
It
has
ELECTROTONUS
As
regards conductivity,
it
577
consequence of
this
fact
that
is
to say,
electric potential.
In
With reference, again, to electrotonic variations of excitawe have seen that under feeble E.M.F. it is the anode
bility,
applicability of
This conclusion
known
which
will
as Pfliiger's
be discussed
P P
CHAPTER XL
INADEQUACY OF PFLUGER'S LAW
Law
Reversal of PflUger's
E.M.F.
Investigation
practice.
we found
that,
it
in
nerves, in
excitability
Now
and
this conclusion,
is
Biophytum,
in
Thus,
a given experiment
in the case
it
was found
of the sensitive
that, using the
anode
at the
which the
make.
itself at
anode.
579
interrupted.
;
of the current.
excitatory reaction
tiated at the anode,
at the kathode.
These
are
normal
the
under Pflliger's
Law, which holds good within
effects, falling
medium range of
But when the E.M.F.
a certain
E.xM.F.
is
much
higher,
find
that
become
Thus, employing
reversed.
an E.M.F. of 220 volts, it was
these normal effects
found
that
excitation
took
Fig. 358.
Upper
figure
excitation
kathode.
effect
shows
make,
produced at
Lower figure shows
effect at
being
at break,
now produced
excitation being
at anode.
initiated
at
At
COMPARATIVE ELECTRO-PHYSIOLOGY
S8o
at the
that an excessively
fact
may
effects,
reversal of these
when
effects
the
intensity of E.M.F.
when
to say,
is
it
is
is
the excitatory
when administered
one
and,
in
of a stimulating agent.
effect
effect,
in
chemical
moderate or very
will, if
These
of exaltation.
reversals,
under varying
intensities of
Thus
chemical phenomena.
formation
of
the
known
that
in the
well
it is
'
normal
negative,' a stronger
and a
positive,'
We may
still
more intense
of recurrent reversals.
Returning,
excitability,
we
then,
to
the question
of
polar action
on
In
this,
which
is
sometimes
muscle
is
the kathode
is
is
no
inter-
stage
excitation.
in his first
58
either.
make
ments.
derives
For, using a
particular effect of
The
is
is
differences of opinion.
specimen.
It
therefore occurred to
me
that
the effect of
a feeble anode in enhancing excitability might be demonstrated conclusively in the case of the vigorous intact animal.
With
experiments on certain of
carried out a
number of
my
on
If
may
non-polarisable electrode
COMPARATIVE ELECTRO-PHYSIOLOGY
582
intensely painful.
during
its
This
is
make and
On
restored.
Fig. 359.
Effect of
is
anode, with
in Human Hand
By means of reversing key, R, E in connection with the wound-spot may be
made anode, and e' kathode, and vice versa. By alternately pressing
the keys, k' and K, feeble or moderately strong E.M.F. may be em-
ployed.
the
same E.M.F.
The
as before.
sensation
now experienced
On
was again
restored.
In these experiments
we have
excitability,
it,
of
583
the
current,
verification,
undertook investigations
on a dozen different
anode and kathode,
as the E.M.F. was gradually increased from '3 to 2 volts.
It should be mentioned here
that the subjects of the
were
experiments
totally ignorant of the object of the
investigations, and were simply asked to describe their
therefore
sensations
at
different
some
points.
As
eighteen to twenty-five.
it
may
The
following case
may be
from
may undergo
taken as typical
volt
i-o
1-5
Effect on wound-spot
when kathode
Effect on wound-spot
Marked
Marked
Slightly soothing
Slightly soothing
Increase of irritation
increase of irritation
increase of irritation
Indifferent
Slightly soothing
Increase of irritation
Painful
2 -o volts
when anode
Soothing
E.M.F. of
-3
volt.
Here, with
-3
volt,
was the
excitability w?.s
COMPARATIVE ELECTRO-PHYSIOLOGY
584
The
critical
to be slightly below
effect
observed at
kathode, while
of the anode,
in that
The
it
in all
cases.
kathode varied
between -6 and i
varied from i to 1*5 volt.
in different cases
it
volt
In a
under varying
intensities
when
in more
of E.M.F.,
other forms ot
detail.
critical
It
may
point of
different individuals,
and
The main
may
be
still
contrasts.
better demonstrated
is
full
Having, then,
-5
volt
and another of
volts
of 2
volts
'5
volt
was
applied,
E.M.F. of
'5
now gave
rise to intense
585
The lower
2 volts respectively.
On
excitatory pain.
voltage
the cessation
was
restored,
on now applying 2
and
salt,
slight
this
volts,
The
to induce
application of
"5
volt
An
depression of excitability.
E.M.F. of 2
volts,
on the
sequent pain.
These
shown
results are
Method of
E.M.F.
'5
volt
Intense pain
From
E.M.F.
Wound-spot kathode
E.M.F. of
2 volts
Soothing
-5
volt
Soothing
it
will
E.M.F. of 2
volt
Painful
low,
is
it
is
and
facts, is
in
that
of low
E.M.F.
the
last chapter,
is
is
moderate or low.
The
COMPARATIVE ELECTRO-PHYSIOLOGY
586
medical application of
electricity, since
feeble
which
is
to be expected under a
of that intended.
It
Variation of Excitability
fails
when
effects
the E.M.F.
is
is
Law
Pfliiger's
either
E.M.F. then,
it is
Under a low
is
It
fact,
of the Polar
fact that
with low
E.M
This important
F. conductivity
and depressed
in
phenomena
of nerves
CHAPTER XLI
THE MOLECULAR THEORY OF EXCITATION AND
ITS
TRANSMISSION
Such opposite
Arbitrary nature
of term
excitatory Pro-excitatory and anti-excitatory agents Molecular
distortion under magnetisation in magnetic substances Different forms of
sponse under magnetic stimulation Mechanical, magneto-metric, and electromotive responses Uniform magnetic responses Response exhibiting periodic
groupings Ineffective stimulus made effective by repetition Response by
Two
re-
'
re-
resistivity- variation
electric radiation
Molecular
Effect
model
Response
ductivity
of inorganic substance to
Conduction of
magnetic excitation
The
excitability
It
is
is
brought
As we do
power of molecular
vision
we have
is
perforce to be contented
complicated as
they are
apparently mysterious
sponse and
its
by the
properties
concomitance of other
of living tissues.
variations, however,
be
in
If re-
others depressed.
It
determine what
the nature
is
COMPARATIVE ELECTRO-PHYSIOLOGY
588
may
be due
what
it
made
to
a particular stimulus or on
to depress, the
excitability of the
same
tissue.
It is
the
phenomena
upon the postulation of a hyper-
physical 'vital
however,
mena
to be
force.'
In the course of the present work,
has been shown that not only the simple pheno-
it
of response, but
met with
in
their
all
complex variations
also, are
must be sought
As
it
The
sensitiveness.
vinus was
made
the
greatest
When
This
local expansion.
On
its
is
we observe a movement of
The pulvinus was next
kathode-make, and we observe a responsive
At kathode-break, however, we have a recovery
to
contraction.
by expansion.
We
sponsive
effects,
stimulating agent
and contraction
make
re-
of anode,
at that of kathode.
or to
conditions of alignment,
it is
589
correspond to
The
Now
this.
two responsive
be
somewhat
it is
effects
arbitrary.
been
thus
normal
taken
the
as
and
excitatory
Having once
adopted such a nomenclanegative.
ture,
of course im-
is
it
portant that
strictly
if
it
adhered
should be
Thus,
to.
mal
response,
any-
then
tends
which
enhance it must be
thing
to
vexity.
re-
traction
Aw =
convexity induced at
Kb responsive concavity at anode-break.
K/ = induced
concavity at
kathode-make. Yib
break.
anode-make.
retards
as
it
This word
depressing.
depressing
'
is,
by it might be indicated a
permanent depreciation of the
the normal
Moderate
response
rise
is
tissue,
temperature,
of
expansive tendency,
while diminution of
such depreciation.
possible without
lessen
will
for
the
example,
contractile
know
more
kathodic
the
intensity
initiation of contractile
employed be
Pfliiger's
Revert-
stimulation,
is
response.^
its
response
we
necessary, for
Should the
(p. 187).
mode of
the visible
'
to
with
Law
is
applicable.
COMPARATIVE ELECTRO-PHYSIOLOGY
590
same
direction as that
which precipitates the excitatory response, hence kat-electrotonus should prove to be excitatory.
But a moderate
anode, with
its
to be regarded as normal.
If,
instead of contraction,
we had
We
all
agencies,
must be regarded
as depressory,
or better anti-excitatory.
From what
be understood that it is
the direction of the molecular derangement which determines
the character of the response. That molecular upset, which
expresses
itself
it
will
as excitatory contraction,
we may
call
the
value
much when
distortion
was proceeding
at
a condition of permanent
It
for
is
reason
this
that,
most pronounced
either kathode-made or anode-break,* and not so much
59
is
To
recapitulate
'
some of the
excitatory
distortion
or
'
principal facts
enumerated
K-effect,
then
anything
which
K-efifect,
known
as K-tonus.
induces an
incipient
distortion
in
the
and
will, therefore,
In
the examples
given,
the
opposite
K-
or
A-effects
And
properties.
molecules
In
in
close
chains,
magnetising forces
directive
manner.
distorted
in
reverse.
It is
of distortion
of stable.
excitation induced
would be more or
COMPARATIVE ELECTRO-PHYSIOLOGY
592
usually a
is
its
original position.
we have a means of
Here, then,
We
force.
external
it
increased
is
or diminished in a
effects
modifying
various
agents on
the response, as
all
we have
But besides
we have
in
also,
the
living
tissue,
mission
tions
to a distance
that
and
is
to
this trans-
is
which
may
be induced
in the
conducting nerve.
Simi-
be able to show
larly
shall
magnetic disturbance
induced
in the
Thus,
is
in
possible,
is
conducting wire.
we
are
its
magnetic substances,
methods,
the
modifications of these
will
be recorded.
From
responses
so compre-
record,
It
is
will
then
be realised
not alone of
its
own
593
we must
find
it
Though
this variation is
very
One end
device.
indiarubber.
is
fixed,
coil,
small,
much
easier,
COMPARATIVE ELECTRO-PHYSIOLOGY
594
electro-motive method.
the magnetising
coil, c,
iUUUUL
Experimental Arrangement
Magnetometric Method of Record
Fig. 361.
for
M, magnetometer
B,
c,
magnetising
Fig. 362.
a, ammeter ; r,
;
K, key actuated by metro-
balancing coil
rheostat
Photographic
Record of Uniform
Magnetic
Responses
coil
of Iron
nome.
to a freely
mirror
(fig.
second balancing
coil, B,
attached
its
is
placed on
by the magnetising
coil.
is
inside C,
experi-
action induced
is
The
then introduced
netometer,
drum.
The
M, thrown
intensity
mag-
on a revolving
of the
exciting
current,
Fig. 363.
Photographic
Record of Periodic
Groupings in Magnetic Responses
is
the rheostat,
R.
The
duration of appli-
In
fig.
deter-
experiments.
in this
is
in successive
595
living
in
effect of
Under
molecular
This
is
strain, the
recovery did
Such
persistent strain
become complete.
not
muscle.
persistent
to
in strongly
excited
may be removed by
mole-
being
the
Magnetic
result
become
ineffective,
by
of
tap.
stimuli, individually
repetition.
In
effective
fig.
365
is
Photographic Record or
Fig. 364.
Response and Recovery of Steel
under Moderate and Strong Magnetic Stimulus
Photographic Record
showing Ineffective Stimulus
made Effective by Repetition
Fig. 365.
recovery.
seen a record of
this.
induced
it
became
effective.
be seen
Under
certain
same
mag-
conditions,
we
moreover, to be fully
enhances the
subsequent responses, while under other conditions, by inducing fatigue, it brings about their depression.
described
later, tetanisation,
as
shall see,
fact
that in addition
QQ2
to
COMPARATIVE ELECTRO-PHYSIOLOGY
596
the
now
have
studying
means
also a third
is
how
explained
extent of
the
a magnetic substance
distortion induced in
molecular
by an external
or measured by magnetometric or
For the detection of similar changes,
however, in matter which is not pronouncedly magnetic, it
is necessary to devise a method of record of more universal
application.
Such a method we have, as already said, in
the record by resistivity-variation.
It is here desirable,
however, to give a more detailed account of this and the
can be gauged
force
electric indications.
principle involved.
Our
induced by stimulus,
known
us
let
consider certain
briefly
stimulating agencies.
of
sign
disulphide,
any
change.
is
subjected for
light, there is
solubility
Its
well-
by various
in
however, has
no
carbon
dis-
means of
tion
is
this
'
developing
only possible
'
solution.
when
molecular or
the
somewhat
to say,
when
The development
allotropic
stable
is
that
persistent.
impermanent
of
any
after-effect
substance in the
stability.
This case
is
analogous to that of a
force
is
the acting
induced
When
magnetisation-
is
fugitive,
that
and there
is
is
self-
to say, the
no
after-
effect
and we observe an
persistent,
To
permanent magnetisation.
of sulphur,
is
it is
name
of
that this
more or
less
induced
allotropic change.'
is
is
after-effect in the
we single
under intense
But,
force.
597
As
it
a matter of fact
we
see
made
we had no means
current
the
of recording
by the
fact that
intermediate changes
The next
records
question
is
as to the possibility of
making such
taking place.
This
may
be accomplished, as
shall show,
two
and diamond.
Let
force
it
We may
to a small extent
latter the
extent
force of restitution
original
neutral
will
condition.
it
If
the
distortion
limit, then,
does
its
not
on the cessation of
and
distortion,
known
all.
will
be very
much
Such more or
less
protracted,
permanent
COMPARATIVE ELECTRO-PHYSIOLOGY
593
substance
panying model
(fig.
366),
if
we
The non-conducting
aspect
represented by
is
particular
ABC.
With the
substance
sensitive
posed
C.
in
condition,
this
inter-
meter would be
zero.
is
the
semi-conducting aspect
of
the
substance,
which we
may
under
imagine the
corresponding deflection of
the galvanometer to be 50.
B
is
aspect,
the
corresponding
The model
Molecular Model
representing
put
in
are
aspect
electric
the sextants
AA
to
The
is
opposite sextants C C
represent
the
non-conducting
aspect
The
communication.
tinfoil.
sensitive substances,
599
The
stimulus.
aspect
excessive,
stimulus,
its
original
galvanometer deflection
will fall
stimulus will
deflection
strain
the
cessation
of equilibrium, and
from
i(X>
of,
to the original
class,
the
distortion will be in
of
position
If the substance
value 50.
If the
the
to
The
the electrodes.
be
to
resistance.
The
coil
the
sensitive
distortions
on one
The
molecular
invisible
are
thus
revealed
galvanometric indicator
by the
the
effect
stimulus,
may
flection as ordinate,
abscissa.
It is
thus
subsequent recovery.
I
shall
resistivity
I
now
describe
variation,
we
how
in practice,
obtain
by
this
method of
The
COMPARATIVE ELECTRO-PHYSIOLOGY
600
sensitive
of radiation
is
photographic plate.
Or
may
substance
the sensitive
be
Taking
first
selenium as the
its
On
light.
A
page
series of
such responses
3, fig. 3.
