1058
PHYSIOLOGY:
TWENTE
AND
TWENTE
PROC.
N.
A.
S.
The
au ho
wishes
o
hank
M .
Robe
A.
B ookshi e
o
his
excellen
assis ance
du ing
he
cou se
o
his
wo k,
and
H.
Rickenbe g
and
G.
Williams
o
aluable
sugges ions.
*
Suppo ed
by
g an s
G
24052
and
GE
3612
om
he
Na ional
Science
Founda ion.
1
Beauchesne,
G.,
M.
Leboeu ,
and
R.
Gou a el,
in
Regula eu s
Na u els
de
la
C oissance
Veg ale
(Pa is:
Cen e
Na ional
de
la
Reche che
Scien i ique,
1964),
p.
119.
2
Le ham,
D.
A.,
in
Regula eu s
Na u els
de
la
C oissance
Vege ale
(Pa is:
Cen e
Na ional
de
la
Reche che
Scien i ique,
1964),
p.
109;
and
e .
6
(below).
3Mille ,
C.
O.,
hese
PROCEEDINGS,
47,
170
(1961);
and
e s.
13
and
16
(below).
4Le ham,
D.
S.,
and
C.
0.
Mille ,
Plan
Cell
Physiol.,
in
p ess.
6
Le ham,
D.
S.,
J.
S.
Shannon,
and
I.
R.
McDonald,
P oc.
Chem.
Soc.,
1964,
230.
6
Le ham,
D.
S.,
Li e
Sci.,
2,
569
(1963).
7
Ke o d,
K.
P.,
Science,
142,
1495
(1963).
8
Mille ,
C.
O.,
in
Mode n
Me hods
o
Plan
Analysis
(Be lin:
Sp inge -Ve lag,
1963),
ol.
6,
p.
194.
9
McCalla,
D.
R.,
D.
J.
Moo e,
and
D.
Osbo ne,
Biochim. Biophys.
Ac a,
55,
522
(1962).
10
Hu lbe ,
R.
B.,
H.
Schmi z,
A.
F.
B umm,
and
V.
R.
Po e ,
J.
Biol.
Chem.,
209,
23
(1954).
11
Jacobson,
K.
B.,
Science,
138,
515
(1962).
12
Khym,
J.
X.,
and
W.
E.
Cohn,
J.
Am.
Chem.
Soc.,
76,
1818
(1954).
13
Mille ,
C.
O.,
Plan
Physiol.,
37,
xxx
(1962).
14
Loe ie ,
J.
E.,
and
J.
Van
O e beek,
in
Regula eu s
Na u els
de
la
C oissance
V~gg ale
(Pa is:
Cen e
Na ional
de
la
Reche che
Scien i ique,
1964),
p.
77.
15
Fox,
J.
E.,
Plan
Physiol.,
39,
xxxi
(1964).
16
Mille ,
C.
O.,
and
F.
H.
Wi ham,
in
Rlgula eu s
Na u els
de
la
C oissance
V~gg ale
(Pa is:
Cen e
Na ional
de
la
Reche che
Scien i ique,
1964),
p.
I
(e a um).
REGULATION
OF
HIBERNATING
PERIODS
BY
TEMPERATURE*
BY
JOHN
W.
TWENTE
AND
JANET
A.
TWENTE
UNIVERSITY
OF
UTAH,
SALT
LAKE
CITY
Communica ed
by
Hen y
Ey ing,
Augus
5,
1965
Hibe na ion
o
golden-man led
g ound
squi els
(Ci ellus
la e alis)
is
cha ac e ized
by
pe iods
o
do mancy
which
a e
in e up ed
a
in e als
by
a ousal
wi h
a
e u n
o
he
homeo he mic
s a e.1
We
ha e
shown2
ha
he
du a ion
o
hibe na ing
pe iods
o
his
species
is
ela ed
o
he
body
empe a u e
achie ed;
hibe na ing
pe iods
a
40C
a e aged
app oxima ely
wice
as
long
as
hose
a
11
'C.
These
obse a ions
sugges ed
ha
he
ollowing
should
be
in es iga ed:
o e
wha
empe a u e
ange
is
hibe na ion
possible;
is
he
empe a u e
dependency
o
he
du a ion
o
hibe na ing
pe iods
con inuous
o e
he
possible
ange;
and
i
he
empe a u e
dependency
is
con inuous,
wha
unc ion
does
he
ime/ empe a-
u e
ela ionship
bes
app oxima e.
Me hods.-Adul
golden-man led
g ound
squi els
o
bo h
sexes
we e
caged
singly
and
main-
ained
undis u bed
in
wo
da k,
ela i ely
soundp oo ,
cons an - empe a u e
ooms.
