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GALVANOTROPISM
i;v

f
/

IN

THE CRAYFISH.

i:.

mili.kh

l!''j>riul'''i

mill

ill.'

.lijurnai
.;.

ut

/'hifsiuiui/i/.
-j:.,

Vol.

\.W

Nm.

M.mli

I'.tUT

^L^ uAOo

yrv^^

^ft^/^-^

Vii'.

XXXV.

X(,.

',.

M,rh

l'.(07.1

GALVANOTROPISM
B\
F.

IN

THE CRAYFISH.
Universlt;/

K,

MILLER.
of Tnnmtn.)

(Fnim

the I'hijsUiloijIod LnhuniViries in the

The
when

earliest observer of tlie iiheiioiiienun of galvanotropisin


first

was Mach',
that
to

but the

detiiled account was given by

Hermann-, who found


were subjected
rest to

free-swininiin;! tadpoles or saltnon-cnibryos

the

influence

of a ,L;alvanic

current tlioy

came

with

tlieir

heads
;

^m

Subsequently he extended his earlier stuili(>s the central nervous system wrvs stimulated by an ascemling (,r honiodromic current, depressed or jiaralysed by a descending or ant idronuc current. Decapitated specimens also manifested the same peculiarities, e.(/. unrest

towards the anode.

ascending, rest in descending, currents.


-spinal

Even

pieces of

tail,

provided

they contained

an entirely similar manner. In 1S!:3 Blasius and Schweizer^ cairied out an e.xteiidjd series of observations on both vertebrates an.l invertebrates. The discovery of
in

cord,

behaved

galvanotropisui in the crayfish


to develop the

is due to these authors. They attempt view that a descending current causes anelectrotonus of the brain with consequent depression of reflexes, while an ascending'

current, on the other hand, leads to katelectrotonus of the brain with consequently increa.scd reflex-irritability. Hermann and Matthias'
criticise the position of

Riasius and Schweizer. The brain

is

certainly

not essential for the reaction, as one gets practically the same results ir. decapitated animals. Moreover it is impo.ssible lo determine the exact
physiological electrodes,
central nervous system.

many

of which undoubt.'dly exist within the subject has be.ii approached

More recently the


both
.-is

by

Loeb and

his students,
in

Loel) and .Maxwell",


'

regards the higher and lower forms. an article on different erustaoMiis, state that
:,,!.

Ma.-li,
1

/)Vi,, ;.,;;;.<, ,;)/;,;,, I, p.


r.ii),"i.

rii,,,i

In

Hi-,.,i,.r, s,lui,i>t,. .I.r lf/,;,,T

j/.ii.;.
'

XIV. p. 27.

Vjlinter't .Irch.
Ihi,l.
/'')./.

xwvu.

p.

1,"7,

Hsri.
1

xxsix.

p.

lU. Hsc,
IVil.

/,,,
//,,,/.

,,,,

|,

,,,.,

].,,j.,

i.vn. p, .TJl.

i.MU.

p.

l-.'l.

IHW.

I'll.

xxx\.

15

ft

21(i

F.
inr.liiiin->tii'ii'4th

It.

M//Jj:i!.
arc
iiir
i;-i'il
ii.'t

wlit.'u
l.'itiii'^f

I'lii'iTiits

(lie

ahDiL-lncntiniR'd stiiiiu'I'licy

.iml
,

ill

|ii

r,--iii_;
iiir;iii.>
t'l

iiilliuih'.'.if

iilivirvi.l

cNiihiiti
tlii'i i-y.

tlio

;i)i|ii

(liaiM
Ill

>

liv

an

iii:;.

iii^iH

tliough
wliiili

iiitr:r.it<'

aMilioii
lii-lh
i-

th>'

aliM\r

wiiik

lias
,,:,

I,

cm
the

canicil

cmt

I'll

the

t'iiini-.

much

ha- aUo
(if

hci-n

ihni,.

I'ictuZ'>a;

;^riat

ileal

nl'

this

has

iict'ii

[luicly

|)h_vsiii|iij,nca!

character.
]ill\sical

Sniiic

iihsciAii^
:

!ic\vi\cr,
(

li;i\c 'a

arh'iii]ilcil
'

tn
11(11

s^ivc

various
it

cx|i!anatiiiiis
lif

fur c\aiii|ilc.
;

rlL^rc n

al tci

tn explain

hy means an inter-

katiiilinrcsis
-i--,
I'll

Lneli

aiiil

lliiilnifl- hy the
aiiil

acliiiii

nf the priHhic.s of
tjiveii

eJictiiiU

while mi
I

lately
|iiiiiit

Coehii
iiilcicst

llanatt' have
I

im

ta

inn

he

of view of

|iliy^ic:il

heiiii-try.

In

\ie\v

of the

Kieat

attacliiiiLj

to

the

|iIienoiiieiioii

it

is

sfraii'^e thai so conijiarat i\( Iv litth'

work has heen done on the


hieliei'

craytisii,

which

is

lather

nnii|iie

aniiHL,'
1

the

tonus

in
if

manifesting,'

a
to

uaUaiiolrojiic

atliaclinn.
ilil

lia\e

tiieiefoie

thought

doiraMc

make

the tollnwino c

rihul imi

lo this Mihjccl.

ooiiinioii Canadian cra\ti-h, The animal The current employed was thai installed in the Astdi'iis ilarinlilis. lalioiatoi iis is diri\ed from the cily-mains and is cut down hy it means of a|i]iroiiiiatc ihcostats. l''i.r the most a current of (i volts
Mctli'iil.

used was the

WIS used, hut


line

for

some experiments

it

wa^ increased
drops of

to la volts,

Tlio

L;railations of
lO'

cm rent were

secured by a rheo-tat.
a lew
I

eon>istinL,' of

a
:

hath tiled with


into
thi-^

'iiS(.),

to

which

L^SOj weru added

dippcil
a-<

\eitically
streiiL^th

V-shaped copper ehitrodes, which could he moved Well a> hoi-izontally throiiirh the solution and the desired
t\\..

of

current

in

this

w.iy

olitaiiied.

The experiments were


Liki;

carried out in a f;lass

di,-~li

measuiini^ 20 x a x lOctii.

Blasiiis

and Schv.ci/.er the writer made use of sheet-zinc electindes, the crosssectional area of which exposed to the watei was IT.'iO sip nun. For
Very
the
larii;e

animals a
oi

l,ir;;er hilli

wa> used,

measiirin;.^

."id

x 10 x Scni,,

area

the

ekctiides
a vcta

heiiin-

measured hv means of
ihicw
.1

4^00 si|, mm. 'I'he current was accmale )'Ar.-oii\ al Lfalv.iiioineter, which
1

heam
'o
I

of

h.eht

upoii a C'llnloid
i'ctioii

scaii'.

The
IIK)

instriimeiil

was
the

graduated
It

hat

l-Miiin. ih

(.orrespoiided to a cui rent

throu^^li

of (l(H)Nf^
il

milliam|iire>

lis

re^i.-laiicc
a

was

olims.