The
parallel
ticles of
Method
of Resistivity
of the
Variation
Sensitive metallic particles placed in
"
gitnreZc'rLrr''""
this
,
o^t
be understood from
On
^S- S^?.
method,
m
.
obtaining records
the
find, as
,
first
pointed
chapter, that
of twO
'',
The
dincrent types.
of which aluminium
may
In living tissues
highly excitable
positive.
to,
we
cases,
stimulus
is
liable to
is
it
feeble
or
6oi
Certain
strong.
again,
now
shall
give several
under
conditions.
various
The
first
example
The
two responses exhibit incomplete recovery, which becomes comNote that warmth, increasing force
and also diminishes amplitude of
response, as seen in the two succeeding records.
first
given, that of
aluminium powder,
will
be of the negative
by diminution of
resistance.
When
recovery
the contraction-remainder
motive negativity
in muscle, or
its
heat, at
This
(fig.
is
favourable to recovery
the end
of the
second
recovery
is
persistent electro-
is
is
an immediate completion of
refer
368).
the place
to
COMPARATIVE ELECTRO-PHYSIOLOGY
602
anomaly,
where a
slight rise
the
P'lG.
warmed
substance
The next
is
seen to
now
at the
of
still
further diminished
height and
substance
in
is
same time
The temperature
When
its
succes-
com-
370
fig.
is
by a
tissues
liable
is
This
to
is
hastened
because
under very
reversal
in
case
this
adjust
critical
value
feeble
better
is
is
living
stimulation.
It
in tungsten,
tap.
It
in
same thing
603
is
easy to
is
it
below the
fall
excitation.
for
serve
a moderate degree
possesses
Using a mass
of sensibility.
of tungsten particles,
under
found
strong
intensity
of electric radiation
brought
that
substance
the
response
Fig. 370.
Photographic Record
of Response of Tungsten
was
The incomplete
varia-
or diminution of resist-
recovery
is
hastened
fig.
364.
be given
later.
now proceed
excitation.
transmission
to
We
of
Resistivity
Methods of
of
its
varia-
record,
the
state
we
of
in
living
tissues
lies
in
the
COMPARATIVE ELECTRO-PHYSIOLOGY
604
transmitted
would be reduced
stimulus
zero
to
transmitted
is
to
that
(2)
short
while a
moderate
a stronger
distance,
by
others.
I
shall
now proceed
to
show
is
Fig. 371.
changes, both, as
shall show,
observation.
which
is
receiving
The conducting
wound
the
coil,
When
galvanometer.
is
it
which
turn
causes
sound
in
the
of simplicity,
we
the
reaches R,
in
coil,
telephone, or a
605
The
coil
coil
to the
mode
electrical
of
stimulation.
Electrical
most
is
excitation
when it
The same
effective
excitation
transmitted to
effectively
is
that
in the case
it is
is
great distance,
relatively ineffective.
at the instant of
excitation
general no
at
is
longitudinal,
true
of
is
effect.
kathode-make,
is
at
anode-break.
response
metric
excitation,
excitation
seen
is
at
also
the
its
moment
of
continuance.
obtained, at break
K-magnetic
The same
of A, or south polar
magnetisation.
We
but
also
some
obtain
to
is
insight
into
the
sequence of
movement should be
opposite
with
the
it is
We
also
know
the
further
is
most
latent period,
effective
in
The
COMPARATIVE ELECTRO-PHYSIOLOGY
6o6
we
In
any
case,
molecular
it
is
second period
maximum.
at its
is
distortion
that
we should expect
to
the
see
K-make
of
afterwards,
not
the
distorted position.
movement of recovery
a sudden
is
direction.
If
now
its
in
the opposite
particular-directioned
and it
an afterThe anode-make, on
as one of depression,
effect
of depression at kathode-break.
one would
expect,
its
is
depressory.
But
at
the
break,
is,
as
the
same
by K-make, must be
excitatory.
may
be polar
in their
character.
The
fact that
certain distance to
nil.
Keeping
this
efficient
now have
to evoke
to be
moved
shall
delicate
as
effect.
conduction and
phenomenon of
its
607
fully
described
in
effects at E'
high
phenomena
degree
in
of
Fig. 372.
s,
delicacy in
the
study of
may be
similar
obtained by
magnetising
is
coil,
T, tonic coil,
the
(fig.
in
stimulator,
which
S,
shall
now
describe.
The magnetic
wound in
opposite directions.
When
a magnetising current
is
suddenly
COMPARATIVE ELECTRO-PHYSIOLOGY
6o8
one to the
towards
towards
right,
E,
to the
left,
e'.
E' at
efifects at
move
the balance
till
This
found.
is
S,
fig.
E was placed
373.
seen as up-responses.
is
The
coil
The
coil is
now moved
FiG, 373.
still
is
enhancement or depression of conductivity. Resulting upresponses will indicate enhancement, and down-responses
A well-known agent for the enhancement or
depression.
depression of the conductivity of the nerve
is
the
polar
also
be found applicable
excitatory or kathodic
in
effect
will
be facilitated
in
An
trans-
if
the molecules in
its
disposed to respond
Hence the kathodic effect
excitation.
whereas
it
now study
is
more
easily trans-
We
shall
polar or K-effect.
arm of
is
a state of K-tonus,
in
is
Normal
bered,
is
pre-
in
mitted
609
is
a tonic
is
it
should be remem-
surrounding one
coil,
other,
in
may
be induced by
bringing near to
or
of
pole
south
permanent
magnet M.
I shall now exhibit the enhancement or depression of magnetic
A
conductivity by K- or A-tonus.
balanced
record
is
first
obtained,
^1
-J
J
^
then mduced
andJ K-tonus
Successive
right arm.
citations
are
now
xu
the
applied, starting
resultmg
records
responses
of the
duction
K-make ex-
Effect of k- and
A-Tonus on Magnetic Con-
^^^- 374-
are
recorded,
break-effect
''"^^^t^S^ ^J.
wards the
"
JTf
effect
of K-tonus
^^.^^"Tt"';^^
with
resultant
down-responses, show de-
series,
pression
by A-tonus.
being
a de-
R R
COMPARATIVE ELECTRO-PHYSIOLOGY
6io
pression of conduction
of the balance in a
is
induced, as
downward
direction.
We
tivity
strong
next turn to the question of the variation of conducinduced by K-tonus, when moderate or excessively
;
and
it
is
considerations what
is
to be expected
It is
under varying
intensities
them
But
ductivity.
will
enhance con-
maximum
position,
These anticipations are seen fully verified in the accompanying record (fig. 375), which shows the opposite effects
on conductivity of moderate
and strong K-tonus. The
upsetting
of
the
balance
upward direction,
K,
shows the effect of
moderate K-tonus. Strong
K-tonus was next applied,
in
Fig. 375.
strong
an
direction,
k'.
Thus we
see
is
that,
while
under moderate
enhanced, under a
much
greater
it
6ll
made
This
response.
to induce
fail
is
with
concomitant enhanced
its
an
conductivity as
after-
effect.
or habit
is
shall
meet
with again.
excitability
by
that
means
of
ductivity
Balance
determine
the
Conmight
the
we
variations,
not
In
mag-
area
end of
at the right-hand
the balance
may
be made either
by bringing
K-tonic or A-tonic,
it
ward
in
Fig.
a down-
wards
(fig.
still
of kMagnetic
Effects
on
while
From
and K enhancement,
of excitability.
376).
more
interesting case
is
itself fashions, as it
The
is
receiving coil
placed
at
its
own
conduction.
and
Ex-
citability
under
K-tonus excitability is enhanced,
the resulting response being updirection
376.
A-Tonus
COMPARATIVE ELECTRO-PHYSIOLOGY
6l2
(fig-
370
vening
that,
tract,
but
little
excitation reaches
little
in
stimulus
S,
immediate neighbour-
inter-
Excitation at
it.
therefore transmitted
is
second
further, bringing
little
about the same predisposition still further on. Thus an improved conducting-path is made, in a substance formerly but
an
indifferent conductor,
In this
tive,
way
by the
transmitted excitation, at
becomes increasingly
effective
relatively ineffec-
first
(fig.
It
377).
have obtained an
itself.
effect
very
is
exactly
For
by mechanical
response, in
first series
-eP"^-
rll^Tft^tfus"'
It
^"ld
sometimes
be
only after long repetition that conductivity would be gradually restored, as seen in the initiation
and subsequent enhancement of responses given at the distant
responding-point.
It
may
be
that
few phenomena
home
to us, so effectively
responsive manifestations
central
termination,
is
of the living.
locally
excited
The
by
nerve, at
electric
its
shocks,
invisible
nervous impulse
initiates
new
series
centre.
It is as if
set free
moreover,
is
and
wire
line
The
energy.
latent
conductor,
where the
circuit,
of
store
613
return
must
wire
be
In
the nerve
a message, as
again,
it
is
from the
physical
grosser
of
vehicle
while
forces,
psychic impulses
the
conceived of
is
nerve
the
played
as
Under opposite
accentuated.
are
conduction of impulse
is
influences,
interrupted.
Similar
again,
the
results
are
Under
electrotonic action
the transmission
block
is
removed.
Or
electrotonic
action, again,
may be
electric
may be
mission of impulses.
facts
character of the
phenomena
in question.
the
physical
domain
show
also
a striking manner,
that parallel
and
in
phenomena occur
in the
COMPARATIVE ELECTRO-PHYSIOLOGY
6l4
excitation applied at
powder
cell
in
The
source of energy.
as
recorder,
and a
voltaic
on
reaching
responding-point,
the
to
an
electrical
This
indicator.
of
excitation
electrical
the
sensitive
disturbance
substance,
secondary
causes
in
consequence of
'
thus seen
transmitted
that
motile
this
need
stimulus,
the galvanic
response,
not
be
recorder.
It
by the
initiated
proportionate
to
its
circuit
itself.
moreover, as
in nerve, to
changes induced
takes place.
by an
in
Excitation
electrical block
may
and
be
arrested in
magnetic block.
the
impulse
distortion
stored
is
within
It
is
so
it
we
are
means of a
arrested,
the
rod.
Molecular freedom
block,
is
next
and we
re-
find
inhibited,
is
signal.
^:.>
cfj
CHAPTER
XLII
Anomalies
of response
Explicable only from consideration of antecedent
molecular changes Continuous transformation from sub-tonic to hyper-tonic
conditions
first
by means of
Abnormal
characteristic curves,
response characteristic
or sub-tonic state
transitional B state
after
history.
We
in
have seen
living
ditions,
phenomenon of response
very definite.
tissues
is
is
These abnormal
even reversed.
about,
it
by
feeble
effects
stimulation, or
may be brought
by changes
in
the
associated
certain
in
particular
with sub-tomcity.
modifications
The
of the
fatigue-changes
effective in bringing
It is
admitted
be
of
phenomenon of response
further
itself is
COMPARATIVE ELECTRO-PHYSIOLOGY
6l6
There
is
and
it
And
possible to
which occurs,
make
for
by
their
means
always
is
it
As
chemical
versus
nature
of
response-phenomena,
have
make
this
clear.
We may
instance, the
take, for
photo-
This
is
re-
is
liable to disappear,
has
that
is
and
in
to say,
in
is
out of the
question.
Stimulus in general
distortion, the
strength
we have
persistence of which
is
dependent on the
We
self-
have
as
between
When
is
molecularly
strained
and
unstrained
areas.
617
accreted,
may
such
reversal,
It is interesting to
recurrent.
a photographic
sometimes,
reversals
subjected
plate,
becoming
fact,
durations
various
to
negative
in
or
of
positive
From such
response and
antecedents,
facts
its
is
it
variations,
and not
to
we must look
its
to
its
molecular
consequences.
If response phenomena in general, then, are
determined by molecular conditions, as such, it follows that
order to unravel
in
anomalies which
the
we must attempt
occur
in
the
to ascertain those
matter in general.
is
pointed out
Response
In the
in the
work
in
my
and which I
on
investigations
^
abnormalities in response,
'
course of
the
said
convenience,
we observe
its
is
however, brings
'
Bose,
On
to say, to positive.
the
Strain
molecular
Continued stimulation,
condition
to
normal,
1902.
See Response in the Living and Non- Living (1902), pp. 129, 130.
as
Soc.
6l8
COMPARATIVE ELECTRO-PHYSIOLOGY
evidenced
by
the
progressive
response, culminating
This
is
lessening
of
the
positive
in
that
is
alike
by the
staircase effect
seen,
tetanisation,
This
evidenced
is
not
after
now undergo
fatigue, that
is
overstrained.
The
responses
negative
less
negative
a change from
is
to
to say, appears.
may
much
decline to zero, or
We
may
exist in
The
were referred to
*
an
in
shall
earlier
passage,
may
be elucidated
now
as positive.;/. C.^ B
'.'...>....
...;.;
by
this consideration.
explained, in the
6ig
first
chapter of
any one of
are
several concomitant
(a)
(d)
resistivity.
By means
of the
changes.
physical
contraction
and
first
(c)
These
or expansion
of these
we
investigated
animal
and
By
exhibits variations
The
unknown transformation of
dition.
This being
so,
is,
whether we could
matter,
will
we can
ways of obtaining an
COMPARATIVE ELECTRO-PHYSIOLOGY
620
changes occurring in
First of these
it.
method would
lie
in taking a
And
the second
In the second,
arrived at
Fig. 378.
characteristic curves
Magnetisation
and
finally, if
response of the
own
first
to have
characteristic
its
This
under
the action
of
an increasing
shows
the
relation
between
as will
molecular
is
(fig.
external
afforded
induced
378).
force.
The
and
distortion.
characteristic
curve,
essentially
obtained from
is
621
of a substance belonging
filings
Taking
379).
we
condition,
the curve in
That
almost horizontal.
induced
is
stage, B,
first
find
is
which
shall
its
earliest
stage, A, to be
call
We
next
arrive,
transitional,
however, at a
where increasing
force
now approaching
the molecules
Fig.
379.
Characteristic
belonging
motive Force
Particles
to
their
maximum
distortion.
Metallic
Electro-
characteristic curves
occasionally.
starts
in a
still
circumstances
the curve
in
But
it
Under such
is
to
below the
tendency of the
first
zero-line, crossing
it,
remain horizontal
decline, owing, as
we
part of
however,
When
may
may be a
B.
c, it
622
^
;^
COMPARATIVE ELECTRO-PHYSIOLOGY
we
If
under
Fig. 380.
Similarly,
when
{b)
an
In
the after-effect
Referring
is
condition to b\
is
represented by b in
In
or
this
continuously operative
self-recovery
be a certain
there
'
Bose,
On
the
fig.
case
the
we
and continuously
will
378,
find
if
a change of con-
again
new increment
marked
in
substance which
that
found to persist
after-effect is
ductivity.
fall
of
On
substance possessing
back from
persistence
increasing.
its
been
to
after-effect,
b.
But
then a
Electro-motive Variation.'
force
would, when
raise
again
would
applied,
and so
to ^",
it
effect
this
in
That
on.
case be
is
substance to
the
after
623
b^
additive.
Tetanisation
will
This
teristic curve.
will
close
381),
which
tetanisation of nerve
{cf.
fig.
313).
Fig. 381.
(fig.
In
in fig. 378.
exhibiting Transient
all
these
we
horizontality, in an abrupt
the photographic
steel (fig. 381), a
served.
is
horizontal
On
state of balance
is
disturbed,
positive
variation,
in
is
the sudden
brief
COMPARATIVE ELECTRO-PHYSIOLOGY
624
recovery
The
rapid.
is
The
may
ways
by
the response, or
And
by means of the
if
a substance, moreover,
convenient
mechanical,
that
is
The
feasibility of
such records
is
it
may be
Of the
various
no
length.
change to be recorded
is relative,
point which
fectly,
is
invariable.
some
This
or less per-
tis.sue for
to as
little
We
shall
now proceed
characteristics of certain
first
6^25
molecular
states.
