Male mice are enclosed in cages of different dimensions (cage A - 23x8x8 cm., cage B - 36x27x17 cm.), in an alternating light/dark regimen, at an ambient temperature of 22 to 23 C. The successions of the behavioral sequences of ultradian activity periods are noticed by direct observation during 11 consecutive hours in light. The experimental situation modifies the mean duration time and the behavioral organization of each activity period. However, the comparison of the overall activity time lengths and the comparison of the overall behavioral frequencies suggest that the energy spent per mouse is constant

Guillot, A.

English

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NATIONAL AERONAUTICS AND SPACE ADMINISTRATION
WASHINGTON, D.C. 20546	 APRIL 1982
Ay ws#
NASA TECHNICAL MEMORANDUM
	
NASA TM-76848
BEHAVIORAL STUDY OF ULTRADIAN ACTIVITY PERIODS OF OF1 MICE
ENCLOSED IN EXPERI'^1TAL CAGES OF DIFFERENT DIMENSIONS
A. Guillot
(RASA-Tlt-1bd41i) bEhAVIONAL STUDY OP
	 BUZ-22dlU
ULTRADIAM ACTIVITY PERIODS OF GF1 MICE
ENCLOSED IN EYPERIARNIAL CAGES OF DIFFERENT
DIRENSIONS (National Aeronautics and Space
	 Unclas
Adainistration) 13 p HC A021ME A01 CSCL 06C G3/51 09632
Translation of ''Etude comportementale des p griodes d'activitLf
ultradiennes de souris OF1 placges en cages experimentales de
dimensions diff4rentes," Comptes Rendus Des S gances de la Socigtg
de Biologie et de Ses Filiales, Vol. 175, No. 3, 1981, pp. 295-302•
t
=—	
X
OF P OR
OF PO QUA,-rfY
ity periods has been accomplished by calculating the matrices of
	 r
the most frequent successions of behavior observed during 36 to-
tal periods. The results indicate that behavioral organization
is the same in both situations. Certain behaviors were never or /20-0
rarely associated (taking of food and construction of the nest=
exploration and washing at the end of activity). The most fre-
quent succession of behaviors is described in Figure 2, presented
in the form of a cycle based upon the same behaviors of cleaning
of the face and body.
Al
onn'VS. M+	 V'Ig, ; pingi3	
^In `n-vat	 ^rinkin$it^,1
	
iaa 
h:^L^T^! 
	
G
the es t
'	 EXPLORATION«1
gashing	 Feeding
in '^
► ^ c	 ngnest;
Nibbling	 Wa.-+%Ing
the ba=rs	 out6ide
the nest
Th_ 
..h *4gr. pp ::: .e arrows is pro-
	