In

expi rinieiiN

was shiintrd

ai;ainst

re.--istanie of

current traversing the wdiole circuit


'

determined.
l;l,

OT ohm, aiiii tiio The currents were

Ar,h.

I.

An. It.

u. I'liiisi..!. p.
ji.

1;

I.

I'ltiiiii'r'i

Aicli. i.xv.
f.

.'lis.

ISDT.
[i.

'

I'eruuiii's yCticlir.

d. al!;icm. /7(i/ii.(. v.

1.

Hill.).

r,ALVAX()Tl{tH'ISM IX
ostinintcd
ii)

<

'1,'A

VFISlf.

217
nun.

Hir:iinmi',^ units, g

I'ljiials

I(l()()

iiiiHiiuii|ii'ir [iir Si|.

of aroa ixjjosid.

The
current.

craytisli

when
weak

iihiOLil

in

a i!i>h .-nch as

ih.;

abuve
tii

witli

/.inc

t'loctrodt's at

ii|)j)iisiti'

ends behaves as fulluws


enrreiit
flie'

in resjiniise

tliO

constant

Witli a

nninial, if ah'eady facing the


in the ty|iical coordinate'd
cra3ti>li'.

anode

(antiihomie positi'm),

niuve.-.

forwards

manner
a strong

described by Betln' as noinial for the


It

If not in this position


befoi'e.
\\

turns to Ms->nnio
tlic

it

and then moves forwards as


in
thi;
is

ith

current

animal
if

antidromic position mov(^s


excessively strong

forwards very

rapidly, but

tlie
is

cuirent

the' lindis

becnme
is

rigid

and the
tlu'
first

ai\iinal

unable to advance.
in tin'

]f a .strong current
it

jiassed

through an animal

hniucidiHude pnsiiinn
:

moves backwards by
in this

powerful and characteristic lail-flexiuns

it

docs not

instance

assume the antidronuc [msition, but darts backwards to tlii' anode. It is true the animal sometimes goe~ to the kathode, but it is undoubted that the main tendency is to move to tin; anode, where it invariably
remains the longest and goes mo>t
readily.

Such irreguhuities arc


iii'iinent

probably liue to the intiuence of the electrolytic products of dissotnation.

When

the

lii'ad

is

towards the auude, at the


is

of closure of
:

the current, the telsnn

closed t'lLfther, but so()n si>ieads out a'^iin


clo.->ure

the cliela also undergoes a preliminarv

followed

abdomen
tliorax.

is

at the s;\me time oxtjnded.


is

When

byopenirc: the moving backwards to tlie


tlie

anode the telson

spread out and the abdomen ticxed underneath

When

using currents of truxlerate strength, one cannot obser\'e any

such definite orientations of the limb.? as

for Pahviiwnetes
is
it

possible to

Loeb and MaxwelP describe Only when the animals are expo-ed to strong currents make out anything (kf this kiinl. The appearances of
and tigwred
ojie ha.s

Addcus under
the

these circumstances liave been described

photographically by these authors.

In the antidnaidc position

abdomen

stretched out. the chelipjiis tiexed and the


jio.sition

claws firmly

closed.

In the homodromic

the

abdomen

is

tiexed, the telson

antennules bent underneath the body, and the claws stretched open. Loeb and Maxwell had noticed that the muscle which opens the claw was iiere
tiie

spread out, the antenna'

are bent

backwards,

in

a state of contraction.
tin,'

When

the current

Ls

pa.ssed
tlexi.'d <ui

tran.vcrsely

through
tile

animal one notices that the limbs are


open,

the side of

anode, extended on the side of the katho(li', the claw on the anodic
is

side

closed, that on the k.itlio(Uc side


'

is

I'jliiijei'i

Arch. Lxvm.

i'.

l.ju.

1,8'J7.

-'

Im.

cit.

i:.

f>)

2 IS
It
i^

F
.luiibtliil

n.
r..irb

MiLi.Ei;.
aii^l

wlirtliri-

M;i\wrll,
iiiti'l

allhoii'^Ii

drscribiiifj
'I'liC
t.i

(liesf

aiiji' iu-.ui''i'<,

I'larrd

lllr

(iirrct

inrlat ion

llpmi

lilclil.

Lcliavioiir of tlu'
\\\\i\v

claw in the almvr in-tancfs really ^ivcs thr key

tlio
is

laohlrni.

When
is

the

l.tM'l

is

tnwar.K the
tlie

ariml,.
is

tli.'

law

cld.-ed,
iinc
ii

when

it

towaitls the katlimlc


all

elaw
Ijc'
i

opm;

the aiipearentirely

es

|iresenleil

hy

the

tnuseles

may

\plaineil

by

ref,'ivnc<> tn

the well-known Kitter-ileliet


of

iihenorienen'.
stnilied

Hitter,
i-tVeet
i)t

at

the

bt'jiiiiiiinij;

the

nineteenth
'he

eentnry.

the

stiiiiuiatini,' tiic sciatic iiefve in

tVeir, wit],

liie

niusclcs ef tlu- lind)

iataut.
ilexoi-

He

foniid tliat with

weaker
ly

sliniiili en<' i,'ets


litidt,

contiaction

(if

the

muscles, and censei|nent

tlexien eftlie

wliile with stnmirer

stimuli the exlensef muscles eoiitiaet and

marked
results

I'xtensinM isnbtaincd.
witli

Kollctt- in IST-t veiitied these earlier means and e~lal)li~hed the |irii'ciple hat
I

mere

suitalilo

,antaL;eni>tie muscles,

such as
other,

the tlexers

and
the

exteiiseis ef

ii

limb.
It
t

|.ii~se^s,

relaluely to

ea. h

ditVerent de.;r<'es orirritabihty.

has

sin<'c

been found that the


is

iirinciple

involved

in

lii

er- l!ol let

phen.uneiKjn

but pari of a t;ieat

Thus riitzncr found that one streneth of stinuihitioii general law. apjilied to the vagus caused narrowing of the glottis, while a stronger Indeed it is very probable that wliurever stiniuhis caused opening.
(!

antagonistic groups of inu.sclcs


obtain.
It
is

e.xist

these
this

ditlerenecs

in

irritability

])racticaily
in

ceitain

that

principle

plays

very

important n'de
atfoided by
tirst to

I'robablv the best


is

the noinial t'unctiouing of these ditfeieiit structures. and most interesting example of this phenoineiioii

the

Kichet and l.,uchsiMger^ were tlie claw of the craytish. show that a weak ^tinudatiou of the nerve of the lindj led to
fi

openinj; of the claw, while

stronger one caused closure.

known
two

the opening and closing

movements

are carried out by

As is well means of
ot

ant.igonistie muscles, a small abductor


tif

and a large adductor, the


a weaker, the hitter
a
restdt attained

foruHT contracting under the intiuence


stroiu'cr, stimulus.