We may
take
mode
when
is
Fig. 382.
and
here,
employing
is
it is
apt to
all
we
sources
of
into a condition
fall
sloping line at the beginning shows growing elongation due to subStimulus here causes positive response.
Fourth, fifth, and
Responses become normal
sixth responses are distinctly diphasic.
and increasingly negative after the seventh, with marked staircase
Molecular transformation is seen to be very rapid, above the
increases.
Record is a photographic reduction, half original
B point of transition.
size, of tracings obtained on, smoked glass.
The
tonicity.
record
by an increasing abnormal
given
above.
When
action of stimulus,
its
progressing towards
the nerve
elongation, as in
is
now
tonic condition
is
subjected
fig.
382,
to
the
gradually restored,
is
at
first
The
expressed
from
positive
to
negative,
or
from elongation
S S
to
COMPARATIVE ELECTRO-PHYSIOLOGY
626
contraction, that
is to'
say, in stage B,
we
The second
test,
consists,
we have
Thus,
stimulus.
seen,
the
in
nature of
reply to
its
with
its
tendency
positive.
From
As
the process
is
Or
we do not wish
if
but merely to
observe the
transformation
terminal
into
negative, or
rapid
of stimuli, or tetanisation.
series
according
circumstances,
to
We may
here,
from,
normal
or (2) diphasic, passing into normal
negative responses
feeble, becoming
enhanced, negative
or (3)
negative
been
put
forward,
has
as already said,
idea
The
response.
that tetanisation enhances the responsiveness of the nerve,
by some supposed evolution of carbonic acid. That this,
however, is erroneous, has been shown by numerous experipositive transformed
to
ments already
related,
and by the
fact that
even
in inorganic
Nor
is it
may
the question of
curve at which
invariably true, in
is
applied.
its
The
decisive element in
result will
immediately
after
be an enhancement.
of response by fatigue.
In
the
fact
that continuous
CYCLIC MOLECULAR VARIATION
627
shall
nerve.
frog's
fig.
is
shown by the
upwards.
It is also in-
continuous
the
formation
of
trans-
the
in-
abnormal
the
positive
also
is
noticeable
point of transition
the
is
undergoes
staircase
383
fig-
record,
marked
increase.
is
In
given another
obtained
Fig. 383.
Mechanical Response of Frog's
Nerve, shov.'ing Conversion of Abnormal
Positive into Normal Negative Response
after Tetanisation
Note
with
also the shifting of the base-line upwards, and that the individual period of
positive is shorter than that of negative
responses.
an intervening period of
cisely similar
are observed
in
vegetal nerve.
If
that
we now
turn to a different
by the electro-motive
mode
variation
of observation
the
We
in
records
^say
will
be.
every particular,
same
COMPARATIVE ELECTRO-PHYSIOLOGY
628
continuous
transition,
negative, as
before,
The
negativity.
may
the response
276).
Or when
the
tetanisation
of
effect
is
to
is
intervening
abnormal positive
substance
the
stimulation,
to
is
that,
verted
into
lastly, since
tion
is
that
when
transformed by
and
normal
is
con-
negative,
and
successive
responses
in
that
becomes the
enhanced negative, after an intervening period of tetanisation. The
or moderate negative
Photographic ReFig. 384.
cord showing Conversion
of Abnormal Down ' Response in Tin to Normal
'
Up,' after Tetanisation
underlying transformation
'
line
upwards,
in
is
thus
indicated
now been
in all
transformation equally.
Results in every
substances.
response,
'
In
fig.
way
384
is
represented
as
down,'
converted
into
normal
In
the
629
next figure
in
Fig. 385.
The
transition will
at
Fig. 386.
In
fig.
386
is
seen
in Tin,
how
the normal
enhanced
after
COMPARATIVE ELECTRO-PHYSIOLOGY
630
response
is
a specimen of
We
tin wire.
by the Conductivity or
mode
Resistivity Variation.
selenium
/UUuLwuV
4^
it
aA\\
Anf^^
it
vV^^
V
V
Fig. 387.
..
cell
in
find to
which
Fig. 388.
it
be sometimes
will
in a certain
molecular condition
by an increase of
resistance.
a transformation, attended
variation, of resistance.
light.
Tetanisation
is
found to induce
by a diminution, or negative
After
this,
place
normal
by the diminution of
that
63
now
to say, they
is
resistance.
Fig. 387
Selenium
effects.
take
gives
cells,
again,
condition
be
diphasic
positive
by
followed
negative.
seen to
further
still
further
in the
The
is
now
amplitude in aluminium
period of tetanisation
We
have seen,
(fig.
390).
lastly, that
interesting
by employing
this
(fig.
In order to
specific
enhance magnetic
of the fact
the result of
thetical assimilation
will
is
of record, that
mode
be
391)-
some
may
And it
molecular response
shall
now
determined
and
COMPARATIVE ELECTKO-PHYSIOLOGY
632
from living
bodies.
As
tissues,
Fig. 390,
variations,
follows
./....'
it
after
.."Tetanisation
633
follows that
it
not by some
are determined,
wise,
may
first
vertical
columns
{a)
have
abnormal response
(fig.
a mechanical
by
392)
record
we
of
expansion,
Fig. 391.
in frog's nerve.
In {U)
we observe
FiG. 392.
Vertical Series of
formation of
Abnormal
into
Normal
Response
in
Tetanisation
Inorganic
Living and
alike in the A phase
after
metric positivity
is
here converted by
by
is
tin
electro-
a,
Mechanical response of
nerve to electric
frog's
stimulation
b.
Electro-
variation
in
mechanical stimulation.
COMPARATIVE ELECTRO-PHYSIOLOGY
^SA
is
tetanisation.
be noticed,
It will
in all
from
abnormal
depends,
normal
to
response
shown independently,
also
is
that
is
shown the
to say, upwards.
is
In the next
of tetanisation in
effect
remembered,
molecular
point of transition
normal.
In (a)
of
the
above
the
characteristic
is
condition,
is
just
from abnormal to
(d)
we have
the enhanced
motive response of
frog's nerve.
electro-
In
(c)
we
selenium
^v\iVV
Fig. 393.
cell,
And
light.
Series showing
how
finally, in
Tetanisation ehhances
in the
,
is
see the
Normal Response
B Phase
Mechanical response of
frog's nerve
r,
635
after
mechanical
transformation to
which
this
these cases,
transition.
The responding substance, however, in consequence of the after-effect of stimulation, now passes into
Photographic Record
Fig. 394.
showmg Responses
corresponding with
Abnormal subtonic
d,
Staircase
e,
c,
Uniform
Fatigue reversal.
d,
Fatigue decline
These may
be
short-lived, or
We
shall
more
or less persistent.
next study
all
post-maximum E
Selecting as
conditions, in order.
our specimen for this purpose the nerve of frog, the different
is
of these
is sub-tonic.
It
is
its
characteristic
COMPARATIVE ELECTRO-PHYSIOLOGY
636
abnormal positive. In
however, we have seen this
relaxation to be arrested, and to pass into growing conconsequence
The
traction.
stage
of
is
stimulation,
is
negative.
On
reaching
the
this,
we
We may
staircase increase.
arrive at
arid undergo a
an idea of the rate of
succeeding phases,
{cf.
fig.
We
313).
we
rate of contraction
We
have now
observe
to
the
responsive variations
For
this purpose, a
method of procedure
is
as follows
We
first
take two or
phase.
tained at the
This test-stimulus
same
When
intensity.
subsequently main-
is
till
we
arrive at the
responses to individual
stimuli
stage,
is
is
applied
is
is
now
field.
At
the com-
shows the
Thus the
first
of the characteristic
We
changes
From
response records.
6lJ
that
the
all
responsive
The
attendant variations,
An
inspection of
is
thus complete.
D shows
onset of fatigue
is
This process
the responses.
relaxing state.
The
seen
accentuated, to the
shall later
D and
mediate between
The
E.
ordinary tissues.
upon was
its
One
characteristic
particularly
insisted
On
response,
the two.
show
we
During the
staircase
series of
first
increase.
is
This
uniform responses.
in both, the
process
no
essential difference
between
And
followed,
in
both,
by a
of contracture
gives
place to one of
COMPARATIVE ELECTRO-PHYSIOLOGY
638
relaxation.
makes
its
When
The only
appearance
in
response by tetanisation,
I
stated that here it was not
which formed the determining factor in
bringing about the increase of response this was rather due
to a phasic molecular transformation, induced by tetanisation.
If the substance happen to be in the transitional B phase,
then and then only will tetanisation enhance its response.
If, however, it should happen to be in the optimum C phase,
then the same tetanisation will have the effect of carrying
it into D and E, the phases of fatigue.
The response here,
tetanisation, as such,
Fig. 395.
Photographic
Enhancement of Response
This
is
tungsten
when
moderate
(fig.
tetanisation
reversed.
enhances response,
but
also
is
seen
the normal
contractile
{b)
seen
is
reversal of the
the
639
normal electro-motive
being
also
we have
In {c)
electrical.
And
of Mimosa.
in
given a
is
(<af)
under
variation,
already
by
obtained
given,
been
have
employment
the
of
We
may,
of
Hertzian radiation.
results
stimulus
the
teristic
transformations
about by
We
all
effects,
forms of stimulus
may
stimulus
brought
as
give
alike.
one identical
rise
to
opposite
condition
of the
responding
tissue.
may be brought
about by the
These considerations
will, I
think, be
Series showing
396.
reversal of Normal Response by fatigue due to
FicT.
reference
actions
of large
to
the
and
strong
fl,
small
doses.
Tetanisation
in-
opposite
Mechanical response of
nerve b. Electromotive response of Di'ofrog's
sera
effects
r,
electro-motive
;
response of pulvinus of
Mimosa
by
Response
d^
variation
of tungsten powder.
resistivity
similar to that of
stimulation
Hence
its
effect
C.
Too
long-continued action.
COMPARATIVE ELECTRO-PHYSIOLOGY
640
of the moderately stimulating agent, sodium carbonate, at first induced an exaltation of response, followed
action
by depression.
later
{cf. fig.
297),
we found
In
the case
that the
same
(fig.
facts
first
Even
298).
make
in the
same
Here an agent which proves
a stimulant when given in minute,
their appearance.
Thus
doses.
in
agents on growth-response,
following
summing up
64I
action.
of sugar
With copper
to cause depression.
case
Here,
different.
is
the
constant absorption
of
the
From
all this it is
may
to deal.
Thus,
must
.sive
be determined what
first
is
is
in
examin-
seen to
arise.
And,
finally, as
by
of the tissue.
another hastens
revives
it
in
death.
its
is
transition
response.
there is an intermediate
stage of diphasic
Before exhibiting this in the case of nerve, I shall
were
large,
is
T T
COMPARATIVE ELECTRO-PHYSIOLOGY
642
series then
recorded
contractile
Fig. 397.
rise to
nerve of frog
(fig.
398),
we have
The
abnormal
positive.
characteristic curve
Fig. 398.
a,
we
of
the
before, a
repetition, but in
reverse order, of
all
the
phenomena seen
increasing negative.
The two
being, as
643
The
cycle begins
The
dying
tissue, in
necessary
to revive
a period of rest
To
revive the
is
it
is
essential.
TT
CHAPTER
XLIII
Responsive
in
nerve:
i,
Stimulatory
Electrical;
2,
THE
Mechanical-
Electrical,
brain
as
motile,
their
Law Elimination
ot
and
resultant sensation
Explanation
nerve Effects
and
It
is
retardation.
nerve by
The
the brain.
changes induced
by
falls
physiological inquiry
and
it is
drawn from
may
which
direct experiment.
vanced
in the
sensations perceived in
They
psychological
will
in
some
cases
from their
strictly
it is
much
to hope,
will
tions
645
much
and
same or of
different natures.
The
point lay
in
by
by
sensation.
two
cases,
one exhibiting
it
accompanied during
indications.
It
their
company
the nerve.
It
could
tion
be
was
also
shown
rendered
does
in
fact ac-
extremely
delicate,
ranking,
in
Employing
brain.
mode
this
of investigation, then,
we
in either direction in
this
known
We
have seen, however, that this is not the case, the mechanical
and electrical expressions of the molecular changes in excited
nerve being of a nature essentially similar to those observed
in
muscle
seen
that
also.
Even
in the
The
kind.
But the
relation
between
COMPARATIVE ELECTRO-PHYSIOLOGY
646
quality
What
is
fidence.'
The fundamental
may
it
Taking
well
known
it
'
'
with
'
negative
'
refers to
the reverse.
'
point, sensation
becomes,
first, less
and
less positive,
and then
Or a simple
is
to say,
when employed
or painful sensation.
IS
in
It is
will,
often repeated
when
intensity or duration of
{\^<)\),
\).
2,^T.
The
Stimulus.
647
a physico-physiological
basis
of psychical
effects
Now we
on nerves,
transmitted
as
designated
the
pulse
of
we
The propagation
wave.
positive
This
expansion.
is
have
of this
that
which
is
we have
we found
called
it,
The velocity
with
conductivity, be
alone
So
(p.
made
530).
far then, as
method.
The same
less sensitive
by means of the
facts
method of
may
delicate mechanical
also be demonstrated
ele.ctrical
response,
by the
according to
COMPARATIVE ELECTRO-PHYSIOLOGY
648
also
energy,
form
under
response
of
moderate
in
the tissue
may
stimulation
and
be
Thus we
we
that,
obtain
as
the
is
positive
less
As
and expenditure.
general
rule,
it
will
be
under
cases,
first,
that
in
;
tone.
We
induced by stimulus
attached indicators.
same change
being
shown
electrically
by galvanometric
positivity.
649
In
terminal
motile
indicator
also,
different
signs
namely,
by very
more
I
feeble stimulus,
is
obtained
easily
succeeded
in
Bearing
in
in
when the
is
is
depressed,
At
followed
by expansion,
by recovery.
Just as
in
a nerve in
somewhat depressed
successive
condition,
evoke
stimuli
positive
Fig. 399.
Abnormal Response of Muscle by
Relaxation, followed by Normal Response
of Contraction
The
first
two responses by relaxation
followed by two contractile responses.
are
by
normal
into
so,
the
in
negative,
muscle-pre-
In
fig.
399
give a photo-
all
we
obtain,
by means of the
is
merely
As
diverse
different
registered
changes.
two
COMPARATIVE ELECTRO-PHYSIOLOGY
650
We
say,
to
psychic
the
to
perceiving
organ
conveyed to
it
by
mode
the
of indication, that
registered
effects
positive
and
is
central
waves
negative
We
the
in
have already
contraction
It
matter
is
of
universal
experience,
already
as
rise to sensations
of
same kind
will
or painful.
We
expansion and
when
stimulus,
We
will
galvanometric
intense, will
is
negative
give
give
rise
a wave of
to
rise
positivity,
the
while
same
to a negative wave.
pleasure-bearing, and
Numerous experiments
ferous.
character
will
positivity, as
its
be given,
in
some
being of a crucial
full
support
to this conclusion.
This
fact,
that the
positive
or
the
traction,
negative
painful
sensation
with
muscular
con-
indicator,
will
hold on
the
duced.^
Of the
hold the motor reaction to be secondary to the mental.
it has been argued by James that the feeling does not cause,
The bodily changes, according to him,
but is caused by, the bodily expression.
follow directly the perception of the exciting fact, and our feeling of the same
'
Many
coarser emotions
PHYSICAL BASIS OF
If the sensation
SENSATION
65
physico-physiological
is
known
Weber-Fechner's
theory, further,
ought
tion,
if it
be
to
capable
of
not
explaining
only the
quantitative
relation
demonstrated.