portional	 tra.,:.:Oequency,of, },.h:e .sequences
for each behavior considered.
out continu-
ously for several Jlridter.
Figure 2. Diagram of the succession of behaviors
during ultradian activity periods during diurnal phase.
Correlations were calculated on the frequencies correspond-
A. Guillot
INSERM, U. 123, 44 chemin de Ronde, 78110 Le Vesinet. Ethology Labora-
tory of Communications, Lyon. Fsychophysiology Laboratory, Paris-X
Sum rev,
Male OF1 mice are enclosed in cages of different dimensions (cage
A = 23 x 8 x 8 cm, cage B = 36 x 27 x 17 cm), in an alternating LD
regimen, at an ambient temperature of 22-230C. The successions of the
behavioral sequences of ultradian activity periods are noticed by di-
rect observation during 11 following hours in L.
The experimental situation modifies the mean duration time and
the behavioral organization of each activity period. However, the
comparison of the overall activity time lengths and the comparison of
the overall behavioral frequencies suggest that the energy spent per
mouse is constant whatever is the experimental cage.
Introduction
Direct observation for 11 consecutive hours of male OF1 mice
placed in an alternating LD regimen (12:12) reveals the existence of
an ultradian activity rhythm and resting rhythm. This behavioral
rhythm, cited within some older studies, to our knowledge has not been
the object of systematic investigations.
Since a previous study has shown the influence of the experimen-
tal situation on the behavior of isolated or grouped animals C?3, one
will be interested here in the effects of the size of the cage in
which the animal is observed in order to state precisely if the dura-
tion and behavioral organization of ultradian periods remain stable re-
gardless of the experimental conditions.
*Numbers in the margin indicate pagination in the foreign text.
t ­= L-1411 -1) t) = Vm0, K,
1
Procedure
Sublects
Four 6-week-old male OF1 mice (IFFA CREDO Oncins), in an al-
ternating LD regimen (12 :12) for 2 weeks, were enclosed alone in
polystyrene cages measuring 23 x 8 x 8 cm (cage A) for four days,
then in polycarbonate cages measuring 36 x 27 x 17 cm (cage B)
for four days.
Four other male OF1 mice of the same age, in an alternating
LD regimen (12:12) for two weeks, were enclosed alone first in
cages B for four days, then in cages A for four days.
The results therefore concern eight males enclosed within
cages A with an area of 184 cm2 , and the same eight males en-
closed within cages B with an area of 972 cm2.
The animals are provided with a litter (sawdust), nourish- /U7
ment (food UAR), and drink (fresh water) ad lib itum, and with a
cotton ball for construction of a-nest.
Experimental Conditions
The illuminated phase is produced by the functioning of
white floureseent tubes (100 lux) from 0600 to 1800 hours. From
1800 to 0600 hours red flourescent tubes (10 lux) create a dark
phase, and permit eventual observations during the active period.
The temperature of the observation room was maintained at 22-230C.
Protocol
Observations were made on the fourth day in both experimen-
tal situations. It is a question of direct observations of four
males for 11 continuous hours (from 0800 to 1900 hours), consist-
ing of 10 hours during the diurnal phase and 1 hour during the
nocturnal phase. Cages were separated from the observer by a
transparent glass partition, and separated from each other by o-
paque partitions. Successions of behavior were surveyed for one-
2
Vminute periods. The behavioral sequences chosen are the fol-
lowing:
Coding
Washing
In The Nest
DEB (*)
FIN (*)
PDT
HS
EXPLO
ALIM
B OIS
NID
ENT, NID ( **)
FLAIR (**)
MOR BAR (**)
REPOS (**)
Behavioral1 Sequence
Washing of the face and body before excursion
outside the nest (prior to activity)
Washing of the face and body after the last re-
turn to the nest (after activity)
Washing of the face and body during activity
Brief washing4 of the face outside the nest
E cplorator yl f,iovement
Taking of :.4urishment from the feeding trough
Taking of drink from the feeding bottle
Construction of the nest
Entry into the nest
scenting: animal is immobile and scents, heed
raised
Nibbling the bars of the cage or of the feeding
trough
Immobility in the nest, eyes closed
*The results concerning these behaviors correspond to their dura-
tion, estimated through surveys at one-minute periods, seeing that
these behaviors are carried out uninterrupted for several_ minutes.
**These behaviors will not occur except during study of the behav-
ioral organization of an activity period.
Results
Ultradian Activity Period
Number of Activity Periods from 0800 to 1900 Hours Figure 11
The number of activity periods varies from 5 to 8.
Four out of eight mice develop more activity periods in cage
B.
Mean Duration and Distribution of Activity (Figure 1.1
The results deal with complete activity periods. Mean
duration: cage A:40.2 min. (s.d.* 25.4), cage B :25.9 min. (s.
d.w 10.3). The difference beiveen the two means is statisti- /0-8
cally significant (t-2.99, d.f.= 7).
3
ORIGINAL PACE 19
OF POOR QUALITY
AM
ecp "Ae'tT#T y period CAUS 4,
O	 C3	 O	 C=
A
•
•	
i
n 	 n
O	 00	 O	 O
A
•
b
M	 I	 =§
C3	 C	 C1
w	 M	 M	 NOW
A
•
c
'	
Jl
A
43	 O O	 O	 O	 no	 O a
! d
n
O	 O	 O O 0
A
•
p	 O	 0	 13
A
•; 2
A
O	 0	 p	 0 p • 3,i
A
4:3 	 i t=	 C:1	 C3	 O1	 i -A	 i s ,
t	 10	 /1	 12	 /S	 »	 17	 1t	 »Is
Figure 1. Distribution and duration of
ultradian activity periods in diurnal phase.
The mean duration of rest intervals in cage A is 93.5
min. (s.d.=56)= that of the first rest interval is 118.2
min. (s.d.:58), that of the last is 36.5 min. (s.d.=24).
In cage B the mean duration of rest intervals is 86.6
min. (s.d.=46.4)= that of the first interval is 120.7 min.
(s.d.=34), that of the last is 50.6 min. (s.d.s26).
In both cases the frequency of activity periods in-
creases at the end of the diurnal period. Observation car-
ried on one hour after the white light is turned off indi-
cates likewise that relative darkness does not immediately
stop the activity of most of the mice.
Behavioral Organization
An analysis of the behavioral structure of ultradian activ-
4
ORIGINAL PAGES
OF POOR Q i
ity periods has been accomplished by calculating the matrices of
the most frequent successions of behavior observed during 36 to-
tal periods. The results indicate that behavioral organization
is the same in both situations. Certain behaviors were never or /1-00
rarely associated (taking of food and construction of the nest$
exploration and washing at the end of activity). The most fre-
quent succession of behaviors is described in Figure 2, presented
in the form of a cycle based upon the same behaviors of cleaning
of the face and body.
^
^onSir gctio2 	 Y'^,t:i^ing
ns# T ^in ncat	 yr ekingfashJ'	
: f^4	
"Pi
E s t	 En k to1	 theNe
ws
PLOR
..J
^^whing	 Feeding
'fin K%.# c c ng	
%
'. nest; 	 (	 C )
Nibbl-ing	 WaV*Ing
the bars	 ouU ido
the nest
Th y, +r : phi= oxF 	 crows is pro-
portional	 tha,-I..:. sequency , of the -sequences
for each behavior considered.
_ ­1 ..' out continu-
ously for several .Ariuter. .
Figure 2. Diagram of the succession of behaviors
during ultradian activity periods during diurnal phase.
Correlations were calculated on the frequencies correspond-
ORIGINAL PAGE IS
OF POOR QUALITY
14 to each complete activity period. The test used is the cal-
culation of coefficients of the ranges of Kendall ( Table 1).
TAI LE 1. LINEAR CORRELATIONS BETWEEN BEHAVIORS, DURATION JF
	 /299
ACTIVITY AND OF REST IN SMALL CAGE -
 W AND LARGE CAGE (B) •2
I,--
-- 1--- ---- _ _--- ---°--	 --	 -_..^^ _. !VWarming -	 _ C
Itt:l
_- i
1` 18	 FIN	 D + t 3 RXI'1.	 ALiB	 isol, Nip	 DI'3 	11.3	 Is3
---^1 DI:8 0,51 - 0,23
llsWa ' ' !wi+b 	 FIN	 - 0,26 0,64	 0,40	 0,390 ,31	 ``-	 0,62
0,26	 0,47_	 -_
0,56
0,41	 0,23
D + F
	 0,29
CXPL 0,46	 0,43	 0055	 0,63
0,39
0.83
n
ALIN 0,40	 0,35	 0,23 	0,69 `a--	 _ 0,61 -	 0,51
	 0,58	 - 0,35
BOIs 0,86	 0,36	 0,26	 0,79	 0,67	 1 ---	
__ 0,63NID	 - 0,26 0,41	 0.52	 0„i3
	