The

question

now

is

whether the

in aiiv case

is

not merelv

the outcome
is

ol

a struggle between the two


coiitraciioii

muscli's or whi'ther there

such an arrangement that


its arit;igoidst.

in

one muscle
'

is

a-s iciated with iidiiliition in


thi-; i>li.Mi.miciiim is ;;iv,'ii liy

This
i

(pie.-^tiou
'Jl-'i.

\i'"t\ iic.M^iut iif

OsswaM.
1S71 TO.

/'/I V;;.

'.-

.I;v;i.

i..

\\.

Islll.
-

s:i:in:.,.h.

,1.

ll'i.H.c

Jhnl. h\\. lAsi.


lss:j.
,-l

xxu. hy

C'ile,i

liy

1',

icilei

UKUi

ii

7.7.'.7i,,;,;/7,M, ,;,!.(/,.
l!r,:^l,iii. r tn-r.l.
*

/l.-rlir.
.,1.

I'.ia.

Ca,

,1

I;

inlc nioiii
Iss'J.

ii

IJ,'rlri<ji]iii.-ii,},il,-.

Hii'lol.

Villi.,,

,l,i

,iiu.-r:,\-

ilis

lu'i^. p. -'71.

l,iu'h>iii;;(

r,

I'lliKjii's

Ar,>. \\\in

). til

Iss'J.

I'itiil i>y

!'

unlui

ma

ii ii

IJicin'jihij.uuhjif.

f/

-#
GAi.v.\X(rn:i)i'is.][
h.-.s

ix cuAYFisir.
1

'21

'.I

'

bcL'U answoroil
iini.-cli'.s

by Pint to wsk
.-f]);ii-alily
;

',

uIid hn stmlii^
llii>

tin'

:;tnlurtc
it

adductor

workin'^ in

way

hi- fcniinl

po.^.^ililu

to (ibfain gciiuiiH' iiiliihiiMi-y frtVcts by appropriate striiigtbs of stimulation.


It
is

particulaily iiitorcstitig in this conn.'ctioii to


wliicli

noto

that

the strength of stiiniilation

pnxhicos contraction

in

one muscK.'
e'

produces inhibition
to act in absolute

in

its

antagoni-t.

We

liave

here

vionsly

ii

remarkable periplural

nieeiianisin, wliicli

enables the ojiposed ninscles

harmony

relatively to each other.


<iuitc
is

The anatomical basis has been worked out very fully Mangold-. He finds that each of the> claw nuiscles bv
two
to
sets of nerve fibrt;s: not only so, but all the
to possess a (hiuble

recently

have been found


impulses.

by body muscles stiidi(Ml innervation. It is but natural then


suppli<'d

assume that one

set of fibres conducts motor, the other inhibitory,

Itdiibitioii

by stimulation

of peripheral

nerves

has lieen

observed

among

other Invertebrati'S.

For instance

I'awlow-'

found

the mnscles which close the shells of molluscs to be supplied with motor
an<l inhibitory fibres.

EtTccts

Vertebrates.

somewhat analogous to the above arc to be noticed in the Thus if the He.\or-reHex c^ the knee-joint is induced by
f(j\ind to

stimulation of the foot the extensor muscles are

be inhibited'.

The

inhibition necessary for the coordination of the antagonistic muscles


is

of the crayfish claw

seen to be a peripheral jihenomenon.

Among

the W'rtebrates, on the other liaiid, it is as.sociated with, the interThe cause vention of the gray matter of the central nervous system
.

of this j)henomcnon in the craylish

is

not at

all

certain.
" in

It

may

be,

however, that the

.so-called "

interference

phenomenon

nerves plays

an important

role".

Who
mental

the influence of a constant current,


jieculiarities of
is

can doubt now that the behaviour of the crayfish, when under is not associated with such fundaits

nenro-nnisculature as

these
is

\Vhen
closed,

the

head of the crayfish

towards the anoile the claw

when

In the first case we mn.st suppose the towards the kathode it is open. stimulus passing down the peripheral nerves to be of such a cliaracter

that contraction

is

induced

in

one set of muscles, aiul inhibitii'n

in their

Tlii.s ,;ul(;,m^ XIV. p.

ir,:;.
it.

is'.i:!.
\<.

'-

rrnriDvr.s /.tsflir.f.
Vjlihjer't .Irch.

iilujein. riiijfii'l. v,

l.'r>.

I'.M.I.'i.

' *
"

xxsvn,
/'/,<

p.

ti.

Iss,',.

Sherringteii.
Il'iil.

lnl>-iir.ilir,'

Action nf thr

.V, n-./ii.-

S//.7.

m, p.

s,'*.

\%<r,.

p. .).
/;(/.;;/rr'..

I'liia.

.l/v/(.

i,\\\i. p. :tsl,

1;hiI).

2-20

F.

n.

MII.LKR.
iuhibitiun has imt been dcinDiistrattd

aiitn<4(iiiists.

As

far as 1

am

.'iwan'

in thi' ('tliiT b'.ily iniisrlfs, vft

it iiiiiluiililrdly iK-curs, for

the anatomical

basis

is

incs.'nt. as iiieniiun.d above.


..nlv
ilie

We may
ilaw

be

(juite certain tliea

tliat not

apiiearances

in

tin-

ih^iii'ml

on

tliese facts,
i^

but
the

also the entire orientation of the animal.

Wiieii the ciayli^i

nervous system ui aiiiiiiioiiMe jMsition the eiirieiit ac'ls on the centra! stren<;fth suited to sneii a way that the impulses ori-inatiui; are of ,a
causi' the (..nlraetion of

one

(,'rnuii

of muscles, thosr of the limbs, body


..'ron|i;

and
is in

telson with inhibition of the antagonistic

when the ammal


i.s

the honii>di-omic position the ..iber


tir.st

>,'roiip

of

muscles

.stiinidateil,

while the

j;roup

i.s

now

inhibited,

i!ie<lernianii' and all


111

other inv. sti^'ators

,a,L;ree

that the central


It

rv

Ills

s\-i.'ni

is

the chief point of aeiion of the L,'alvanie cnirent.

was not iiiconcoivable that nlh-N stimulation oeeu-r.-,i fi,,m the s.aisory rieently cNcludid by eii.liiiuv in the skill, but thi- po-^bilify ha- been
prohabh that the periplieial ne.ves must al>o stimiila'Mii, and ihi- n: i\ be r^a lily shown by the followinr. cave \pi nm.iit. 'I'lc elielip. d IS cut off and placed in a dl-ii of simple
Dreiici-'.
It woiil.l
s^

em

w.itcr,
th.it

thioiigh winch a eui'ivBt

i>

thiii

]ias>ed.