We
more
shall,
crucial
painful
If
tests.
it
be
true
still
our sensations,
that
negative wave
is
We
have seen,
for
instance, that
it
the
its
component
themselves
in
parts,
succession
is
that
which
;
may
also be analysed
thus be
made
to exhibit
the emotion.
all
may
flexion.
COMPARATIVE ELECTRO-PHYSIOLOGY
652
Of
the
may
scheme thus
down, the
and the second
laid
part will be
first
in the succeeding,
chapters.
it
may
be well
central perceiving
apparatus.
As a detector of nervous changes the brain is
undoubtedly the most delicate of instruments, surpassing in
this respect
not only the galvanometer, but also the
Kunchangraph. It fails, however, strictly speaking, as an
accurate
apparatus.
metrical
It
is
means of
and- pleasure
make
are,
it
differences of
receives.
In the pain-
to
extent, of
certain
sensation,
finer
which
able to
is
it
merely qualitative
two extremes
The
But
not only
is
the
a single individual
it
different conditions.
By
deliberate attention
or
inhibition, as
by
raising or
we
Pursuing
find that in
this
coil,
further,
with
its
one pair of terminals, there are many coils with many pairs
of terminals, receiving impulses from every part of the
organism. Confining our attention, moreover, to any single
circuit
among
these,
we
find
again
that
the
impulses
it
653
The
previously absorbed.
is
not
but from
many
circuits at the
same
The
a vague tremor of generalised consciousness.
in
particular
stimulus bears
individual sensation evoked by any
to the rippling surface of this consciousness the relation of
Thus we
will
There
simile of the galvanometer
individual stimulus.
will
be,
the
first
to
the
suppression
of
By
caused
interference
all
revert to the
and, secondly,
by the semi-
may
they
inhibition.
be depressed
to
which
and the
be exalted, while
in others
The extent
may
it
is
claimed
that
this
be carried, would
have myself known of an
practice,
I
appear almost unimaginable.
authenticated instance in which the pulsation of the heart was
In India, indeed, it has been
arrested and renewed at will.
held, from very remote times, that such practices are capable
of reduction to a science.
that
all
It is
one be inhibited,
until
the attention
is
may one by
concentrated on a
any interference
Regarding the physical
aspects of these processes of inhibition and concentration,
more will be said at the end of this chapter.
There are again other elements calculated to bring about
given
point,
in
further
v/hich
variations
lie
more
in
or less
COMPARATIVE ELECTRO-PHYSIOLOGY
654
well
lies in
Even here
posed.
owing
there
to changes in
is
com-
a fluctuation of sensitiveness,
But
neural
the
whose
apparatus,
excitability
the
extremely
is
influence of fatigue,
maximum.
is
then
It
undergoes a decline,
we know, a masked
as
reaches a
it
passing into a
positive
component, which
by appro-
methods.
With
increasing
stimulus
Expressed
is
in
till
a limit
negative
the
reached.
is
effect perceived
pleasurable.
range,
in
positive,
finally passes
commencement
neutral
that
point.
effect,
it
This
is
zero-line
of a somewhat extended
masked
its
must be
not,
however,
same
the
as
absolute
initiated,
effects,
over the
undergoes increase.
see that
shall
till
resultant negative.
tains,
we
gradually augmented
for
zero
at
compounded of
it is
positive
tone
is
complex,
and nega-
655
is
theoretically
by suppress-
tive (pleasure-pain).
it
It is
moment
determines
At
is
it
It
in
general
will
be
sufficient here
to
make
which
may
a cursory reference to
occur.
We
by a
have found
tissue
deter-
is
its
The
to be negative.
illustrated
different effects
several
in
The
cases.
slightly
excitable
We
shall
now proceed
further
to
is
factory
in
variations induced
by
The
it.
standard,
and
in
the
According to what
is
known
as Weber-Fechner's
Law,
COMPARATIVE ELECTRO-PHYSIOLOGY
656
in
order that
The method
arithmetically.
may
increase
result
and even
urged,
its
supporters claim,
Fechner
starts
for
it
only a very
with the
It
or sign of sensation
is
excitation at
more
first
He
expressed an ultimate
generalisation
relation
or, in
It is
now
away from
specula-
and simple
have seen how delicate and free from
We
experiment.
complication
nerve.
effect
is
the record
In these tracings
of the
we have
mechanical response of
the record of the direct
itself
In order,
sliding
electric
stimulation.
inductorium
657
manner by
the
fixing
The
of stimulation.
by increments of one
in-
time
at a
(fig.
is
a sudden responsive
relaxation, followed
by
a partial recovery.
As
is
successively in-
the
creased,
positive
response undergoes an
the maxiincrease,
mum-positive response
Fig, 400.
stimulus-intensity
is
3.
in Nerve of
the Effect of Arithmetically
Record of Response
Gecko showing
increasing Stimuhis
was shown
earlier, is
in fact the
will
9,
This
where
response
to stimulus
Above
is
find, as will
be seen
in the
negative response.
decline,
u y
COMPARATIVE ELECTRO-PHYSIOLOGY
658
we can
lies
rapidly,
first
transition,
increases very
increments of stimulus-intensity.
In order to test the universality of these characteristics of
nerve-responses,
specimens.
obtained
The
with
records
many
said,
different
sluggish condition.
The next
(fig.
On
r,
in
somewhat
2, 3, 4,
it
subjecting this
maximum,
declined,
came
ally
after which it
and the response be-
With gradu-
diphasic.
increasing
positive
the
stimulus,
element
in
the
re-
Response
of Nerve of
Bull-frog to Stimuli i, 2, 3,
.
.
12, increasing in Arithmetical
Progression
FiG. 401.
.
Up
this
point,
of
response,
as
seen
characteristics hold
have
motor nerves.
to
in
same
nerves,
The
I,
2,
3,
659
Records were then made under increasIt will be seen from these
4, 5.
.
is
reached.
The same
characteristics
response of plant-nerve
(fig.
find
403).
in
the case
some
specific
same must
Mechanical Response
403.
of Nerve of Fern to Arithmetically increasing Stimulus
Fig.
And
themselves,
we
further, the
same
next
Even
in
figure.
Hence
it
is
is
by the
COMPARATIVE ELECTRO-PHYSIOLOGY
66o
In
fig.
404 we have a
ampere
to 2
responses at
amperes by steps of
first,
ampere
'2
The
at a time.
rate,
at once apparent.
is
Molecular Variation,
We
curve.
molecular
characteristic
there
saw that
just
beyond
state,
became slower
it
formations in the
by which
phase B to C, and
nerve,
in
higher
the
The induced
condition
it
passes
trans-
of
the
from the
of the
They
Photographic
Record of Magnetic
Responses in Steel to
tion.
stimulus
Fig. 404.
phenomenon of
will, for
intensified,
sensa-
example, explain
proves at
given
first
at
regular
intervals
become
obscure charac-
may
same.
interesting
manner
PHYSICAL BASIS OF SENSATION
being equal, the intermittence
An
per minute.
intensity
falls
of
66l
this
radiation
gether.'
later, fusion
first
sight inexplicable.
next chapter
its
two elements
we have
and nerve
in
particular,
is
this
382).
{cf. fig.
will
to one above
owing to
transformation, the character of the response is changed
If
it
But,
it.
be positive,
it
will
be
be moderately negative,
It will also
solely determined
by the
is
is
not
modified
by the existing condition of the nerve. And this latter undergoes a change by the action of stimulus itself Thus there
are certain times of the day when, owing to sluggishness
of the tissues, our power of perception is dull.
But acuity
of perception becomes enhanced with each successive stimulus
responded to. This is also true, more or less, of each individual undertaking.
These facts are easily understood from
a consideration of the different responses which are characof the ascent of the molecular curve above the point
teristic
of transition.
We
maximum,
with increasing
The molecular
variation,
in
consequence of the
ii.
p.
998.
COMPARATIVE ELECTRO-PHYSIOLOGY
662
by the shifting
upwards of the base-line. We may here refer back to fig. 3
where the responses of two different tissues to increasing
stimulus, show, in the one case complete recovery, and in
the other,
a persistent
The
after-effect.
base-line
the
in
and
owing to molecular
till,
maximum
reaching a
distortion
was reached.
In the second
Owing
to
this
fact,
the
line of
relative
modified equilibrium.
maximum
of
variation
this,
we
the
is
changed condition
moment
of
the
nerve
of
In our sensation
are
line
the
itself
absolute
zero,
any given
which is our
at
this
chang-
ing zero.
The
record which
have given
in figs.
response
is
response by sensation,
we
sensation attains a
maximum
After
663
line of trans-
The
fact,
applied,
efficient to
become negative
as
is
will, if tetanically
it {cf. fig.
383).
We
its
will.
stimulus which
attention, a
may be raised
may be
ceptible
time, again,
excitability,
Thus, by
to sensory prominence.
The reaction
The very
diminished by attention.
As an example
tion
by the
given stimulus.
tion of
of the latter
may be mentioned
the inhibi-
will of the
It
the in-nervation of
antagonistic muscles.
is
held to be
is
It
would
unknown
through
an
character,
it
is
'
result
molecular distortion
is
facilitates the
this
transmission
is
hindered or
blocked.
Thus an
COMPARATIVE ELECTRO-PHYSIOLOGY
664
by the opposite
inducing tonus.
an external force be thus capable, according as it is
positive or negative, of inducing opposite molecular dispositions, by which the conducting power of nerve is so proeffects of the
If
it
for the
it
not
is
under-
difficult to
may
also be varied
The molecular
dispositions themselves,
source.
by which these
effects
are brought about, are no doubt curious, but they are neither
for
we have seen
or
another,
that
by molecular
induced
artificially
may be
in
either
accelerated or retarded.
We have
is facilitated, if
Now
attitude of attention,
we can
considering the
in
if
We have again
found that while incipient contraction or K-tonus enhances
conduction, an intense contraction, or very strong K-tonus,
inhibit,
will
is
already
maximum, and
little
further effect
that
the expression
altogether fanciful
(p.
*
or
external
6io).
steeling
It is
the nerves
metaphorical.
is
stimulus
one of
can produce
to
The
'
is
not
attitude of pre-
rigid contraction.
it
has been
The
negative, on the
rise to
a sensation
665
The
transition,
and slow
later,
equal increments
which
is
at first rapid
is
converted,
We have
may
moderate stimula-
be altered.
In
a magnetic
substance,
the
incipient
depress them.
By
or inhibited.
is
either accelerated
In
will,
like
circumstances, the
may
transmission
accelerated or inhibited.
of excitatory impulses
is
CHAPTER XLIV
DISSOCIATION OF COMPLEX SENSATION
Conversion of pleasurable into painful sensation, and vice versa, by electrotonus
The Sensimeter Mechanical stimulation Stimulation by thermal shocks
to demonstrate,
by means of decisive
with
certain
physiological
appropriate modifications.
response
stimulus
negative
We
shall
now study
sensation with
stimulus,
I
and
changes induced by
under different agencies.
accordance with the chief aim
show,
in
in
the responsive
sensation.
resultant
transmitted
apparatus, was, as
we
effect
recorded
in
the
The
responding
transmitting tissue.
ately the effects
We
6^y
of various
We
stimulation
is
above
rendering
it
pleasurable.
it,
if,
In
this
painful
sensation
lowered.
agents
may
depression
of excitability at
only for
will,
but
the
also
The
ideally
exaltation
for
the
or
rapid
COMPARATIVE ELECTRO-PHYSIOLOGY
668
is
that of
electrotonus.
made kathode,
is
or pleasurable
when anode.
If,
make
it
intensely painful,
it
into
then the
on antecedent physiological changes, we may expect corresponding reversals to take place in sensation, according as
the anodic or kathodic applications are feeble or moderate.
we might expect
And,
teristic
lastly,
changes
to
in responsive sensation,
which
is
way
as to cause a sensation
Under normal
fused
and painful
the
individua
being
responses
become
sensation will
resultant
short-lived
into
and
discrete
669
positive,
the
pleasurable
will
tabulated
summary
in
it
the form of a
Anode
E.M.F.
Kathode
Moderate.
discrete.
Feeble.
discrete.
These generalisations
shall
of stimulation.
have to be
The
fulfilled.
first is
all
so to adjust
its
but complete.
The second
may
fusion will
stimulation
adjustable interrupter.
I
shall
first
when
out.
by means of which
Mechanical stimula-
electro-magnetic arrangement
current
ture
is
is
pulled
the current
is
down
When
is
released,
and the
which
is
determined
in its
turn
by the
intensity
of the
COMPARATIVE ELECTRO-PHYSIOLOGY
670
This latter
current.
a toothed-wheel
of
rotation
the
however,
speed,
is
by means of a carbon
adjusted
rheostat.
certain range
is
axle being
may
be
once
in
four
seconds.
continuously adjusted
This
within
When
is
numbers of
different
governor,
teeth,
Fig. 405.
it is
By
The Sensimeter
possible to obtain
of
frequency.
For
is
this
The
stimulator attached to the circuit, instead of the tapper.
the
regulated
in
be
shocks
may
frequency of the thermal
usual manner, their
intensity being
is
dependent on that of
6/
so
adjusted as to bring
about a
neutral
nail.
In order to study the electrotonic
one electrode was applied by means of a piece of
cotton, moistened in norm^al saline, and placed on the
After
receptive area, the second being on a different finger.
being moderate
that
made
is
The
resulting
By now
made
anode,
and
the
sensation
resulting
was
not only
In
E.M.F.,
employed
indifferent sensation
anode, to
soothing
a feeble E.M.F.
is
When
same
frequent
consequence of fatigue.
It is
way
the opposite
experiments
are
apt to be blurred
is,
and with
therefore, advisable to
different
till
modes of
start with
'Oi
is
stimulation.
volt,
and increase
most pronounced.
it
On
COMPARATIVE ELECTRO-PHYSIOLOGY
672
still
Similar effects
are
now
obtained
also
The
into neutral
on
employing the
electro-thermic stimulator
tapper.
The indifferent and incompletely fused sensation
becomes continuous and painful, on making the receptive
point kathode, under a moderate E.M.F. of 2 volts.
On now
making the receptive point anode, the sensation becomes
converted into a markedly discrete positive. The reversal of
these normal effects under a feeble E.M.F. was obtained in
a given experiment by thermal shocks, when the polarising
E.M.F. was -03 volt.
Turning next to the chemical mode of stimulation, it will
be remembered that an experiment has already been
described (p. 582) where a continuously irritating sensation
was caused by the application of salt on a wounded spot,
this moderate sensation of pain being rendered soothing
when the spot was made normal anode, and intensely painful
when normal kathode. Effects precisely opposite resulted
from the application of a feeble E.M.F. It will thus be seen
that those same conditions which depressed the normal
convert
the
positive
tone
of
sensation
into
negative
or
next
briefly discuss
the
effects
We
that
distinct
effects
and
responsivity,
receptivity,
(p.
induced by
were
conductivity
6/3
493) we saw
it
on the
of
tissue
tion
is
an
effect of exaltation.
by
its
or negative
wave
is
At
And
by causing sudden
this
relief,
will,
as
it
may
Long and
minor operations.
depressing a reagent
Effects, in
some
it is
is
employment
On
for certain
will,
of
the after-effects of so
unnecessary to
respects parallel,
narcotic, there
its
intense alcoholisation
dilate.
may be
observed under
a cessation, not of
may
be 'dissociated
pressure on a nerve
and
it is
'
also
known
Sustained
to depress conductivity,
nerve-trunk will
practically undiminished.