0,45	 0,30	 0,42 0,39
DU 0,52
	 0,45	 0,67	 0,60	 0,51	 0,51 0,40
1P 0,28	 0.42	 0,37 0,36	 0.30	 - O
Is	 - 0.29 0,52 0,45	 0,30	 - _+
Large cage (B)	 N s 34	 T 11b 0.235
1Test of coefficient of Kendall ranges. Only significant re-
sults (p s .05) have been entered.
?-Commas in tabulated material are equivalent to decimal points.
3L + Fa DEB + FIN=	 DU= duration of activity period= 	 IP w du-
ration of rest period preceding activity period=	 IS s duration
of rest period following activity period.
Total Observation Period
Overall Activity Time Lenath:j During 11 xoyZs(In Minute s)
Comparison of the overall activity time lengths, carried out
by paired series, shows that mice are active just as long in cage
A as in cage B (t • 1.39, d.f.• 7). The difference of mean dura-
tion time in the longer periods reported therefore compensates
for the difference in the number of activity periods between the
mice in cages A and B.
6
-a
0FJjQ SAL PACE IS
Of POOR QUALITY
TA- '.E 2. OVERALL ACTIVITY T+ME
LENC' 1 ' !S DUR TWO 11 HOUR: ( IN MINUTES)
Cage A	 285	 197	 210	 198	 225	 144	 155	 151
Cage B
	 21L	 18;	 183
	
222	 172	 145
	 143	 176
Qyeral1 Be avioral.2reyuen_cies
TAi13LE 3. SUM OF THE FREQUENCIES OF
fA.11TORS DURING 11 HOURS IN CAGES A AND B
	
11 I \ ':'Uf OFI; PDT	 11S	 FIX I.%Pt	 1111	 1101 N111 I11,114imte
83 (' 54 0 41 29 144 24 16 14 285 1
97 0 55 1 16 41 72 32 10 31 197 2
05 0 53 6 22 36 115 37 23 24 210 3
Cage A
	