Maximum
empe a u e
luc ua ions
a
he
speci ic
places
in
cages
whe e
hibe na ion
occu ed
( he
mic o-
en i onmen al
empe a u e)
we e
less
han
±0.50C.
Thi y-gauge,
i on-cons an an,
Te lon-coa ed
he mocouples
(±0.050C)
we e
implan ed
sub-
de mally
1-2
mon hs
p io
o
expe imen a ion
wi h
he
sensing
elemen
ied
o
a
lowe
ib
in
he
mid en al
egion.2
When
g ound
squi els
we e
cu led
in
he
hibe na ing
pos u e,
empe a u es
eco ded
om
he
ib
a ea
we e
no
di e en
om
hea
and
li e
(co e)
empe a u es
as
eco ded
by
a
hypode mic
he mocouple
p obe.
Tempe a u e
eco ds
we e
p in ed
a
6-min
in e als
on
VOL.
54,
1965
PHYSIOLOGY:
TWENTE
AND
TWENTE
1059
ime-s amped
cha
pape
by
wo
Leeds
and
No h up
Speedomax
G
16-poin
eco de s
calib a ed
om
0
o
50'C
(
±0.080C).
Con inuous
eco dings
o
body
empe a u e
we e
measu ed
du ing
wo
hibe na ing
seasons
a
11
di e en
empe a u e
le els
anging
om
2
o
250C.
Expe imen s
in ol ing
hibe na ion
a
mic oen i onmen al
empe a u es
abo e
250C
we e
no
a emp ed.
Di e en
g oups
o
g ound
squi els
we e
u ilized
o
he
wo
hibe na ing
seasons
and
o
each
empe a u e
le el,
he eby
allowing
o
he
de e mina ion
o
a ia ion
wi hin
he
labo a o y
popula ion.
The
du a ion
o
he
hibe na ing
pe iod
was
measu ed
in
hou s
om
he
ime
he
body
em-
pe a u e
eached
20C
abo e
he
mic oen i oninen al
empe a u e
du ing
en ance
in o
hibe na ion
un il
he
ini ia ion
o
a ousal.
The mocouples
we e
placed
in
he
nes
benea h
g ound
squi els
wi h
su gically
implan ed
he mocouples
and
con ol
g ound
squi els.
The
du a ions
o
he
hibe na ing
pe iods
o
a
empe a u e
le el
did
no
di e
be ween
he
wo
g oups,
he eby
indica -
ing
ha
no mal
hibe na ing
beha io
was
no
al e ed
by
he
su gical
p ocedu es
o
he
p esence
o
pe manen ly
implan ed
he mocouples.
The
a e age
du a ion
o
he
hibe na ing
pe iod
o
each
g ound
squi el
a
a
empe a u e
le el
was
calcula ed
om
eco dings
o
wo
o
six
successi e
hibe na ing
pe iods.
The
a e age
du a ion
o
he
hibe na ing
pe iod
o
each
g oup
a
a
empe a u e
le el
was
calcula ed
om
indi idual
a e ages.
300
200
150
100
50
3
0
o
F-129
_
£
:
F-501
20-
*
O he
Squi els
I0
2
4
6
8
10
12
19
25
30
36
Tempe a u e
(c)
FIG.
1.-The
a e age
du a ions
o
hibe na ing
pe iods
o
indi idual
g ound
squi els
a e
plo ed
agains
he
body
empe a u es.
A e ages
o
he
popula ion
sampled
a
each
empe a u e
le el
a e
indica ed
by
lines.
Pa e ns
o
hibe na ing
beha io
o
h ee
emales
(F-94,
F-501,
and
F-129)
illus a e
long-pe iod,
a e age-pe iod,
and
sho -pe iod
hibe na o s.
The
da a
epo ed
in
Figu e
1
ega ding
he
du a ion
o
hibe na ing
pe iods
we e
ob ained
om
74
undis u bed
g ound
squi els
wi h
su gically
implan ed
he mocouples
which
had
exhibi ed
all
es ablished
pa e n
o
win e
hibe na ing
beha io .
Tempe a u e
eco ds
ob ained
om
he
he mocouples
o
h ee
ac i e,
nonhibe na ing
g ound
squi els
main ained
a
250C
showed
pe iods
o
7-11
h
daily
a
which
ime
he
body
empe a u e
ell
om
he
mean
empe a u e
o
ac i i y
(app oxima ely
380C)
o
360C.
These
eco ds
ep e-
sen
pe iods
o
quiescence
and
may
be
indica i e
o
sleep.
Resul s
and
Discussion.-The
du a ions
o
hibe na ing
pe iods
o
indi iduals
a
each
empe a u e
le el
we e
ela i ely
consis en ;
successi e
pe iods
we e
usually
close
o
he
a e age
du a ion
a
each
empe a u e
le el.