XX
COMPARATIVE ELECTRO-PHYSIOLOGY
6.74
We
shall
We
and
the
other
negative,
Thus
rise
to
the
two
separated
to arrive at the
impulse,
we
The
touch or contact.
rise,
shall
later-arriving negative
wave gives
as
the
case
may
be.
If
the
long,
the
receptive
In other cases,
of
negative
or
painful
This
negative
is
made
the
positive or sensation
With
pain.
greater depression
still
of conductivity,
its
675
two
the
negative
inferences.
and that of these the positive travels faster than the negative,
is demonstrated by the well-known experiment in which a
smart tap is applied on the ball of the foot, with the result
that we perceive, first, the sensation of touch, and then, at an
appreciable interval afterwards, that of pain.
of this demonstration in a normal nerve
is
The
possibility
same
will,
however,
dissociation
The
'
have
dissociation
seen to be effected
considerations which
pain
is
is
first
'
and
'
delayed
being very great, the patient could handle burning coal without pain, the resultant perception being entirely positive.
From
the diverse
previous chapters,
is
phenomena described
it is
in
this
and the
nerve, which
is
negative
Of anything which
effect.
internal energy,
it
may
positive mechanical
and
be said
increases
it
will
With stronger
induced.
thus
in general that
and
stimulus, as
electrical
concomi-
we have
responses are
the
induce
COMPARATIVE ELECTRO-PHYSIOLOGY
eye
masked
positive
diminution.
effect will
this particular
and negative
tones.
stimulation, finally,
expenditure
will
will
be
assume a pre-
The complex
positive,
elements.
it is
is
By
masked
component
capable
of
dissociation
into
its
may
In this
be rendered not
It
CHAPTER XLV
MEMORY
Memory an
after-efifect
after-effects in retina
Spontaneous revival of
their revival
After-
on
excitability
Up
and
in
to the present
effects
entails,
the immediate
We
phenomenon
matter of
known
as
after-effects.
the
that
of
feeble
stimulus.
In
more
is
persistent
very sensitive
the
impression
recovery.
The
perceived.
follows,
The fact
of strong stimulus
in
psychological
general, the
retention
at
than
neurile
is
we
more
of
an
less
rapid
than
required
it
from
feeble,
for the
may
has
its
fading of
COMPARATIVE ELECTRO-PHYSIOLOGY
678
for a longer
manner as follows.
with magnesium powder and
interesting
may
be exhibited
simple design
is
in.
an
made
and
flash
closes
obscured as
any
definite
The
eyes.
his
instantaneous
is
it
impression.
In
the
retina,
at once
flash,
produce
scuring
phenomena of
the eyes
were
We
required
This
is
'
excitable
tissues,
for instance,
nonsense
Certain
find that
Ebbinghaus,
quick,
we
such
found
syllables,' at first
time.
nerve
as
and cardiac
we have
again, under
The
multiple in character.
retina,
which
may
by a galvanometer
be detected
(p. 426).
the eyes
after
strong
Another
light.
or activity.
is
proof
lies
in
This
on closing
that
these
Thus, early in
the morning,
when
fresh
from
and
growing fatigue.
In a given instance, the period at 8 A.M. was 3 seconds,
rest, this
day
to
period
find to
become gradually
be at
its
longer,
shortest,
owing
to
later in the
MEMORY
679
P.M.
and was
On
photographs.
other, instead of
inclined to each
slits
The
inclined cross.
noticed
By
is
eyes are
now
closed,
referred to,
pears, after
At
first
seen with the eyes closed, are very distinct, so distinct that
any unevenness
the
'
made
cuts can be
objective
'
There
out clearly.
is
here no doubt of
after-oscillations
This recurrence
may
with
concomitant
visual
recurrences.
The
recurrent after-image
is
repetition.
the image
is
it is
difficult to tell
or merely an effect of
'
memory.'
There
is,
into
the
In
other.
connection
no hardone merges
in fact,
whether
with
this,
it
is
Visual
for
impressions
and
It
their
recurrence often
persist
some
in-
COMPARATIVE ELECTRO-PHYSIOLOGY
6So
Thus
in
given case
some
in a
time, after
which
On
completely forgotten.
retiring
extinguished his
lamp.
intense.
and
in this case
it
of bright swords.
made
impressions
in the
distracting influences
withdrawn,
is
any
single impression,
is
will,
Since an
significant.
is
or even in spite of
when very
intense,
it
it,
follows
may become
in recurring automatically.
Examples
We
which
in
But we encounter a much more difficult problem when we come to the question of the revival of an
image long after it has apparently faded. It has been suggested that this process of revival depends upon the existence
stimulation.
of some
certain
'
scar,'
disposition
persistent
created there.
brain, or
tendency to
or
on a
movement
It is
some
is
time, yet,
when once
healed, no
It is
of
clearly understood.
are
the
first,
phenomenon
if
we
recognise in
We
at a true explanation of
it
two
distinct factors,
MEMORY
properties
68
upon the
We
sensitive surface in
shall
now
first
tial
revived.
We
must here recall briefly the results which were established in Chapter XLIL, on the modification of response
under cyclic molecular variation. It was there shown that,
under the action of stimulus, the molecular condition of a
substance undergoes a progressive variation, exhibited in
its
effect,
not
to
is
Owing
We
also
saw
that, in
consequence of
frequent repetition
made
a conductor of excitation.
it
In a sluggish
condition, that
is
to
Let us suppose next that by the action of stimulus the subis raised above B.
On the cessation of stimulus a slow
stance
recovery will
indefinitely
prolonged.
The substance
will
may
be
thus approach
COMPARATIVE ELECTRO-PHYSIOLOGY
682
it.
may
mode
We
of inspection.
therefore
be certain areas
the
in
We may
tive surface, in
Thus
excitability.
may have
portion
B,
their
whose
them
latent variations of
represented
by
zero,
another
take a sensi-
consequence of pre-
excitability
effect of stimulatory
excitability
We may
in
artificially
In
still
a third portion,
depressed
tin
wire
or
;
abolished.
its
the
C,
The
dilute solution
of
As long
as the
MEMORY
683
was no effect detected in the galvanoBut as soon as the exploring electrode rested on the
area B, the latent enhancement of excitability there showed
itself by a sudden responsive up-movement of the galvanometer. When the electrode again passed over B and reached
But when it
an indifferent area, A^ response disappeared.
reached c, with its depressed excitability, there was another
indifferent areas there
meter.
responsive
movement,
direction.
It
is
thus
Fig. 406.
this
made by
the
406).
(fig.
way.
We
may be
it
condition.
mined what
From
is
that
till
the whole
is
This
is
kept
reduced to a uniform
we have deter-
a previous experiment
duration of exposure, T, to
a given
COMPARATIVE ELECTRO-PHYSIOLOGY
684
escent response.
stencilled pattern
is
now
placed on the
On now
light
in the
condition, which
is
ferent
A.
feeble
differential action,
We
prove
be effective
We
is
An
to say, an
its
image.
by
The
this
of obtaining a negative
possibility
The
image.'
reversed
or
memory
'
B the substance
rises
in
We
saw that
excitability.
But
region
indif-
now, by
shall, therefore,
that
stimulus should
background
diffuse
in
in
the
the
shall
D and
E,
now describe
Let the observer stare at the incandescent filament of an electric lamp, preferably with one eye, say the
The right eye
right, the left being kept closed all the time.
experiment.
is
after-images will
is
further covered
now be
by the hand.
till
No
Multiple
the impression
trace
of the
MEMORY
latent
image
When
this
now
is
eye-lid,
withdrawn from
light in the
685
suddenly
The
the retina.
is
its
diffuse stimulus to
revived, as a negative
that
is
latent
image
is
Thus the
essential condition
would seem to
The
revival of the
moderate or intense.
I
have already shown, by actual experiment on nervous
tissues themselves, that the differential excitability induced
as an after-effect of moderate stimulus (memory-impression)
will give rise, on diffuse stimulation, to one kind of response,
and the
{cf,
figs.
after-effect
311,
stimulated area, on
induced
In the
its
over-stimulated
area,
on
re- stimulation,
is
reversed, the
becoming galvano-
metrically positive.
The
differential response,
A similar
differential effect
on
is
the sensitive
It
will
thus be
elements,
made
COMPARATIVE ELECTRO-PHYSIOLOGY
686
Diffuse
stimulus,
marked
indication,
psychic.
This
is
may
which
stimulus
be either motile,
applied
is
to
the stem
of Mimosa.
organ
rise to
electrical, or
As
far as this
may
In
leaf.
electrical
by the
delivered
fishes,
will
the
similarly,
internal stimulus,
memory
constitutes a psychic
upon a
will
by the presence of a
It will
latent image.
phenomena of memory.
The
effect
From
the sensory or
latter case
again,
is
memory
is
less
we understand
that
Very
intense stimulation,
its after-effect,
phenomenon
and
is
seen
after
the
When
a considerable
interval
has
elapsed
CHAPTER XLVI
REVIEW OF RESPONSE OF ISOTROPIC ORGANS
Laws of response
Opposite
responsive
of
expressions
true
excitation
and
Position
in
molecular
Effect
sub-tonic positive
and
Similar effects in
nomous response
reversed fatigue
Unmasking
Different expressions
positive
of
tetanisation
Multiple
and auto-
element by overshooting
of antagonistic
of a single fundamental
Explanation
of characteristic
distribution
electric
cylinders
Relative
so-called
Unreliability of response by
positivity of
dead
tissue
plant
in
Reversal
and
of current of injury
negative variation.
all
diverse
muscle
Excitatory
the
in the
response
takes
place
by
contraction
and
galvanometric negativity.
2.
eiFect,
The
first
of these effects
is
the
second,
stimulus
responding point.
In
is
applied
In order to demonstrate
at
consequence
a distance
iof
sudden
from the
local
con-
COMPARATIVE ELECTRO-PHYSIOLOGY
688
This
positivity.
wave of
is
its
short,
is
the
or
conductivity
the
great,
excitatory
masks the hydro-positive effect. But this hydropositive may again be unmasked, by various forms of
selective physiological block, which depress the conduction
of the true excitatory, without interfering to any appreciable
negativity
In this way,
be separated from the contained negative,
the response being thus rendered diphasic positive followed
Or, by the complete suppression of the
by negative.
the positive
may
may
its
amplitude.
The
by the molecular
condition of the responding substance, and the different
character of response
is
modified
also
may
by the
characteristic curve.
From
pass
the study of
and
living tissues.
when
normal
it
is
stimulus
external
in the
will
is
for
When
the energy of
extremely sub-tonic
be absorbed
excitatory expression.
par, that
is
condition,
Response
will
then
be
abnormal
successive
At
this
stage,
only reversed
arrive at the
the previous
normal,
but
At
the
Following
this,
increase.
we
to
D.
Stimulus
680
great.
is
The
by
are
fatigue-
forms of
all
the
sub-tonic
in
tissues.
exhibit a short-
to be indefatigable,
under extreme
fatigue.
is
These are
an intermediate diphasic.
extreme sub-tonicity, and
As
over-stimulation.
definite conditions
regards
the
first,
it
which
siipply of energy
it
from the
Under
forms a part.
brought on by
to be
is
by the
condition of
(i) the
is
rest of the
remem-
maintained
organism of
energy or
formation
may
stimuli,
when
through
an
be seen
in a series
of records to successive
intermediate
diphasic.
Or,
an
intervening
Abnormal
here
in
or reversed response
its
is
abnormal
also seen
sub-tonicity.
In
the
latter,
But
To
Y Y
690
To do
case, stimulation
first
is
is
necessary.
In
necessary.
anomalies are met with, of which there has not hitherto been
any
explanation.
satisfactory
different
give
to
either
tissue
Thus
at
the normal
two extreme cases alike, of subtonicity and fatigue, the response of nerve is abnormal positive
Numerous other examples of this fact have been met
(p. 636).
it
in the
(p. 311),
and that of
seen
in
in, for
how important
is
shown
condition converts
It will
thus be
same
This
tetanisation
the abnormal
is
strikingly
which
to normal, in the
Again,
if
in the
will
tetanisation be
example,
may
be
cited
the
mode
transformation of
tungsten,
after
of record, resistivity-variation.
in
the case of
by the electro-motive
tin,
variation,
abnormal
tetanisation
been shown
radiation,
(fig.
and
mechanical
also
the
As an
386).
(fig.
and
finally,
was shown
fatigue
691
in
it
The
(fig.
stimulation.
change
is
In the
shown
part of this
first
curve a progressive
is
seen to occur.
This
is
illustrated
in records of response
Mimosa, as a
65).
.(fig.
fall
of the
followed
leaf,
Electrically, this
is
by
re-erection
its
These
some cases be exhibited only
phasic
alternations
may
in
quiescent, continuous
(fig.
The
141).
distinction
between
the tissue which gives only one such alternation, and others
The
is
phenomenon of wide-
stimulus,
is
whose predominant
to find an
expression
We may
first
relatively
effective
in
increasing
the
internal
factor,
with
Y Y 2
Comparative^ eleCtro-physioLogy
6gi
At
power of recovery.
the
and
is
to say,
are balanced
and
(figs.
in
vegetable structures
in certain
(p. 536).
on the molecular
factor, is
The
phasic
may
The phenomenon
is
change
(figs.
in question
It will
variations of
any one of
The
these.
therefore be possible
effect of stimulus
may
absorption
nution
changes
changes
expansion;
or increase
of
resistance
galvanometric
of
in
or
resistivity
and
negativity
or
that
first
positivity.
three methods.
by th6 electro-motive
by dimi-
(4) electro-motive
an attendant secretion or
(2)
But
variation, the
The
may
be
in the
method
way
first
of doing this
one
is
This
693
may
be done by interposing a
is
to
The method
viously
employed,
in
had pre-
study of
main
in
similar
difficulty
phenomena
in
living
in
a-symmetrical
tissues
also.
The
excitation,
or
variation,
the
consequent
differential
on
excitability
structure.
which
this
due to
excitatory
movement
change,
the
in
electromotive
response of
is
Similarly,
are taken,
mode
is
of
stimulation,
unless
special
precautions
For this
was desirable to devise some non-electrical form of
stimulation which should be capable of quantitative application
and this I havQ been able to secure by no less than
reason
it
COMPARATIVE ELECTRO-PHYSIOLOGY
694
an
constituted
vibration
effective
angle of vibration.
to
The
vibration
moreover, was
tissue,
My
(p. 31).
The third
(p. 291).
mode of stimulation employed was that of
thermal shocks. The area to be stimulated was, in this case,
determined by the number of rotations
non-electrical
produced by the
in
this
The
case be varied
by
And
have shown that the drawbacks incidental to the electrical mode of stimulation might be over(p.
38).
come by
finally
As
intensity of stimulus
the
gradually
is
increased,
it
On
response.
there
then,
is
an amount
find expression
Under such
latent,
finds
response
tissues in
stems
which
and
this
As examples
may
petioles
is
later
tissue.
held
in
find to
be of very
be observed,
(fig.
of energy
expression
of various
unable to
extensive occurrence.
the
excess
responsive
is
circumstances, the
and often
which
of energy
in the single
may be mentioned
plants
(fig.
138), the
695
to a single
conditions
will,
that
to say,
is
requires,
will
responding plant
merely.
then
It
responds
multiply
by a single
When
stimulus.
is
Or
may
be put
into,
and maintained
a state of pulsation
in,
by the
action of sunlight.
It
is
from
surroundings
raised
as
From
this
that
it
the
maintain
to
will
energy
stored-up
the
condition
tonic
so-called
its
be seen
derived
from
of the plant
autonomous
is
its
so
activity.
no
the
of Desniodium gyrans,
it
ment due
of
the
to recovery,
organ,
is
lateral
relatively
excitatory impulses, in
tively,
is
is
of
1*5
to
i.