!'

will

bserved
is

111.'

iiUL-cles 'Mv ditbavnlly at'feelel


'I'hat
ill.'

wlan the cuir.nt


.hit
if
I

passed in

oppnMti' directions.
I.UII-, lit
,,i

appear.m.'.s are
slmwii, for

lo ilio aeti-.n of the


ina.lc in
t

..n

the

IK

rve can
s.

b.' iva.iily
111
-

tw- hoh's are


in.

fill,'

pr...\i!iial ai..

o|ih

and

phil

iniiiii ele.

insia tcl,

hmugh
that

whi.h .iirirnts
l,in.l

pa-s.'d. oh.' ..blaiiis Minilar


i'arii..l
.'ii

i.

suits.

Stmlicsoi this
tin. Is

l.av.'

aU.i

b. cii

by

I',

.d

.'

ma
iv

n ir. wli.i

...n.ial th.' >l..sui'4


III,
1

aii.l
ill.'

op.

II 111 J:

mn.-.l.'s

b,

have
II.

in .in .'Na.-lly ..|iposile

;i,

i.'^ral.U
f.t.'l.ii-,

c..n~laul
,\

cuir.lil.

lli.iiw.'

have

an.'tlier

nii.l.iiibl.'.l
,,|

ih.iU'uh

S.C.. Hilary on.', in


a. II.
.11

pr..ilu.'in^ lie
iil
..ii

..ri.'iil.ili.uis

||.

ria\li-h,
I.

Sm.'.'

th'
.

.'I

lli.'

.airi.
1

th.'

p.'npln

i:il

II.

'U.

mainlv

iii;iiii r.',M
1.1.
III'.'

.1

at

tli.
..i|.

mak.'

,'iii.|

;ik, lli.'

t.'nic C'Hi.lil h.n

,,t

tl,,,
I,,

mil-. I.,1,,.
'
I

pi...

ill.'

lilatl.iiis ..f
II.

til.

mla.'t

.immal
c

lilU-t

!..

,!,,,,

._.,,!\:,i,,-all..|i
l:,|. I\

..f
.1

th.'
,1

r\.' c.'ll,e.

of ihe
ill

ea n'4;
I'

'111.'

.hilli,

I'.iiMi,

li.i^

aiii\.

al

siiiiilai
111'

ni.l ii-h Hi
is

'.;ard

to
!
I

ti-h.
'ii

'n,,

.pi,-.!i..i
....

w
-

niiL;hl
I

lai-.il;

tiie
III.

si

imulal
..I

wln.b
ni'r\e-

iii,,|,,iil.i. .Il\

UI

Ml

li.'

e.

mill
'I'll,

n. r\..il- sysl.

mainly

h.

lil,i,,

tin

ii.i\,

i.n>'
.1,

answ.r
i>.

is

tint

uii.loubie.jlv

b.. ill

arc

'

r,ii.

/../-.../,
(/.

1.

r.i."i.

I'.iO-i.

Sii-.uiigfli.

Wiriitr
p.

Akad.
)1.

rxiv.

\<.

\'>

I'.'u"..

Hhilmphyinlvitie,
Sil-.uii'i'h.
il.

IHW>.
p, IT,
1

H'/rHcr .^A^^ niv,

hi"..

GALVAXnTll()I'IS^[ rX CRAYFISH.
Ktiinulat-(1:

221

the norvc-fibivs
1..'

uliirli

],af<s

in

a ^'oiuMal autero-ro^trrinr
i.'us aiul

(lii-ootinn iiiii-t
iis

stiniiilat.'.l

by

tl'o

incvriuent of
t.i

coliouls jiisl

wv

haw shown
'.naniier

tho

p.-iipliei-al

nnvos

ho stiinuhitul.
tliou^i;h

The

nervo-

cells are also

uu.iuestionahly stimulated,
oxiilaitio,!

probahly not in the

peculiar

by

Loob

an.l

Ma.wvull.

They had
in <.ne

assumed that when the current


direction tho e,ll-hodirs of
tie-

is passed throncrh the ueunuics suiiplying one

ai.iiual
sot

of muscles

are in katelcctrotonus.
tonus.

whde
t.)

They

also sujiiiosod

those of tluar antagonists are in anelectrothat on-.' set of nerve-elmicnts ^represent

a cnissed relationship

it is true the muscles which tbeV supi)ly. seen, the axoues of which cross that anatomically ccr'ain neurones are distribution of the-c is each other in the middl.' line', hut tlm ultimate As far a> the assumpti(Ui (if kat- and aie-i.rtroimtorionslv uncertain. which I'.ie.leriiian n tomis is cmi.vrucd it is a ].ure hypothesis, regarding indeed Loch', hims.lf, expresses hims, If as rather sceptical^ finds that the l)ehaviour a subsequent co.nmunication on gaUaimtropiMn, thi- animid is exposed to a of the skin-linds ..f AniMystoma, wii.n by suj.posiug the central .tant cnneiit, can o.dy b,- interpret. d I

%,

attempts to justify ins system in thcrv 111 regard to crustaceans by stating that the nervous may behave dilVerently, an assumption -roups of animals ditr.r'riit very general which lliedermaiin coii>id.Ms would not meet with
nervous system
to act a^ a nnif .rm Ah.ile.

He

agreement'.
Ihietly then the orientations
follows:
in

which are observed can

hi-

,.\plainedas

when the animal is an.l sends out one direetioii the central nerv.ms system is stimulated mu>.les, pe.s.s.Miig certiiin group of iiiiiMilses of such a strength that a antagonists Ijeiiig all irritabilities, is cans,.,! |,, contract, their
ade.inate

subjected to the intluenci' of tlie <-uneiit

reversed the londi'ioiis are precisely the ivous .-y>l,an is 'IdiH slimill.Uinn .M'.airj-ing in the ,a-ntr,d m ,,pro>ile. Combining n. rve-tibrcs. partly of the nerve-, 'ells and pailly ol the the p, npheial iiervis with this pimcipl,' the actmn ,.f the lairient on is al.-o char from the al)ove it explained, l|. orumt.ations are r.adily craUish is acted upon as a that the ceiitial lervous ..yslein of the no .xc.ptioii to the g,-iieral rule nmfaiu whole and tlius proves
inhibited; wli.n the current
is

l!,t/iu<.
p.

!:r,l.

rnl.rgn.h.

I.

\<.

\.

l-'.'ii.

.\\...

.\\\vu

i.hi.i,

.I-ian. Mic.

S,-i,-i,.<;

XWM

*

nil<l.

1^'tt

trifl:

I'hytixl.

I.

I'.

I'.fJ.

1'"-'
f-'.':.
I

ViVl<,fr Ar,h. i.xv.


lu'ifh.
,1.

\<

:ii's
1

riiij>:>'i.

I'

"'-

.222

F.
f..r,ns.

H.
I

MILLER.
shall

estal.lisluMl

for ..tlior

ivfvr R,l,sc,|uc.itly

to

the

real

Lfirin-

if

anv, of these ori.ntath.iis

o!i galvaiiotroi.isii).