Indi idual
a ia ion
seldom
exceeded
20
pe
cen .
Di e ences
be ween
indi iduals,
howe e ,
some-
imes
exceeded
100
pe
cen
( e .
2
and
Fig.
1).
No
sexual
di e ences
we e
demon-
s a ed
in
ega d
o
he
du a ion
o
hibe na ing
pe iods
o
undis u bed
indi iduals.
Gene ally,
g ound
squi els
which
hibe na ed
longe
han
he
popula ion
a e age
o
a
empe a u e
le el
beha ed
simila ly
a
all
empe a u es
s udied
as
illus a ed
1060
PHYSIOLOGY:
TWENTE
AND
TWENTE
PROC.
N.
A.
S.
by
Female
94
(F-94,
Fig.
1).
A e age-pe iod
hibe na o s
(F-501)
ended
o
be
nea
he
popula ion
a e age
a
di e en
empe a u es,
and
sho -pe iod
hibe na o s
(F-129)
we e
usually
below
he
popula ion
a e age.
A
line
be ween
2
and
250C
was
i ed
o
he
popula ion
a e ages
(Fig.
1)
by
he
leas -squa es
analysis
(log
hou s
=
2.425
-0.0405
body
empe a u e).
The
di e -
ence
be ween
he
a e age
du a ion
o
he
popula ion
and
he
compu ed
a e age
o
each
empe a u e
le el
was
less
han
1
7
pe
cen
o
he
expec ed
du a ion
a
each
poin
excep
o
7°C
(-
16%)
and
130C
(+12%).
These
discon inui ies
in
he
linea
ela ionship
can
be
explained
on
he
basis
ha
he
g oup
ep esen ed
a
70C
had
a
p eponde ance
o
sho -pe iod
hibe na o s,
whe eas
he
g oup
a
130C
consis ed
p ima ily
o
long-pe iod
hibe na o s
as
de e mined
by
he
pa e ns
o
hibe na ing
beha io
exhibi ed
by
hese
indi iduals
a
o he
empe a u es.
The
da a
ob ained
a
a
body
empe a u e
o
360C
which
we e
in e p e ed
as
possibly
ep esen ing
sleep
i
he
ex apola ion
o
his
line.
.051
o
.01
-
.005
325
335
345
355
365
IO5/K
36
25
19
13
9
2
C
Tempe a u es
FIG.
2.-The
ecip ocals
o
he
a e age
du a ions
o
he
hibe na ing
pe iods
o
each
popu-
la ion
a e
plo ed
agains
he
empe a u es
exp essed
as
10/0K.
Co esponding
empe a-
u e
le els
a e
indica ed
in
'C.
We
ha e
assumed
ha
he
a e
o
he
p ocess
limi ing
he
du a ion
o
hibe na ing
pe iods
is
p opo ional
o
he
ecip ocal
o
he
du a ion
o
he
hibe na ing
pe iod
in
o de
o
cons uc
a
g aph
which
is
equi alen
o
an
A henius
plo .
This
g aph
(Fig.
2)
shows
he
ela ionship
be ween
he
log
o
he
ecip ocal
o
he
a e age
du a-
ion
o
he
hibe na ing
pe iod
and
he
ecip ocal
o
he
empe a u e
in
OK.
The
poin s
on
his
g aph
be ween
2
and
190C
appea ed,
by
inspec ion,
o
ep esen
a
s aigh -line
ela ionship.
A
line
bes
i ing
he
da a
be ween
hese
empe a u es
was
calcula ed
by
he
leas -squa es
analysis.
The
da a
ob ained
om
he
ela i ely
small
numbe
o
animals
a
25
and
360C
i
his
line.
The
appa en
ene gy
o
ac i-
a ion3
ep esen ed
by
he
slope
o
his
line
has
a
alue
o
15.5
kcal/mol.
Because
o
he
linea i y
o
he
da a
plo ed
in
Figu es
1
and
2,
i is
emp ing
o
assume
ha
he
du a ion
o
he
hibe na ing
pe iod
may
be
limi ed
by
a
single
egu-
la ing
p ocess
which
ope a es
simila ly
a
all
empe a u es
be ween
2
and
250C.
Since
he
da a
ob ained
a
he
sligh ly
dep essed
body
empe a u e
o
360C
i
ex apola ions
o
he
lines
in
Figu es
1
and
2,
i
is
sugges ed
ha
a
ela ionship
be-
ween
he
du a ion
o
hibe na ing
pe iods
and
he
du a ion
o
sleep
may
exis .
VOL.