This
COMPARATIVE ELECTRO-PHYSIOLOGY
696
explains
the
peculiar
which
gyrans^
are
electrical
responses
concomitant
with
leaflet.
of
find
Desmodium
autonomous
the
that, corre-
two
electrical pulses.
wave
is
nearly
1*5
These electro-motive variations are expressions of fundamental excitatory effects, and not dependent on the
mechanical movement of the leaflets.
For when the
responding
electro-motive
be seen
in the
An
fig.
145.
demonstration
excitatory
This
reaction
of the
which
that
fact
expressed
is
the accurate
in
in
is
single
different
ways
temperature, there
excitatory spasm
If a continuous record
is
occurs,
point
definite
marking the
at
which
initiation
an
of death.
a sudden contraction.
In
an anisotropic
of
the
death-point,
sudden uncurling.
method of record
the
increasing
positivity
the
increasing
is
seen
to
resistance.
Taking,
become
of
the
Finally,
resistivity
suddenly
It is found,,
how-
697
ever,
and
electrical
constitute
by
the
fact
case
true
that
of excitatory response,
induced physiological
The response
is
later
much
is
proved
depression
Fatigue
also
may
thus
as 19 C.
the
excitatory galvanometric
moreover,
effect,
to
of
after-effect
persistent
greater
is
or less
is
negativity
The
strong.
undergoes
effect
excitatory
As
mitted
is
diminution with
this trans-
distance,
will
is
it
be induced at
isotropic tissue,
shall
we make two
clearly have
most positive
(fig.
we
no).
Two
is
two
relatively
potential, a
to
COMPARATIVE ELECTRO-PHYSIOLOGY
698
by experimental
are
results,
often
is
But
living.
have shown
that
this
negative
is
to
is
of death, a
tissue
excitatory con-
exhibits
Proceeding
point,
it
this
or
dead portion
is
dead
dead
these facts,
is
current
the
is
to the living,
former,
From
this
is
relatively positive
the
positivity of the
From
maximum
But the
decreasingly negative.
of
living
from
direction
either
in
is
it is
on the
(figs.
death-frontier.
it
negative,
is
injury
the
In the case
liable to reversal.
of
and the
clear that
be from the
will
Now, by the
may
as
be
made
positive
to appear,
variation.
is
held
to be
reversal of the
is
sufficient to
indicate
Method of Negative
the
Variation,
699
many
glaring instances.
An
obtain
assumption more or
less current
it
is,
that in order to
an antecedent current, by
can
be
Hence
detected.
the
The
real
contacts
reason,
is
so
however,
to
depress
for
its
thus
injuring
excitability
one of the
on diffuse
that,
as
such,
resultant
response
which
is
response,
may
still
current of injury
is
the
is
may
it
is
essential
seen
That
condition
from the
fact
for
that
obtaining
excitatory
(fig.
116).
CHAPTER XLVII
REVIEW OF RESPONSE OF ANISOTROPIC ORGANS
Anisotropic organs Laws of response in anisotropic organs
Natural current of
due to induced
Electro-motive
peculiarities of skin
negativity in
structures
-Similar
effects
with
inorganic
of
on plant
tissues
after-effects
like,
after-effects in retina
SHALL next
many
I
supposed anomalies
in the
is
at the root of
kEViEW OF
OF ANISOTROPIC ORGANS
RESPOiSrSE
As an example
than
the
In
upper.
stimulation of either
this
case,
^Ol
we may
is more
strictly localised
the
we
response of
anisotropic organs.
On simultaneous
1.
excitation of
two points
and B, the
responsive current flows in the tissue from the more to the less
excited.
2.
current be from
to A,
is
points.
The second
As
in
a simple
we may take
surface,
constantly
and reduced
in excitability.
it
subjected
lily.
to
were fatigued,
and protoplasmically
excitability on
this
defective.
Owing to the depression of
epidermal surface, the intensity of its
in
responsive positivity.
may be
The
COMPARATIVE ELECTRO-PHYSIOLOGY
702.
excitable,
to the less
Such
is
of injury or accidental
effect
more
in the
rest,
is
to say,
excitable
to
B, this reversed
B-^A.
than
yi,
less
A ^
will
induce a responsive
current from
to
more
is
it
is
necessary, as
now
under-
is
lation as the
(i)
the
When
or (3) stimulation
this
it
is
(2) the
by thermal
according
obtained
are
results
For
Vibratory Stimulator;
In order to do
employ
possible to
^O^
to
these
when the
The last-named
may, however, be used without misgiving, when stimulation is effected by equi-alternating shocks.
The ordinary
cations
electrical
form of stimulus
employed.
is
this purpose,
for
make and
The
may
equi-alternating shocks,
(2) a
motor-dynamo
under these
(fig.
172).
The
forms of stimulation,
electrical
responses
may
again,
be photo-
It
all
these
anisotropic organ
was
definite
in
obtained with an
direction, being
always,
by these methods
in
various cases of
normal response
by galvanometric
much
negativity.
stronger
On
simul-
COMPARATIVE ELECTRO-PHYSIOLOGY
704
Here the
outer.
current
resultant
is
brought
by
about
may
which
be represented as
With
j^.
certain
is
great
so
positive, that
strong negative.
The
that
we
this
to
pass,
the
summated
is
effect of the
The
and
has been
conclusion
skins,
may
verified
less
excitable
Among
response of
This
outer.
and that
care, so as to
excitable
outer to the
i.
be said
from the more
is
minimum
two
may
it
from
by experiments on various
reduce to a
is still,
is
From
response
individual
its
is
from the
surface;
it
less
the
Owing
to
The
responsive current,
excitable.
In illustration
may
human
forefinger
(fig.
i8o).
peduncle of Uriclis
are, as
rule,
REVIEW OF RESPONSE OF ANISOTROPIC ORGANS
705
is
opposite,
human
lip, I
Again,
196).
(fig.
on
testing
the
differential
As we approach
regarded as epithelial.
and secretion
The
is
is
But
In
all
these
is
by strong galvanometric
DiUenia
indica,
it is
negativity.
Taking a carpel of
is
from
the outer epidermal to the inner secreting surface, the responsive current being in the opposite direction.
On making very
body of
its foot, it
is
is
As an example
may be
showed
of the
way
in
which the
of '0013 volt
after
zz
COMPARATIVE ELECTRO-PHYSIOLOGY
706
natural current
was
induced at the
excitation
became relatively
With the intact human tongue, further, I found that a
very strong responsive current was induced on excitation,
from the lower to the upper surface, thus showing that the
lower was the more excitable of the two.
The response of
digestive organs
may now be
passed
in
review.
excitation
re-absorbed on
again,
is
know
the pulvinus
regard this
secretion as
an
fluid,
which,
cessation of excitation.
be contractile, and
to
expulsion of water.
the
effect
may
We
therefore
of contraction, causing
differential
action of
the upper and lower halves of the organ, and the magnifying
petiolar index, the
in
This goes to show that it is not impossible that the phenomenon of secretion through a permeable membrane may
shown
In digestion, as a whole,
we have
to
recognise two
sequent absorption.
electrical
Parallel
to
these,
we
and of sub-
find
that
the
organs.
current of
which
is
of Nepenthe in
pitcher
opposite
the
in
and the glandular surface, on simultaneous excithe two, becoming galvanometrically negative
direction,
of
tend to -exhibit
203).
(fig.
a fresh
is
tation
the normal
reverse
necessarily to
With a
rest.
707
206, 209,
(figs.
when
is
number of captured
change,
a phasic
becoming reversed
out above,
213).
to positive
(fig.
205).
This, as pointed
to positivity
In
is
on long-continued stimulation,
(fig.
now
responses
the
negativity, but
that
flies,
it
exhibits
it
and
In DroserUy
is
this
by induced
is
reversed
208).
is
From
the fact
that
the
skin
of the
toad, which
is
also
it
frog
motive reaction as
That
the case
is
its
outer skin.
electro-
however,
is
not
becomes galvanometrically positive, while the mucous membrane of the stomach becomes galvanometrically negative.
The observed current of rest in the stomach would appear,
from Nepenthe, to be, not the natural current of
rest,
but the
The normal
and 212).
stomach of
210, 211,
to the
On
frog, I
ZZ2
COMPARATIVE ELECTRO-PHYSIOLOGY
708
diphasic,
consists
first
materials
are
by the
first
The proof
absorbed.
root,
dissolved
of the
known
corrosion-figures produced
marble
surface.
is
seen
by growing
in the well-
on a
have also been able to demonstrate the
rootlets
phenomenon of excitatory secretion in young roots by allowing them to absorb dilute salt solution, and then under excitation to secrete
This
last
it
precipitate.
The
Colocasia, moreover,
electrical
I
response of young
roots
of
is
is
as-
by the plant
For
in the
first
supposed dead
709
transport of poison,
it
elements concerned
in
are
diffused
throughout the length of the trunk, the death of one individual zone, to which the poison has
when the
of the solution at
its
It is
by the
only
arrival
And
this is
found to be
the
case.
sponding modifications in
the
rate
suction.
very
such as cold
or
anaesthetics,
it.
It
when
depress
or
arrest
suction,
Desmodium
leaflet
falls
is
below
into abeyance,
COMPARATIVE ELECTRO-PHYSIOLOGY
^'l-O
is
when
longer
the plant
its
is
in
tonic
stimulation.
that
is
nervous surface.
discharge
is
is
which
in
it
is
is
direction
that,
yet
in the
has
always
is
organs
electric
homo-
or hetero-dromous.
will
fully
in
same
whether
No
explain
all
theory
these
peculiarities.
I
of
its
nor
is it
this
phenomenon
dependent on any
is
not alone
specific characteristic
The response
of the electric
Hence the
current of response
is
the less
In
taking rheotomic
acting, as has
is
observations on
was attained
'2
the response
electrical plates
of moderate
maximum
that
is
it
which elapsed
With vigorous
of NymphcBa alba,
leaves
was attained
in
This was
mechanical stimulation.
effect
was found
to
virtually
the
tion
7 II
much
shorter time,
or
multiple.
In the
character
is
shown
to
of definitely uni-directioned
dromous
of
is
not distinctive of
assimilation
general,
with
its
and
homodromous
life,
with
dissimilation,
but
consequent
its
of
differential
or hetero-
specific
powers
anisotropy
excitability,
in
was
We
(fig.
167).
COMPARATIVE ELECTRO-PHYSIOLOGY
712
effect of this
normal and
like that of
These
by the
results,
however,
may be
when
of
is
light.
the upper, the expelled water reaches the lower half of the
pulvinus
the hydro-positive
is
half,
it
electrical
itself
is
But as
The
mechanical response
is
now
of
variation at
lower
The
effect
From
this
experiment
it is
(fig.
237).
by mechanical response.
We
see,
moderate stimulus.
Owing
fall.
its
application
is
strictly local,
it is
and
easy to
/1
Thus,
in
by galvanometric
tissues,
and the
negativity,
be conducted to a certain
But we have seen that in tissues which are not
distance.
we
light,
and
positive.
stimulation
light,
of Bryophyllum
plified
(fig.
(fig.
of
)
or
to direct
response
exem-
in
240).
been
has
It
responses
lead
how
explained
us
to
the
these
alternating
phasic
autonomous response.
leaflet of Biophyttim^ or
Desmodium
leaflet in
a state
The corresponding
(fig.
by the
fact
more
effectively
by the
arrest
maximum.
normal alternation is ( +
particular phasic
of external
Thus,
in
stimulus at a
1-), if the stimulus be suddenly withdrawn at the end of the second phase, or positive
maximum,
(figs.
243, 244).
in this
Type
I.,
The
characteristic direct
and
after-effects
and
its
removal are (
In specimens whose characteristic
V -f).
response under continuous stimulation is (+
if
),
1
stimulus
be
the
second
COMPARATIVE ELECTRO-PHYSIOLOGY
^14
phase
in
here
negative
maximum the
after-effects in this
Type
(figs.
response
The
245, 247).
III., therefore,
may be
overshoots
direct
and
represented
or negative phase, or at
first
maximum
of the second
or positive phase.
(- +
,.. ),
while
The response
striking
(a)
is
thus
).
light.
The
to be unlike
and
show response by
the response of the retina was referred
muscle
variation,'
*
were
said
positive variation.'
the
confusion
which
is
apt
to
cornea, in
result
the standard
the longitudinal
'
negative
by
to as
resting-current
so-called
to
surface of the
from
of
making the
reference.
On
an undetached eyeball of
found that
it
is
frog,
excitation on the
the nerve
and
this
715
These are
tissues.
the
in
first
place
and
sub-tonicity,
secondly, fatigue.
the
first
retina,
excitability.
ot
stereoscope
specially devised
may be mentioned
In this connection
(p. 432).
phenomenon of Binocular
the interesting
Alternation of Vision.
The
vegetable
the
under light
tissues
correspondences
in
I.,
we have
highly excitable
Type
find to
the formula of
-f-
flower,
to
and
direct
excitable
be
after-effects
observed.
is
affording
(-1-
mentioned
...).
the
us
Type
In
III.,
as
the
the
In
of
the
highly
of cauli-
correspondence with
response
less
petiole
isolated
this
retina
may
of
be
fish,
supposition
COMPARATIVE ELECTRO-PHYSIOLOGY
7l6
the
...).
frog.
electrical
In
more
The
vertical.
curvature
might
The
question
its
to say,
by excitatory
described
(p.
The
tests.
contraction.
in
436)
which
of cold
is
was subjected
known
its
that
is
this question
application
local
An
to
to
bring
effect,
and
the active
test
of
side,
retarded
it is
This
factor.
electrical
it
gravitation.
is
is
response,
when subjected
in
where
find
that the
upper
namely, by
induced galvanometric
negativity.
particles.
When
the stem
is
tissue,
by the weight of
vertical, in
solid
consequence of the
This, however,
expected to be
initiated.
The shoot
first
it
is
is
This anomaly
is
717
first
sign
probably due
by weight which
may, however, regard ourselves
has
first
to be overcome.
We
when recording
by geo-electric response.
we have seen in other cases,
The
takes
place
as
before,
when
all
responsive
Proceeding on
mechanical
principle,
indications
are
therefore,
restrained.
this
tational stimulus as
to
many
CHAPTER XLVIII
REVIEW OF RESPONSE OF NERVE AND RELATED
PSYCHOLOGICAL PHENOMENA
Transmission of excitation
in
plants
Vegetal
nerve
Similar
variations
of
receptivity, conductivity,
Multiple
excitations
in
Negative
after-effect
subjective response
Under
to
it
artificial
vice
stimulus
at will
after-effect
differential excitation
Same
effects
demon-
The
of stimulus.
known to
known case of Mimosa^ where stimulus
ment at a distance, this was supposed
dro-mechanical disturbance.
do.
is
Even
in the well-
RESPONSE OF NERVE
movement
(figs.
719
and
44, 45,
of the
The
46).
fall
metric negativity.
the plant
is
conditions
which influence
hand,
is
the
in
it
case of the
animal.
is
feeble.
Fatigue, on
The
the other
application of
warmth enhances
to
its
velocity.
depress conductivity.
And
lastly,
the
is
polar effect
of
to induce opposite
applied.
have,
fitted
for the
soft
conduction of excitation.
easy to isolate
in the case
may
is
be
teristics,
of
fern.
about 50
view
and the modifications of
They
mm.
per second.
It
said, in
their response
under given
from
be rightly designated
example, these highly
nerves.
On
may
therefore
isolation,
for
which
their
fall
when
isolated,
conducting power
is
temporarily impaired.