As"'.nentio,UMl

ahove

Her.nanu' and Blasius


i,sitio,.

a,.l
.[Uiet,

Schweize.
those u the
in

antMlro.ine utic..l that ani.nals n, the


],.,,,,,,
,.,.,0 ,.xe,te,l.

were
to

Th.y

foun.l

this

apply

the

h.-h.

by the work of numenms This is a,n,,hihian a.i <Tavtish. ot worki>i- with the hu:d portions unters' l;las,us :m.l Sehwei/er, and (lejir.'ssion in the tho aseendiii.u, excitation froi;s, (ihtaineil strychnmi/.Ml fiogs were -I'he convulsions of deseendin.', eiunnt. an;,inent,d ly as. endinK. etimnts'. ih,hi,ed hv dosceiMli.,^, and to a specific actio,, on the r.,,..n.kv "has shown that t'iis is .lue he , ass.d eunents ito lio-vous svst,,,,: hv nie;,s of needles
snp,.orte,l
i

II

cent.al

their descndiii^ accvleianon of the ,-e-p,ration~, while similar ivsnlts i.c-ios and On, mils also a.rived at cessition. this general propoMli.m. r.,,.dcr,uann,liU'NMs.'. admits the tmih of moderate that hv iis,n,i,' .'urrents of ,,ri, and Maxw.dl l.avo >!atcd -h^piosio,, are not ohserved. s,,,..Mh the otVccts of ex.atalion and howover, in the eraylish, as in the other '1 l,at' thev ivallv .lo ohtam, d The ainmal is tied upon it.s ear, l.o .ho^vn as lollous.

the spinal

co,d:

ax,

ndin,,'

e,i,Tet ^

nndei- these

conditions,

can-.,
t.,

cuiTents h'd

f.,,-,,,-

studied,

i,:i,k

so that

tlio

appeiiifirres

may

l.e

liioic

ivadilv ohseived.

l)nnie,^

the pas-acc of a .le-.vnd,n^' cm-eiit cuirct they a,e in a cond,t,o,. of ,in,e' the pa-aue of an ascend,,,st.o,,;^ .u>T.nts a,v tu,ed It is true that wl.en ve,y ,K,.,| ,,,u,v. o,ientation<. hut the.e oo not either direction one notices o,ds
.,

ida"-es are at re^t, tlie ajipon.

whih

in

^..

,,|,

le,ate

e,,.nts

which

-ive
,i

ri^e

to

excitatl^^n

and

,l..Jv.~io

Other npo,-,ant

f,ct>

make

>t,ll ,o,e ,-e,tain that


1

tins

have lound In ilf cou,-e of th,s wo, k law applies in the caOi^li. vet, a! -ai,-lioic cliam i- stimulated, a> hy tl.at ulie,, anv part of the out ea,-i,e> er,upted f,rad,c curnut, the anal sphinder tl,..
rhvthnncal
,,p,t
inte,
,..

.-onl

actions.

In

suhM..|e,

communication
seen,s
t,,

shall

moie

tullv

ou

tln^

p!,e,,o,i,euon,
liea,
t

wind,

attoid
<

an
.

MIC analogy NMlh the

of lamulu^, a^ desc, ihed


'"'
'<"

hy

'arlson

U.nt.
I,,,..

'

"'
'

>

.,(. p.

y>x.
(.

*
.../.

rnr.,lb.

</.

ir,W.r;,.vi..r."".
IH"?
ln.li..l'<- U'

I-'''

7r,.,(r ,JV7,r(,-.,a^ mr.li,;,U: \'n*,


!-

,.,. ntc

cti.,n,

ntlexrs
.l.-

ft

dimmvie

rcx.itat.ilil*

ny\W."
IW-J-

CiUil

ommn, l.v BUmu.s

^ lUa-,..- >..:as.l.^v, ,. r ' - 1.' -uooU '-' '"'"'" '^' "" ''' """ '" desomlual ..u,k.I,.- V* m-iK.. o ll. vs
'

"

'-"'

.a

l.>

'

anl hcliw.

i/..r.

Er(ifb. d. I'lif'iol.

I.

p. l'<n.

Amrr. Jo'irn

V-'i-.l

n r ''

''"'"

GALVANOTHOPISM IX CRAYFISH.
If th.-n tho an.l ,]>h\uctov
is

223
impulses
the

aftivo

nv,-

havo a

inai.-ation tliat
>s

an. brin-

s,.ut

out

l.v

tho vnlrat .haiu of -an-l.a, tliat


.1

lo say,

PauMiaare
,|,

in a c.MHiilio,, f i.K.vas.
i..

ir.

Ital.iHty.

ihw
,s

tiinls

now that

\.hon a constant cunvnt

thr asr.n.lin- ,l,n-etio

p.ssr,! thron^li

,i
is

,1,,,

s,.hu,..l.ait

lK..,.n.s
:

cunvnt

passt.!

is

unatTo.t.Hl

aciivo, .hi!- wh-n a ..-sc.ndM.i: an this ran only mean that m

,,..n.lin.'

cMurent tho gan-lia are

^litauhitin^. or the ase..|.lin- current is who.,- nT,tal.,lit.v nno,lic region itself

a con^Iit.on of increase.! nntalnhty, is the It is also clear that it


is

increas.,1.

IWasn.s

an.l

Schweizor

%vcre

.,f

iintahilitv while tin- katloli..

opinion that the ano.lic rog.on w,.s i.-ion ha.l its irrital.ilitv uuavaMMl.
.1

.l..pre..so.l

m
We

s.e from the last-n>ention,


is

results, iiowovor, that exactly

ihe op,.os,te

the ca>e, while

it

is

tho at the s..ne tin>o clear that

onfnary results
.liivct
It

c.ntral norvous s^slmi. of ..Irctrotonus do not apply to thabove instances is a the action of the current in the

one
o,u>

That

,1, ,,,,tral

nerv,.us system can be readily

shown

as follows.

.,.,tions the cord

botwem

,, .,h.lniinal
,,,,^,,i,,,,

ganjia

intact,

leaving tho thorax and the abdon.on, thu^ porfornis a sones ot the spluncter tiist

,ine.or has
,;vaMls

Alter the of the cut. ,ont. actions owin,- to iho stimulus when the anode is one again tinds that come to

nM

the
is

tail

ihv.hmical
tl.o

artni.y
If

is

kathode
u-tion of

towards
leniovod

tad.

the

when the abdominal ganglia are now


elicte.l,

but

not

Miccessivelv

until only
is still

the sixth

or la^t

is

lett

the typical
,s

tho curront

manifostod, rv. activity

whon the anodo

bohin.l and rest


,.,no!i.,n is

tho la.st whon the kathode is hohii.d. When, liowcvr. nspon.so to tho coe.,tant curront is at romovod all aotivitv in
bh.ntioal phe

;.o lo.t.
tl,o

application to i.na can bo rlicited by the


;

which a curont is j.issed cord of plaiinum ohvtrodos. through in.aoV and the aimdo is ,.lacrd evon Nvhen the laM ganglion is alono
b,.l,ind
it

witii tho

kathodo

,n iront

ih.Mlimioal

..,iit,

action, aro earriod

out

liy

the ^phmctor.