54,
1965
PHYSIOLOGY:
LORENTE
DE
N(6
AND
HONRUBIA
1061
Summa y.-The
du a ion
o
hibe na ing
pe iods
o
golden-man led
g ound
squi -
els
was
measu ed
a
body
empe a u es
anging
om
2
o
250C.
The
log
o
he
du a ion
o
hibe na ing
pe iods
was
shown
o
be
a
unc ion
o
body
empe a u e
h oughou
his
ange
wi h
no
appa en
discon inui ies.
I
was
pos ula ed
ha
a
single
p ocess
ope a ing
be ween
2
and
250C
may
limi
he
du a ion
o
hibe na ing
pe iods.
*
These
s udies
we e
suppo ed
by
esea ch
g an
AM-05942
om
he
Na ional
Ins i u es
o
Heal h,
USPHS,
and
by
he
Uni e si y
o
U ah
Resea ch
Commi ee.
I
Pengelley,
E.
T.,
and
K.
C.
Fishe ,
Can.
J.
Zool.,
39,
105
(1961).
2
Twen e,
J.
W.,
and
J.
A.
Twen e,
J.
Appl.
Physiol.,
20,
411
(1965).
3Glass one,
S.,
K.
J.
Laidle ,
and
H.
Ey ing,
in
The
Theo y
o
Ra e
P ocesses
(New
Yo k
and
London:
McG aw-Hill,
1941),
p.
1.
THEORY
OF
THE
FLOW
OF
ACTION
CURRENTS
IN
ISOLATED
MYELINATED
NERVE
FIBERS,
VI*
BY
R.
LORENTE
DE
N6
AND
V.
HONRUBIA
THE
ROCKEFELLER
UNIVERSITY
Communica ed
June
10,
1965
We
con inue
he
p esen a ion
o
he
heo y
o
he
isola ed
ibe .'
Fo wa d
and
Backwa d
Jumps
o
he
Impulse.-When,
wi h
node
N,
in
he
cen al
pool,
he
s imula ing
de ice
is
placed
ac oss
he
i s
gap
and
he
ampli ie
ac oss
he
second,
poliphasic
ac ion
po en ials
a e
o en
eco ded
( e .
la,
Fig.
2,
1-8;
i ,
Fig.
2,
1-4)
which
may
be
in e p e ed
as
he
esul
o
successi e
jumps
o
he
im-
pulse,
i s
o wa d
and
hen
backwa d.
Howe e ,
since
he
la e
upwa d
peaks
may
emain
a e
anes hesia
o
he
segmen
o
ibe
in
he
dis al
pool
(la,
Fig.
2,
9-15),
he
poliphasic
spikes
may
also
be
in e p e ed
in
his
manne .
All
he
ac i e
zones
a e
c ea ed
by
he
applied
cu en ,
he
empo al
o de
o
hei
appea ance
being
de e mined
by
he
dis ibu ions o
he
elec o onus
and
o
he
s imula ion h eshold
in
he
in e nodes.
Figu e
1
illus a es
a
si ua ion
o
which
he e
is
only
one
possible
in e p e a ion.
The
si ua ion
is
excep ional
because
i
can
appea
only
when,
owing
o
di e en ial
inju y
done
du ing
he
isola ion
p ocess,
he
s imula ion
h eshold
is
lowe
in
he
dis al
pool
han
in
he
neighbo hood
o
he
cen al
pool
(c .
e .
ic).
Since
node
N,
was
loca ed
a
he
cen e
o
he
270-u-wide
cen al
pool,
a
ze o- esis ance
shun
placed
ac oss
he
i s
gap
blocked
p opaga ion
o
he
impulse
(Fig.
1,
2;
c .
e .
lb).
P opaga ion,
howe e ,
was
es o ed
when
in
addi ion
a
1-megohm
shun
was
placed
ac oss
he
second
gap.
Reco ds
3-8
(Fig.
1)
belong
o
a
se ies
ob ained
a
3-sec
in e als
du ing
a
3-min
pe iod;
hey
p esen
he
ex eme
ypes
o
ac ion
po en ials.
The
a iabili y
o
he
esponse
was
e e able
o
oscilla o y
changes
in
he
low
o
he
dema ca ion
cu en s
( e .
la)
which
caused
small
up
and
down
dis-
placemen s
o
he
base
line.
A
imes
p opaga ion
ailed
o
occu
and
only
a
small
upwa d
de lec ion
was
eco ded
(Fig.
1,
7),
bu
as
a
ule
he
ini ial
upwa d
peak
was
in e up ed
by
asha p
downwa d
peak,
which
was
ollowed
by
wo
upwa d
peaks
(Fig.
1,
4
and
6),
and
a
imes
he
wo
la e
upwa d
peaks
became
used
in o
a
la ge