The
metric positivity.
of
the
galvano-
in this state,
into
the
response into
COMPARATIVE ELECTRO-PHYSIOLOGY
720
ammonia
effects
are the
same
in the
one case as
The
in the other.
same
finally, in
in
as on those
alike
same
salts,
the
characteristic effects
(Chapters
XXXII. and
nerves
was able
to devise
Conductivity Balance
(fig.
291).
ability,
applied
It also
of two
enables
different
The
principle
Balance depends
is
is
exactly
midway between
then zero.
the
When
conductivity of
say,
up
c,
E, or
of the balance
is
it
in
the
RESPONSE OF NERVE
was due
to
72i
molecular transformation
the
condition B
(fig.
to
At
311).
is
to induce
moderate
the
itself,
to condition
excitability.
or E,
As
and the
result should
In accordance with
is
this,
it
is
be one of loss of
is
depression of excitability
found that at
(fig.
2).
economy,
a
them
by the
The effect
caused
tissue,
it
may be
by which
said
that
the
normal excitability of
its
It is
tion, the
It will
of
this, its
is
The
when
The
fact that so
whole
its
tonic condition
long as a portion of
it
is
is
energy of
light.
COMPARATIVE ELECTRO-PHYSIOLOGY
*J2Z
is
whose
have demonstrated.
In
the case of trees, again, the interior tissues whose functions
are of great importance in various ways, are inaccessible to
plant-nerves,
is
far
light.
The
energy as that of
such external
them
existence
in the ramified
But no part of
whose outer
nerves,
An
its
334).
I
the
reaction
It
of
being incapable,
sive
it
was
change of form.
for the
study of
the
It
is
specifically
said, of detecting
I
this
tinuity
In
original length,
or more.
afforded
by
observer
is
my
moderately sensitive
able to study
all
times, which
is
Kunchangraph, the
the excitatory
phenomena
in
Records of the
by
when
may be
RESPONSE OF NERVE
The
positive response.
is
complications.
various
yz:^
we have
the
and not
direct,
is
electrical
employed
differential.
Again, the
almost
universally-
liable
by leakage,
is
of nerve,
is
mode
is
When
of response.
the
By means
trans-i^
An
itself,
may
generally
falls into
an
as
record,
increasing
abnormal
positive response, of
relaxation
and the
this
point an abnormal
sudden expansion.
Successive or con-
from
abnormal expansion being
arrested and converted into increasing contraction.
During
this stage, then, the responses to individual stimuH are transformed from the abnormal expansive positive to the normal
contractile negative, through an intermediate
diphasic.
tinuous
stimulations,
condition
to condition B, the
Molecular transformation
is
(fig.
now have
response
In
ing the
(fig.
383).
382).
and the
An
re-
intervening
we
its
different
change from
The next
are uniform.
stage to be reached
After
this,
we
is C,
arrive at D,
(figs^
275-278).
A 2
COMPARATIVE ELECTRO-PHYSIOLOGY
724
make
begins to
its
appearance.
Up
But
after D,
the
responses
it
individual
to
stimuli
From
of the other.
with
abnormal
its
till
are transformed
They
B.
here
to
C.
fatigue,
brought on by overstrain, with its diminishing normal responses at D, through diphasic, to abnormal positive once
Excessive sub-tonicity and excessive stimulation
more, at E.
opposite.
In the
in the second,
first,
it is its
which
is
is
necessary
required.
Similar effects are also met with, in the case of transIn the sub-tonic condition, conductivity
mitted excitation.
is
By
the action
of
is
abnormal
positive.
stimulus,
is
is
negative
converted
through
from
the
abnormal
positive
intermediate diphasic.
to
normal
After
this,
under increasing fatigue the diminishing responses are converted to abnormal positive through an intermediate diphasic
Another important demonstration was that of the
(fig. 325).
perfect similarity of the molecular changes induced by
stimulus
in
the
afferent
and
efferent
nerves
respectively.
RESPONSE OF NERVE
just
725
nerves
certain
fishes
With
obtained
324, 402).
optic
That
is
may be
The
difficulties
due to
light,
by means of a writing-
By means,
however, of a battery of levers, and using the optical method
of record, it is possible to have a magnification by the
obviated by the use of the Oscillating Recorder.
By
this
muscle.
it is
found
Now
it
this
If the
is
we know
(fig.
to
drying of the
326).
And
is
the
thus
demonstrated
With
clearness.
Under
COMPARATIVE ELECTRO-PHYSIOLOGY
726
On
abrupt
the
cessation
of
tetanising
shocks a
is
that of electrotonus,
-^s
33^)-
by Bernstein
Hermann
polarisation-decrement, whereas
as
known
are
as
polarisation-increment.
those of
That
is
to
say,
may
influence
of an
electrical
in the correct
difficulties
current.
One
of the principal
Pflliger's
Law,
of universal
to Pflliger's generalisation,
have shown
the anode
it is
depresses
it.
This
RESPONSE OF NERVE
727
great simplicity
sensation.
The
irritation.
wound now
anode
at
once made
it
held
1*5
was reduced
volt,
to
'5
volt.
against
have shown
it.
is
to say,
when
mission.
is
same
direction as that
The normal
responses
reversed to positive,
are
then
diminished, or
by the diminution
even
or abolition of the
The
more excited of
two points, the responsive current being algebraically summated with the existing electrotonic current. The greater
excitation of one of these two points was also shown to be
due to the greater intensity of excitation conducted to it, or
increased galvanometric negativity of the
COMPARATIVE ELECTRO-PHYSIOLOGY
728
or to both.
the hydro-
tory or negative.
If the
stimulus
wave alone
will
applied,
moreover, be
be transmitted.
If the
one wave will be superposed over the other, the negamasking the positive. The two waves, however, may
short,
tive
owing to
sufficient
may be made
positive
tively.
we have
also
the
concomitant
electrical
expressions
have been
of
able,
which
may
rise
to
the
be described as
The
positive
tone of
pleasurable
or
at
which
with
its
is
first
different
In various
RESPONSE BY SENSATION
729
is
logically.
may
be held in
the hand and induce only the feeling of contact, without any
sensation of pain.
One
very
difficult
logical response
is
intensity of stimulus
known
tion
problem
in
The
generalisa-
must increase
in
arithmetical, progression.
relation, not as
factor,
On
nerve, given in
figs.
we
see that
We
positive, connoting, as
After
this,
stimulus
feeble
we know, a
the
response
is
From
negative.
point
this
These
maximum
molecular
distortion
facts
such considerations
changes
in the
we
an
obtain
explanation
characteristics,
again,
show a
and
of
in
those
is
mole-
are
not peculiar to
That
That these
the
nerves
and response
be
understood from the response of an inorganic substance to
(fig.
403).
this
relation
indeed
fig.
is
universal, will
404.
COMPARATIVE ELECTRO-PHYSIOLOGY
730
by the polar
action of currents.
more
relatively
has been
It
short-lived,
is
shown that
is
405),
intensity
which
is
If
we now render
thus reduce
its
appear
all
but continuous.
be converted to
If,
next, the
become converted
into
It will
we have
into another,
seen, to convert
it
is
stances, then,
It is
less
ex-
modification
indeed
By
in
any
induced dispositions.
direction,
The
opened up by the
line of
thought
here suggested.
The
phenomenon ot
memory, which is an after-effect of stimulus. The aftereffect of strong stimulus is in general more persistent than
last subject
that of feeble.
to be reviewed
Similarly, the
is
memory
the
of a strong sensation
RESPONSE BY SENSATION
of reviving memory-images.
Long
73
Of
this
metaphor, however,
it
may be
said, that
though, no doubt,
effect
after-
sensation, yet,
itself,
To do
it,
this,
of
the
original excitation
regarding
as a scar,
it
we
There
is
now nothing
visible
by which
to distinguish
In consequence
The
isotropic,
by the impression of
stimulation,
the
differentially
Thus,
isotropic,
has
now become
excitable
On
an-
diffuse
structure will
now
will,
we
and
COMPARATIVE ELECTRO-PHYSIOLOGY
732
may be
On the
if
image
once more appear as a bright patch against a dark background. This is because, as the after-effect of stimulus, the
will
area
stimulation evokes
its
is
acting
sation,
as
stimulus to
a diffuse
light
picture.
The
phenomena seen
responsive
by the inorganic
are no less
matter
living
in
By
wonderful.
ascribing
By
that
siveness
we
attain an
itself,
is
all
by con-
new
possessed of sensibility
are,
is
order,
closed.
any ultimate
analysis.
We
are led
by
it
to the discovery
responses of the
most complex
inorganic matter.
we
living matter,
living,
nomena
suspected.
this
the responsive
phe-
electrically.
responsive, to
all
the one
is
common
indistinguishable
In
excitatory effects,
or
All
Both
alike
are
responsive,
and similarly
RESPONSE BY SENSATION
733
tropic
same
peculiarities of epidermis,
are
plant,
like
the
in
animal,
known
by the agency
the re-
phasic alternations
anisotropic
is
The
and
all
its
As
as nerves.
its
nated,
responsive
and gland
discharge of an
electrical
same
the
The
both.
in
epithelium,
in the plant-nerve,
moreover, so also
two
distinct impulses,
It is
find manifestation as
known
The
are
under
positive
These
and negative, pleasure and
parallel
conditions,
as
the
and negative
positive
And
example of the
differential
finally,
effect
impression
which remains
memory-image.
surfaces,
perhaps,
of induced
made by
latent,
the
most
anisotropy
stimulus on
and capable of
it
significant
lies
in that
the
sensory
revival, as the
1.
PAGE
2
Electromotive response
.......
.....
2.
Response of
3.
4.
5.
Action of
6.
Response by
7.
8.
9.
'
tin
poison
'
resistivity variation
.......
...
11.
Electrical
12.
electrical response
Mimosa when
pulvinus of
physically
14.
15.
16.
17.
18.
19.
...
.......
.......
.
25
25
26
26
27
27
-32
23.
26.
19
25, 38
22.
25.
20
.
24.
re-
strained
13.
in plants
10.
of
... -9
response
.48
...
.
34
43
43
39
40
41
COMPARATIVE ELECTRO-PHYSIOLOGY
736
Mimosa
-55
by negative in Biophyticm
by negative in Mimosa
30. Simultaneous record of positive and negative mechanical and electrical
.
29.
31.
Positive, diphasic
32.
response in Biophytum
33.
Effects
on growth
58
59
61
62
in petiole of cauliflower
.
64
66
73
of temperature on growth
response
74
Abnormal
37.
38.
39.
of
44.
Mimosa
.103
.91
.91
.........
83
.
in
92
93
96
mechanical response
95
95
.....
46.
47.
.98
50.
51.
Differential response of
98
99
.102
49.
100
104
compound
108
strip
109
Cucurhita
no
ill
its
Effect of rise
57.
58.
Effect of
NagCOg
solution
59.
Effiect of
CO2 on
current of rest
60.
on current of
on current of
.114
Variations
56.
rest
rest
119
119
122
122
127
717
Effect of chloroform
62.
.130
...
..........
Effect of
NaOH
65.
Effect of
KOH
66.
131
132
134
.
67.
68.
Effect of
69.
Effect of moderate
135
...
136
136
HCl
138
KOH
139
Method of Interference
70.
Responsive
71.
72.
effect
of variation of phase
its
144
variation
147
147
Electrical after-effect
lation
in inorganic substances
.
-151
74.
75.
stimulation
I55
.158
152
.154
Positive' Variation
due to section
165
.169
of injury
171
174
Effect of Temperature
autonomous pulsation in Desmodium
warmth on autonomous pulsation of Desmodium
82.
83.
Effect of
181
182
183
184
I8S
186
on amplitude of response
on conductivity
.
187
hy high temperature
^
90. Effect of cooling on frequency and amplitude of pulsation of Des
89. Abolition of response
modium
,..,.,.,..
3B
190
188
COMPARATIVE ELECTRO-PHYSIOLOGY
738
Determination of Death-point
PAGE
..........
.
195
198
202
204
Biophytum
Mmiosa
97.
98.
99.
100.
Autonomous response
in ^zVj//^_>'/w
....
sating
Desmodium
leaflet
Record of
105.
212
.214
restrained
209
210
210
.211
..........
104.
is
Z>^j;^^zz^;//
207
208
217
when movement
.
220
221
Response of Leaves
106. Effect of section of petiole of Ficus on the current of rest
107.
108.
109.
227
226
225
Musa and
that of
Dionaa
237
110.
111.
113.
115. 'L&2X oi
116. Y\\.(^Qx
117.
Pterospermum
of Nepenthe
Pulvinus of i^/iw^ja
in
245
248
252
266
256
255
256
268
257
267
283
285
285
286
118.
'121.
122. Eel
123.
739
Response of Skin
124. Current of rest in vegetal skin
...
PAGE
298
.........
.....
....
.......
of grape skin
126.
299
300
Response
129. Response
130. Response
131. Response
128.
tomato
307
301
302
303
308
132.
309
133.
134.
307
300
135.
136. Illustrative
...:.......
......
variation
137.
138.
303
304
310
after tetanisation
311
318
140.
141.
Response
Response
Response
Response
Response
319
315
316
142.
143.
144.
145.
of water-melon
........
........
146.
319
320
322
Normal response
147.
326
328
151.
152.
342
157.
339
339
after
tetanisation
sation
335
338
345
345
after tetani.
346
when
341
fresh
3 B 2
740
COMPARATIVE ELECTRO-PHYSIOLOGY
PAGE
158.
159.
160.
161.
Electrical response of
...
.....
343
347
353
.......
....
164.
355
...........
its
wood by poison
167.
168.
Effect of cold
169.
170.
Effect of anaesthetic
Excitatory
on suctional response
z'<?ri/<j
362
.......
....
......
171.
354
352
^63
366
372
372
on suctional response
375
376
osmotic action
383
384
386
388
173.
397
400
180.
Transmitted
effect
Mimosa
399
to unilateral light
of stimulus of light
401
402
183. Multiple
402
406
light
405
184.
185.
407
408
186.
409
187.
positive after-effect
and
under
light
411
after-effects in plants
under
412
light
Response of Retina
189. Determination of differential excitability as between optic nerve
and
cornea
190.
............
...........
nerve
agents
193.
417
Normal
419
.418
by excitatory
423
427
741
PAGE
194.
195.
after-effect in
Ophiocephalus
430
and
426
'
human retina
in human retina under continuous
light
.....
.
433
430
432
431
432
Geo-electric Response
202.
......
437
436
440
442
Velocity of Transmission
206. Determination of velocity of transmission by mechanical response
447
448
....
....
....
449
449
450
452
method
454
tetanisation
...........
abnormal
through diphasic
positive
to
457
460
462
463
normal negative
.
-465
by increasing
intensity of stimulus
466
22 1
Effect of alcohol
....
..........
ammonia
of ammonia on response
222. Effect of
223. Effect
of indifferent tissue V
472
472
473
474
474
475
475
COMPARATIVE ELECTRO-PHYSIOLOGY
742
226.
into
normal negative
476
after tetanisation
477
Conductivity Balance
NajCOg solution on responsivity of frog's nerve
CUSO4 solution on responsivity of frog's nerve
228. Effect of
229. Effect of
485
485
486
486
KCl
231. Effect of
responsivity
487
488
Na^COg
489
KI
491
Nal
492
492
493
493
494
495
499
....
warmth on conductivity
243. Effect of
SOI
504
505
.....
Effect of
ammonia on mechanical
morphia
response
509
^\(>
516
516
5i7
518
by
tetanisation
.......
.