Amending

cunents thus stimulate the .vntr.d


It.

,,,,oii>

>v-toin and .losrondmg doprois

and

this

bw

is

.mvii to

apply

,.,

M,, oaso of but

a -ingle ganglion.

Strango

to say this interit

,.UU" prmciplo
.

has attract,.! but li.tlo attontion.


.ion,ti,an-.o.

although

appoars
I i

to

bo of groat

Sin.v
prol,

it

,-

on.iioly distinot
,.!

the
ol

niinoiple. of oh.ot,
;,,,,.,,,11. .^lni,

nus

is

.hk

as-ooiat

It 1,

In

pro.,.

thoc.nlralnonoMssystom and al-o . ith tlio pm,1.> manilos, ,,M..nordi.b.on,ia!ionof tho Uttor. The a^o^.
:,nlvad:.uat;onlromel,rtro,onu.
,,
1

anlo.ont

.Uo Iron,
th.

I'ibigo, s
i..t

law

Inav

|,,^,,

olh.i

m.lano,^

uhioli

,n,odo and

thoka-hodo

224
appears \n
l)c

F.

li.

MILLKR.
Tims Lorl)' f.mid
Ilrcucrlias
I

the stimulating elortrodo.

tliat

in

Auiblystoiiia the skin-;j;lainls an the anodic side hi'ranu^ activr dwiir,' to


stimnlatinii

nf

tlu'

("cntral
Dii
(if

lu'rvuiis

system.

dlita'.iird

lateral fiexinn in the tish

the side towards the a!i"de. the eraytish at the


the

have studied
nf the

the

tle.\i<.ns

ot'

the

tail

make and break


ceiitrartinii

current,
stroiiLjest

anil

have
all.

found
'I'he

.amdie

chi>iiiL;'

to

In'

the

of

k.iihodie

iiiieiiin;.,'

eontraet ion,

tliout^di

weaker
IJreuer'
l.icts

than

this, is

-tron:.'er th.au

the kathodii,- elo^inu'


in
tisli.

c^ait lact i.m.

has eht.ained analoLjotis results

'Jdte

eNplaiialion of these
at

and their

he:,

ring on

I'tiiiger's

law

is

not

aU char and fuither

investieat ion ajiiie.ars most

desir.dile.

In an attemi't to .analyse the -al\-;inotro|iic behaviour of the caaytisli


it

a|i|ieaied

de-ir,able

f..

di

termine the inllueliee of the

li;aiu

in

tln.s

r<'g.ard.

Ileimanii*

^iinl

Xage]
worthy.

h.id experimentecl w ith decapitated


is

animals, but the injuiy by this procedure

so gieat

tiiat
tin.'

the results
(i'-"phage,al

(atmot be Considered

tru.-t

In the writer's work

Commissures

W( re cait alter the

route bi'ing -rleetid

manner described by llethe'.ihe doisal by piefeieece. Only large specimens were (diesi.ai
and
resist
ii.s

as ihe\,are both e.isiu to manit'ulati


.smaller ones.

shock nincli better than


;

The

anin.ial

is

seeiired on
ttii'

viaitral stiiliice to a boaiii

one removes the cirapaee behind


the anteiior gastiic
sni]>|ied

rostrum, then goes down,


lie

(uittillg

muscle--;

the cominis,ur(.'s which

deeply aie

with

p.ur of tine scissors.

The
.

(peiun.'

i^

then cai'efully

sealed by
Altta-

means of a mi.xtuie ol' bee.- a.\ and t.illow. some hoiiis tin' ai.ini;il has lecoM'ied from the sIiock .'ind then behaves lU the niahmr diseiibid by l!ethe\ 'I'lnie is the t\pi''al
ab-ence
ot'

sponlnieons
of the

mo\emenls, the amm.d only going foiw.'uds


lor but
h g^.

when
nsn.d

stiinulatid and iIhIi


(

riiinatioii
,ds

When

-hmt disianre, with, however, tiie in.t moving lorwauls the legs

and pleop,
forwaid

perform pendnloii- mo'ienniits.


..f

As

is

well

known the
tor

movements
of

such anim.ds depend on the integrity of the

subo'sophage.'il

eanL;li"n,
pio^'ri;-'-iiin.

which acts a> a coordinating

C(iitie

the

movements
aiiim.'d
1

I'nder no eiieum-'ianees wb.itever does the

\er

^'.i

b.'irLw aids.
it

No
hi',

e\planalion nf

thi-.

Litter peeiiliaiily

is

attempteil by I'.ethe.

may

howi.'Ver, that the

absence of backward

'

;://;/;;.'/

.(o/i. i.w. p.
il.

.'Ids.

1S!)7.

Sit.iiii'i^ti.

Uiriifi-

ikiiil.

rxiv. p. 30,

mO.).

litr. ril.

'
'

IliiJ. p. 4(t,
/Vl--../.)'-.
/.,

.In/i.

1.1.

(...
i>.

IMi-i,

'I

;/)(/.

LXMli.

p. -IV.i

ls;i7.

.((. p, 4lic.

GALVAXOTROPIF^M IN CRAYFISH.
loconi.ition
tlu'
is

22..

due

t:o

tin/

Idiicritiidin;)!

ginnt-fibrcs,

which run through


of a constant

nil ire cnrd,


H(i\v diirs

having

bi^fri

poetinnod,

such
the

nil aiiinial

hrliavp under
as
niisi^ht

tlio irdiucTice
it

current

'

In

first

place

be expceted

never darts

backward

to the anode.

But one kind of moveinenl now obtains and


it

that one forwards to the anode;


aniTnal tnav advance
at other times
it

ne\er

fjoes

to the kathode.

The

to

tiie

anodi' immediately on closure of tlie cuiTcnt,


in

appears to be

a condition of what
is

Blasius and
does not

Schweizer have
forwaids until

called i^'aKanonarcosis, that


is v<

to say,
is

it

move

the current

broken.
ry

This

jiarticularly

the case

when one
it.

coinnieiices with a

weak current and r;raduany increases

have here ]irobably to do with the paralysing intiuence of a There is thus the detimti attraction to tlie anode d.'.scending current. It should be observed, however, just as occurs ui thi' intact animal.
while the noimal ainm.il issomcwl:at capricious in its beh.ivioiir, occasionallv goinL; to the k.athode, as mentioned above, this is never
tliat

We

noticed after the cotnniissures

,-ue cut.

Tlie behavioui' of the

appendages

does not differ in any essential respect from liiat in the intact animal. Whin the anode is tow.irds the head the legs and pleopods are
inactive,

while

when

the

anode

is

tow,.-ls

th(.'

t;iil

there

is

great

excitement manifested by all Ag.ain we see the untenabdity of the position of Hlasiusand Schweizer,
for the br.am,

the append.iges as well as by the anus.

which

]ilays -o iniport.mt a pait in their theory, is clearly,


i..n
,,{'

ininecessarv for the product


de,'<erves

the typical reactions.