521
5^4
528
5^9
53
536
539
554
556
525
Enhancement of excitability
after tetanisation
743
546
548
nerve
549
Electrotonus
.......
...
.....
561
566
567
568
569
570
shunt
...........
572
574
Normal
274.
effects of
.....
....
579
579
582
.........
Magnetic Response
277.
make
of
583
....
594
594
595
595
281. Magnetic
cessation
..........
....
..........
..........
tetanisation,
effect
01
623
608
609
duction
Enhancement
stimuli
of
610
611
612
613
COMPARATIVE ELECTRO-PHYSIOLOGY
744
PAGE
Response by
resistivity variation
Persistent after-effect
290. Effect of
warmth
hastening recovery
in
...
601
603
601
602
characteristic curves
Effect of tetanisation at
in
...
620
......
627
628
.631
Effect of tetanisation at
seen in
...
632
......
........
302.
629
powder
634
631
632
630
633
.......
......
.....
635
642
642
638
and
...
..........
649
657
658
Ophiocephahis
and response
312. Relation between stimulus and response
in plant nerve
in
magnetic substance
659
659
660
of
pleasurable
(mechanical stimulation)
........
sensation
to
painful
under
kathode
670
671
745
PAGE
due
to thermal stimulus
converted to
672
anode
E.M.F.
it
is
.
672
671,672
modified to
.
671, 672
Memory
.
319. Revival of latent impression in metal under diffuse stimulation
320. Revival of latent or ' memory image ' in phosphorescent screen under
diffuse stimulation
.........
683
684
685
INDEX
Additive
effect, 34,
595
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
in metallic plate,
683
,,
in
Biophytum, 211
,,
,,
in
Desmodium^ 212
Bernstein on
'
-1
,,
in,
59
of,
133
method of, 5
Burdon Sanderson, on response oi Dionaa, 12
,,
Characteristic curves of
5,
M
Complex
,,
,,
},
conductivity, 621
by
lag, 674,
675
by
675
COMPARATIVE ELFXTRO-PHYSIOLOGY
74^
on conductivity
,,
z^if^jwj
495
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
495
on responsive variation under alcohol, 494
on variation of conductivity under alcohol, 494
,,
,,
>>
.,
,,
,,
,,
,,
,,
,,
,,
,,
,,
by
by
by
by
by
,,
,,
,,
,,
contrasted
,,
,,
,,
variation of responsivity
,,
,,
frog's nerve,
effects
of
485
and
NaCl
NaBr, 487
Na^COg on
conductivity by
489
Conductivity balance, experiments with
on conductivity, 499
of electric current, 565-568
effect of cold
,,
,,
,,
,,
,,
,,
of warmth, 501
Correlation, 553
of,
various theories
159
decline, 158
positive
Current of
of,
156
rest, effect of
CO2
of,
of,
161
149
on, 122
,,
,,
effect of fall of
,,
,,
effect of
Na^COg
225-227
,5
,,
,,
,,
effect
,,
,,
effect
,,
5>
in animal skin,
,,
j>
in Citrus,
288
,,
)>
,,
,,
,,
224
in Dioncea, 224
in Ficus, 224
in foot of snail and
,,
in frog's eye,
418
its
variation
on
injury,
318
INDEX
Current of
rest, in
749
Nepenthe, 335
in vegetal skin,
natural,
and
its
phasic changes
298
variations, 317
in,
302, 303
126
of,
24
Darwin
in Drosera, 331
170
Death, (^\^&xQXi\. post-mortem symptoms of, 192
Death-point, accurate methods of determining, 194
Dead
on excitatory reaction
,,
,,
,,
determination
,,
,,
,,
,,
,,
,,
,,
of,
initiation of,
219
under stimulus, 212
of,
214
retinal current,
415
in eel,
285
inMusa,
114,
284
,,
,)
in retina,
,,
,,
,,
in variegated leaves,
in pulvinus, 303
419
286
Differential excitation, in
Diff"erential response,
Mimosa,
,,
,,
of
,,
,,
laws
of,
108, 303
109
,,
,,
,,
in,
329
346
339-341
in Nepenthe,
,,
in tortoise,
,,
,,
,,
,,
,,
in
,,
345
of,
,,
of,
gecko
after tetanisation,
256
reversal of,
under
fatigue,
327
346
750
COMPARATIVE ELECTRO-PHYSIOLOGY
,,
,,
Drugs, modification of
effect
significance of effect
,,
Drying,
effect of,
of,
342, 343
by tonic condition, 64
of dose, 639, 640
on nerve, 539
on current of rest in frog's skin, 288
on organ current, 260
on positive and negative polarisation, 246
Du Bois-Reymond
,,
,,
,,
,,
pre-existence theory
,,
,,
Dying
Ebbinghaus, on
of,
149
173
rate of forgetting,
678
,,
,,
in Coleus,
,,
in
,,
in
,,
in
Bryophyllum, 2$i
,,
257
248
Mimosa, 268
Musa, 284
Nymphcea, 246
,,
in pitcher oi Nepenthe,
,,
in Pothos,
,,
in
2.^(i
286
Pterospermum, 255
,,
in plant
and muscle
and dead
cylinder,
tissue,
50,
56
,,
,,
,,
laws of response
,,
,,
laws
of, in
>>
220
in isotropic organs, 75
,,
their advantage,
274
173
INDEX
Excitability, variation of,
,,
,,
formalin, 132
j>
>>
HCl, 138
KHO, 138
NaHO, 134
,,
,,
,,
Excitation, true
Fatigue,
,,
.,,
chloral, 131
chloroform, 130
,,
by
,,
,,
,,
Na,C03, 136
,,
meaning
alternating,
16
of,
98
in response of metal, 7
,,
,,
,,
in
,,
,,
,,
in
,,
,,
,,
,,
,,
,,
in
,,
,,
under overstrain, 96
,,
751
,,
of, in
stem, 558
678
of,
of,
418, 426
,,
,,
in
stomach
of,
302
345, 347
Galena,
response
of,
response in nerve
of,
,,
,,
,,
,,
in
stomach
of,
530,
657
of,
346
,,
435
,,
Growth
Gymnotus,
Haake
temperature on, 73
pulsation, 221
electrical discharge in,
242
on electromotive difference
in plants, 13
COMPARATIVE ELECTRO-PHYSIOLOGY
752
Human
response
lip,
of,
321
,,
,,
tongue, response
300
of,
322
Hydraulic response, 55
method of:
,,
,,
effect
effect of cold
,,
KiJHNE and
Kiihne on polar
Protozoa, 579
effects in
Kunchangraph, 511
Kunkel on electro-motive variation due to water movement, 13
on electric reaction in plants, 13
,,
Latent
Laws
of, in
metal, 683
75
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
excitability
,,
,,
,,
Coleus, 248
,,
,,
,,
,,
,,
,,
,,
,,
,,
Mimosa^ 268
Musa, 284
Nymphcea alba, 246
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
Fothos, 286
,,
,,
,,
,,
Pterosfermum^ 255
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
and growing
organs, 394
multiple electrical response induced by, in plant, 406, 408
multiple mechanical response induced by, 407
J,
J,
s,
of,
411
,,
,,
in retina,
426
INDEX
753
,,
Lip, response of
427-429
human, 321
Magnetic
balance, 607
Magnetic conduction, blocking
614
of,
,,
effect of
,,
,,
enhancement
,,
,,
,,
,,
of,
in,
595
,,
,,
,,
,,
effect of A-
,,
,,
,,
,,
magnetometric record
of,
,,
,,
mechanical record
593
,,
periodic,
and transient
and K-tonus on
of,
after-effect,
623
excitability, 611
594
594
uniform, 594
Makpterurus^ electrical organ of, 242
,,
,,
Mechanical and
Memory, an
after-effect of stimulus,
,,
explanation of revival
,,
persistence,
,,
spontaneous revival
Memory
677
685
of,
,,
Mimosa,
of,
no,
,,
electrical response
under
,,
hydraulic response
in,
light,
401
,,
55
hydro-positive and negative response
,,
light,
,,
of,
,,
reversal of response
, ,
by
fatigue,
of, 56,
59
400
303
326
of, 620
Morograph, 197
Morographic record by electro-motive response, 202
by mechanical response, 198
,,
M
hy resistivity variation, 546
j>
,,
Motor transformer, 281
3C
COMPARATIVE ELECTRO-PHYSIOLOGY
754
Munk
Musa,
on response in Dioncea^ 12
electrical response of, parallel to Dioncea,
Muscle cylinder,
237
150
in Biophytum, 209
in
,,
,,
>j
),
,,
in Drosera, 343
electro-tactile in Mimosa,
,,
,,
209
,,
,,
,,
,,
347
mechanical, in Biophyttim, 208
in
,,
stomach of
Desmodium^ 212
))
>,
,,
in
,,
,,
in nerve, 539
,,
,,
)>
,,
>
>)
,,
341
frog,
}i
J 5
,,
in frog's retina,
>>
J,
,,
in
,,
,,
human
426
retina,
430
of growth, 221
in,
335
,,
,,
,,
mechanical changes
failure to conduct,
Nerve of animal,
in,
509
530
>>
s>
>>
,,
j>
j>
>;
>j
))
J J
)>
>>
>j
,,
>
>)
)>
,,
>
5>
j>
,,
>
j>
jj
,,
>>
>j
>>
,,
employment of electrical
due to, 458
enhancement of normal
stimulus, errors
response after
thermal tetanisation, 462
gradual transformation from positive to
negative through diphasic, 464
>>
J,
of,
constituent twitches
in,
during tetanisation
535
>
>}
>i
,,
effect
>
>
))
,,
effect
j>
>>
,,
,,
>>
>j
>
.,
,,
INDEX
Nerve of animal, mechanical response
755
of, effect
,,
,,
,,
relation
,,
,,
,,
,,
three types
,,
,,
of,
of tetanisation, 536
Nerve of
,,
,,
,,
,,
response
of,
470
conversion of abnormal to normal after tetanisation,
475
^
,,
476
,,
,,
,,
,,
,,
effect of
,,
,,
,,
effect
,,
,,
,,
,,
,,
,,
effect of tetanisation in
,,
472
enhancing, 476
,,
,,
Nerve of
473
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
enhancement
557
after tetanisation,
647
various direct and indirect manifestations
,,
,,
Noll, Haberlandt, and Nemec, statolithic theory of, 435
Nymphaa
response in retina
of,
648
246
554
of,
of,
529
427-429
of,
Pacini, law
of, 242
and Czapek on theory of geotropism, 435
Pfliiger, avalanche theory of, 502
Pfluger's law
its failure with high E.M.F.,
579
with low E.M.F., 581
,,
,,
,,
Pfeffer
turgidity, 305
,,
,,
Phenolpthaline, detection of transport channels by, 360
Polar effects of currents, demonstration of
>>
,j
j>
,,
,,
>,
),
,,
in-
COMPARATIVE ELECTRO-PHYSIOLOGY
756
Poison, action
of,
,,
,,
unmasking
,,
of,
eel,
in,
394
289
546
its
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
,,
response, 549
experimental
difficulties of,
3,
540
600-603
of selenium by, 3
Response, bifurcated expression of, 104
,,
,,
,,
,,
,,
law of isotropic, 75
,,
positive, diphasic,
,,
and negative
in cauliflower,
in potato,
,,
,,
,,
,,
62
64
,,
,,
various forms
of,
Response recorder, 34
Retina, determination of differential excitability
excitatory after-effect in,
,,
multiple response in
,,
response
,,
,,
,,
in
Ophiocephalus, 427-429
in frog,
,,
418
Retino-motor
Wallago, 425
,,
of, in
of,
427-430
human, 430
frog, 426
,,
419
421
Retinal response, conversion of abnormal to normal, 423
explanation of abnormal, 423
,,
,,
,,
effect,
,,
427-429
responsive secretion
,,
Rosenthal on current of
of,
352
stomach, 288
rest in
Sachs on growth
Season, influence
,,
,,
of,
INDEX
Selenium, response
757
of, 3
646
,,
,,
dual elements
,,
effect on,
,,
effect on, of
,,
,,
in,
674
650
,,
,,
positive
,,
,,
Sensimeter, 670
Shoshungraph, 368
Skin, abnormal response converted to normal after tetanisation, 311
response of, in frog, 300, 302
,,
,,
,,
,,
,,
in intact
,,
,,
,,
in tortoise,
,,
'
human, 300
309
307
316
and
resting current
its
,,
,,
,,
,,
,,
,,
instantaneous mechanical, 47
,,
of,
24-27
,,
,,
,,
,,
vibratory, 27
40
Stimulator, electro-thermic, 38
vibratory, 30
,,
Stimulus, effect
of,
on growth, 73
importance
))
,,
,,
,,
,,
,,
of,
,,
,,
opposite effects
of,
i,
in plant-tissue, 40, 41
5,
in nerve of bull-frog,
,)
in nerve of fern,
659
658
tissues,
77
COMPARATIVE ELECTRO-PHYSIOLOGY
758
,,
,,
,,
,,
,,
,,
,,
J5
,,
,,
,,
,,
,,
,,
effect
384
,,
,,
>j
,,
,,
,,
Tabular
of,
375
,,
overbalance, 386
statement
,,
electro-motive variation in
,,
Desmodium
leaflet,
219
,,
-;
,,
,,
Temperature, different
effects of,
on
on conductivity and
,,
effect of cold
,,
effect
,,
of cyclic variation
effect of low,
,,
effect of,
effect
,,
excitability, 187
on
electric response,
189
190
on growth, 74
of rise of, on autonomous response, 182
effect
of
rise of,
,,
,,
,,
,,
in frog's retina,
,,
,,
,,
in
428
Tetanisation, effect of
A stage
conversion of abnormal
Electromotive response,
,,
,,
to
normal
resJ)onse,
629
,,
,,
,,
,,
,,
in
,,
,,
in tin,
in
628
Mechanical response,
,,
,,
<
conversion
from
abnormal
expansive
to
normal
INDE)^
7:59
Tetanisation, effect of
Response by
resistivity variation,
,,
,,
,,
in selenium,
,,
,,
,,
in tungsten,
629
630
stage
Electro-motive response,
,,
,,
in
,,
,,
in nerve of frog,
,,
,,
in response of tin,
462
630
Mechanical response,
,,
,,
in frog's nerve,
,,
,,
in nerve of fern,
Response by
634
554
resistivity variation,
,,
,,
in
,,
,,
in selenium,
Magnetic response,
in response of iron,
,,
,,
632
,,
,,
,,
reversal of
normal response
,,
in Drosera,
in
,,
342
stomach of gecko, 346
Mechanical response,
reversal of
in frog's nerve,
,,
,,
,,
,,
,,
,,
,,
in
,,
,,
,,
,,
,,
in
635, 642
indiarubber,
642
nerve
of
gecko, 530
Response by
resistivity variation,
,,
,,
638
,,
,,
stimulation by, 38
variator,
,,
13
Time-relations, difference
45
J 5
of,
of stomach
307
of,
345
Velocity
of transmission
determination
of,
by phenolpthaline, 36
55
>>
,,
of,
449
448
COMPARATIVE ELECTRO-PHYSlOLOGV
760
Velocity of transmission
determination
of, effect
effect of
,,
,,
Verworn on polar
effects of currents
Vibratory stimulator, 30
Vision, binocular alternation
of, 431
Visual impulse, pulsatory character, 432
Von Fleischl on response of nerve, 278
Waller
,,
,,
,,
on
on
on
on
'
Wood, demonstration
of living characteristics
of,
361
,,
effect of anaesthetics
,,
361
,,
normal response
361
Zantedesci on organ
of,
of,
362, 36
current, 269
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