One more

point

mention and that


:

is

the behaviour of the nut eiinaB and

antemndes
eij. if

in

such an animal
is in frt^it

these invariably point towards the anode,


if it

the

anode
sides.

they point forwaids, while

is

behiml the jinteiinules

are bent

undnneath the thorax and the antenna' l"iit around to the These reactions are aNo shown by the normal animal but to iv
it

that they are inliduted in the is highlv probable It thus impul>cs which ja-s from bel.nid forwards. intact anim:\l bv g.dvaiiotr'ipii- re^Ji^nscs of the er.n hsh .-ire inl imalely appears that the

minor degree;

incorporatid

in

its

maki^ up .and are a most fniidanicntal

propeity of

the creatur.-.
It
is is

clear that the

fundamental eau^-ative
fmin

fa.'tor of galv.-inotropism
I,

to

be found

in thi' licculMiil ies of

the stimulus.
sl.nidpoint but
I

o id)
,i t

and I'udgett'

had given an
lattdv ihruvMi

I'spl.aiiation

liii^

S|

kew

sch

has

doubt upon

its v.ili ::ly.

have tested the very pl.uisible \W'

'

loc. cit.

I'fiHiiiii'- /N.'/.r /.

./.

filli.'iii.

riiijii..!.

V. p. ^>2'.K

22(1

F.

n.

MILLEll.
in
tli.'

thcMiiT of (^M'hii

ami iKinatt.
iiMililirati.Jiis

iiviitiou.Ml
it

iiitr >iliictioii. an.l

tlml

that wilh

iiiiiMir

a|ip'it's to
wliii'li,

tho crayti-h.
is

'I'luir

tluory

was

]iio]io>c.i

tor

I'lirdiinn imii

as

w.H known,
c\|iIanation
slii^litly

nianitr-ts a
is

katiioilic i^alvanntroiiisiii,

Slatcil liiirllv
is

llicii

as follows,

thr ni>'nilirano of the animal


f.ir

snpposiMl to hr

inorc iiurniraljle

CI than

for

Na

ions.

]f,

now, tho animal


is

is ltroiiol,t,

into a nir.iiinii
itsrlt

liko

tap-wator the concuntraiion of wliirh


(lifTiision

loss

than that of

a
iio

slii^'ht

of Cl

ions will

n.vcssarily orcnr.
i<.ns,

The

re~ult

will

that the animal will contain an rxeoss of N.c


chai-i,'!!
'<(
it

or will hear a positive to

electrieity.
is

On now
pull,
it

siihjeetinu;
to
in

the animal
kathoili',

a ^'alvanic

current

detlnit. ly

attraeleij

the
this

and

the

cilia

responding;
test of the

to

the

moves

direction.

crucial
ot

hvpothesis con-i>ts in

iiua-ea-^in.;

the concentiation

the

medium
should

in which the animals are o!,>erved.


in,
giviii','

In this c.i-e the Cl ions


it

outside should pass

the

animal a ne,'atl\e cliaroe and

hence niove

to

the aiiode.
.-re

Indeed

this

is

f,,und aitually

to

occur: when I'nnniKiciit

plac'cd in

t<

NaCl

solutieii,

within the

oouiso of iwtior three ndnutes, durin:,' which time ditl'iisioii is takin<; 'uove Tilace, tho Lrah allot ropie reS|ionse is seen to be reversed and they
t,,

the anode.

The

position of

Coehn

.and

iJairatt

lia^

been

lately

criticised, but, in

my
the

opinion, not w.akeiied by liancrott'.

In

appKiue iho
in

tiie.iry of

Coehn and
.if

liarratt to tie crayti-li certain


latter

peculiaiities
<'on~;doralion.

behaviour
loneiiou
ihi--

the

are

to

be
to

taken

into

.\s .already
it >

slated this form

moves iml

the kathode

but
In

the anode;

older to exphiin

i> just the rever.se of that of I'n f.i iinrrtii i. anodic Ljalvanotropism one nin^t su|ijM.se the

animal
:ire

to carrv a ici^ative chari,'e

of

I'lect ricity
i

en. tluit

its

meiidiraiies
nii,i,'r,Ue
it

nc'iv peimeablo lor

Xa than
i

for CI

-ii-

the

Xa

ioiis

thus

,iut,

leavino an

\'

o<s of

'I

ion>.

In order to produce reversal

will

b-

n.i'e,^>arv to pl.ie..
is

ih.. .miuial in a -dt solution, ihe o.Miiotic pressure


..f

of which
USUI.
ilic'

great.

llian that
ih.'
,

this

-,iiu.> salt
.1.

within

tli.'

animal.

Th.'
l.i

pr.'ssur.'
..f
.1

.it

.lavtish has b.cn

I.

rmin.'.l by Krtoleric.|I.,

P,|,,al

tint

-'l

NaCI
O-^
( ".

s^ilnti.m.

.'..i

iv-p.mdini;

a d. pr. ssion of
,,f

ih.' oe.'/in- I'oint


11, at ol
.all

..f

This

c.

iilraii.m r.'pr. -.iits,


It

c..iir.M>,
t.) t..

th.-

-alts .oiilain. d in
in
'1
I

ill.'

animal.

w.i-

toiiml p.i-sih!.'

brinu' ah.iul

i.v.rsd

h.'

ciaytisli
:!lt2
|..ii

by

sir. iil;I lis

varyin.L; I'r.un
i..f

.".

Xat'l.

-J

Xai

.'.pials

X.al'l

an.l

thus

.oii.M.h i.ibly

iiioh.r than ih.' total coiiceii' lal


1

..t

the ti>Mies
.\No
is.is.

tic animal, whi!.'

r/r. ..<.,;./''..;%</.;.
luoi.
ill

III.

p.

-J

I.
]..

i'""'s:)i.

tiiis Ju.i/-n.if,

wxn.

p.

iii.

r.Mii-,.

r.i.iii/. /.Vv. .'. /;</./. .\xxv.

(iAI.VAXOTIlOl'ISM IX CUAVFIsIf.

X
(

.')

N'aCl, wliicli
iif

((jiiiils

li7" NaCl,

is

somculi

it

It'.-s

tlitui

tliis

tliis

t'Diici-iil I'atinii

Nil

i.s,

liDUcvcf, CL'itaiiily liigln-r than

tliat

uf the

Na
tile

mtaiiici

ill

the tissuis.

The ncecs-aiy

Cijiiilitiniis

fnr

obtaiiiiiii,'

rt'Vftval arc theret'die

fiiitillcil.

Seiecteil pi'otijcols arc given below.

The
fiiilows.

])iMC'eiliirc

in

inakirii;

these experinient.s on reversal

was as
of,

Only

fVe>hly-eiill('rtcil,

healthy animals wtn.' nir.de use


k^jit hiiig in

as

it

was found

tliat

those which

wc'l'e

the lahoratniy becanio


'J'ho

]e>s sensitive

and showed more


liist

irrenularitie.s

than normal ones.


in

animals wore

tested in tap-water to see wiu'ther their respon.scs


])lapi.>d

Were normal.

Tluw were then


once
testi'd in

each

a separate jar of

.l

Xa(,'l

and
this

at

this solution.

Under
the

these conditions they

are

still

anndic.

control animal was ke])t in taji-water in every case


in

and
the

was tested
'I'he

")

XaCl

specimens.
ethei'

rensoik for testin<^ the control in the


is

animals

obvious, as

same time as the other same solution as the conductivity and consiv|ueiit
at

stimuiatiiiLf actinu
I

of the

lia\e cai'iied out

medium are thus the same. I may many nmre e-\periments than those iieie

say that
I'ejiorted

.,.

and am convinced that a reversal can alnmst invariably be obtained. Very rarely I have failed, anil in these cases the failure was due to the ]ioor condiiion of the animals useil. I believe that in fresh, healtliy specimens the revei'sal will occur in eveiy instance. DitVereiit stroiigtlis ot'ciiiieiit were used as measured by the galvanometer. From the .above results it is clear that the law established by Ciielin and llarra", t in the case of I'dniniirc/iiiii hoMs also in tiic
craytish.
fcirms.

We

notice,

however, certain
to be bri^ught

For the

reversa.l

di(TereiiCes< between the two about several hours, instead of

minuies, are leipiiied. with certainty


;

impernu^aliility of the
si 'te

One may attribute this slowness to the greater membranes of the ciaytisii. It is not possible to thr<iiigli whi'. h membranes tlie migration of ions
net improbable that part of the exchange occurs

takes place

but
gills.

it

is

can si/e in this way how the process would be Frederieq' also is of this opinion. The positions of the limbs .and abdomen constituting the orientations are found to be unaltered after the reveisal. These arc therefore not

through the

One

iiecessarilv a slow one.

the cause of the riMction, but

lUiiely

represent a parallel series of

phenomena, depending as pointed out above partly on stimulation of the central iieivoiis system and partly a'. on stimulation of the
>

|iirijih(

r.il

nerves,

.Moreover

tin- ri'lative

iri'itabilities of It

the muscles,

on which these
nervous sv^tem,
sion
in

flepeiid, are founil

to be uiichangeil.

must

also be
ceiiti-al

allirmed that while the action of the constant current on the


in proihicing stiniul.ation
in

the ascending,
lii^t
i.-..
i

and depresis

the descendiuL', direeiion as ohservcil


'

bv
mil

Hermann,

i:,ni.

il.'

i\t,,i<l. I;,,, I.

ii,

/;,/./.

\xxv.'|i

22s

F.

U.

MIIJ.KR.

J! .9

<

<!

s;

l5

^1

-M

Tl

^ ^ O

-I'

-T*

C OO-K
c s c.i
-^

OOo

cc2S=
12 ::

H3ass
?,

3 =
,,

^ - s s

2 ? i

Ji

1^

^-^

o ^5

If

~~

y,

HALVANIlTRtil'ISM IX CRAYFISH.
utlficifiit
iiiiUiiiii,r
ti.

o.jr

Icail
,,,,

mrcl l.'UIIC.lllv
iiiiluc,. (Mlllliit
;,

I.

iliiri
1

iivp iiiifiitatiiinIS ll Micrr


\

IS

in these the

ppy

,,,

ihlhiii,. Mil
(Iniil)l.
,.r

a lot
I

I.

|)i L'

attiacli

Tliat sucli

exist.s

llin\ cviT,
iiiiil,
1

i)r

jy aiiyoiir

wlio ha.s
:

uatohrd
niatl iiatirr
alio,! e,

th.:

ciavfish

tiu-

iiitiiicticf

thr

coiisfaiit. curr-iif
it

no

Us

wli.-it

attitiid.. at
liy

th..

mnnient of cI.^muv,
f'nci..
,i,

is

,lrawn to the

as

It

wcri',

an

in\i>il)h.

The
<

attraction of the animal thus

pen. is n| ion th nair\ tile

iiii;

of

an
the

h-elrir'a! el lai-'fe

and

tins

iii

turn

\>

ass,i,itril

with a

pi'eiili.nitv

It '"^''il>" lH'iiial.lc :lil\ Iliat in timse forms not ..xhihitai,vattra<'tion,,i. uav or ih,. other the .inhranes are iinpenneable. As ,s well known the idoosts are not attra.^ted to either the anode or th.. katho.io, and iiere ue uu^ai assume the

iPllt

of

it i

nii'iiiliianc

which

(io

iK'Hiliranes to
.1

1,,.

nnponncahlr.

view.

]]y eryoseopie drti^rininaiion

Other faet.s l,.nd probability to scl. Dot taz/i' has found the concen-

tration of the l)h,od of marine teleosts to be oncdialf that of sea-water (iarrey-' has found that their oMu.,tie p,es>nre r,.,nnot be altered by phicmg tiiein in more dilute ,.r mo icentrued medi.. unless
,

membranes
integument
.x|ilanati,.n
f.irms.

the

are previously injure.!,

He

tie ivfore roiielu,i...<

that their

and
ot

.;ills are imp,.inieabl,. the absi^nce of galvan.-tr.

Hen;il..

th.^n we have the attiactmn amon.- these "

StMM.\1;V.
orientations of the crayHdi, wh,.,, nn.ler constant current, are jiroiluced by tiie combined
1.

The

tile inl!u,.n,v

of the

the central nervous system and of the Jieripheral nerves. The impulses sent .M>t are of such a strength that one group .,f muscl,..s is caused to contract. ^^ lule the antag.mislic group is at tli.' same time inliibited
1.
111.,'

etreet of stimulation of

as.riulnig

i.crvo,,- ..ysteni,

ciirnait l.^ads to stinndati.m .,f the central while the .iescending current caus,.s .lep.vssion of its
.,f

irntabiltty.
:!.

The niovem..nt

the animal

'll''

an.,d,. ,s preserve.1 after secti.m


-+

f,,rwar<Is, but not backwards of the u>..phag..al c.mimissures

'

to

The

the.iry
d,.p,.n.!s

of

Vovhn and Barratt


(lie

Jammr,a
.-..

that

the

attraction

of

mollifications in the case of


It
IS

on their carrying ele.trical charges applies with


ciavfish.

probabh

Mich forms as

imp^rm.vd.ility .,f the membranes of tehosts det.rtnines tlie abs.ai.-e of any gaivaiiotropio

that

tlie

attraction ain.mg tliem.

K.:s-:.;;i,;;;;!;:';:'~
'

- i--~:i. '''"-.::;:;::"::; ;i;r;i, \:Ti


lll.
l:,.,i,
,/,

li..tta/vi.
/.I../.

.(,,/..
..,

/,,/.

xvviir, p.
/.

i;i.

ls;(7
|..

l:H.

M..r.

/,,,.

ir

///;. ,,11.

-ii;.;,

l;i.-,.