106 0 58 5 17 43 83 19 9 11 190 4
44 0 23 3 37 18 111 23 19 15 225 a
82 0 18 9 20 55 39 14 2 31 144 b
65 15 28 4 26 18 85 5 8 33 155 c
49 13 17 4 10 15 58 9 16 37 151 d
9S — 7 -- All	 - 3 20 39 92 28 ^ - 13 23 215 1
96 13 41 3 9 39 65 16 14 17 184 2
105 25 47 0 13 33 50 23 9 20 183 3
Cage B	 161 0 97 8 8 56 66 34 0 14 222 4
70 0 27 1 21 42 69 17 17 17 172 a
84 0 33 2 5 49 53 21 8 22 145 b
90 16 41 2 7 31 43 19 10 21 143 e
99 28 32 3 10 36 59 23 18 18 176 d
3.3u 2,47 1.22 0.38 6.72 2,34 2.53 O.61) 0.30 1.72 1.40 -Student t (') (') (1 (') (')
(*) Student t test on paired series significant
at p < 0.05.
TDN• sum of the washings in the nest] TOI: es-
timated duration of the activity periods com-
posed only of washinss DUR r duration of the
activity (in minutesg.
Discussion
	
/301
The results therefore show that the ultradian activity periods of
the diurnal phase present a similar succession of behaviors, and that
the animals organize their activity differently according to the exper-
7
imental situations. A sage with a more restricted volume and area
brings about a longer duration of activity periods and undoubtedly in
consequence a different connection of behaviors among them.
Indeed, in cage B the ► duration of ultradian activity, linked to
the preceding rest interval and for the most part to the behavioral
frequencies, appears to express an organized situation. The duration
of washings prior to and after activity varies in inverse fashion,
which maintains a better connection of the total duration of the activ-
ity and that of the preceding rest interval. These correlations, the
last of which is found again in the situation in cage A, could be in-
terpreted as a participation of tkermoregulation on the washing behav-
iors.
However caution should be used as to the function of washings pri-
or to activity. They were never connected to the duration of the ac-
tivity and have a negative connection with the frequency of construc-
tion of the nest. These facts could exclude them from all participa-
tion in the regulation of the temperature of the animal, contrariwise
then to washings following activity.
The less numerous correlations between the different behaviors
appears to express a less organized situation. This could come from
an emotional reactivity, more important when the size of the cage is
restricted [8]. The increase of substituted activities (brief washing
of the face [91) could also confirm it. In cage A rest time following
activity is shorter as the frequency of taking of food increases. The
animal therefore activates itself all the more rapidly as its accumu-
lated supplies are all the more numerous. This result was never found
in cage Bs it is possible that in a cage of restricted size, the ani-
mals only orjanize their activity around the taking of nourishment,
whereas in a larger cage their activity is a function of all behaviors.
The positive connection between the frequency of movements of con-
struction of the nest and the duration of rest intervals following ac-
^	 8
tivity, found in both situations, in noteworthy on the one hand be-
cause the duration of rest following activity is thus predicted, and
on the other hand because it suggests the role (similar to that of
washing in the nest) of thermoregulations indeed, the animal arranges
its nest all the more as more time passes without activity, during
which time its body temperature decreases. The positive connection
between exploratory frequency and that of taking of food and of drink-
ing was found equally in both situations, a connection furthermore
pointed out in us mus us [10] and Peromyscus manioulatus [5].
When the investigation on the total activity of the animals dur-
ing it consecutive hours was completed, one reports that the duration
of that activity is the same, regardless of the experimental situation.
Besides, investigation of the overall behavioral frequencies shows
that the animals reacted to changing situations through a modification 1.102
of the number of exploratory excursions, more numerous in a restricted
cage. The frequency of taking of food, of drinking, and of construc-
tion of the nest, did not undergo and modifications it is possible
that the mice increase their locomotive activity in cage A or decrease
it in cage B in order to maintain an identical level of activity. In-
deed, a locomotive excursion in cage B could represent a more impor-
tant expenditure of energy than an excursion in cage A t the distance
traversed being greater in the former case.
Washings in the nest have a more significant total duration time
in cage B, although washings outside the nest are more frequent in
cage A. The animals therefore have a tendency to reduce the duration
of washings in the nest when exploration is more important, bringing
about a greater frequency of washings outside the nest, with a regula-
tion of the total duration time of this cleaning behavior.
The preceding analyses suggest therefore that the observed dif-
ferences in the two cages could be due to at reaction or adaptation of
the animals to the situation by taking into account constant expendi-
ture of energy.
9
-_	 a	 MW
The manner of surveying behaviors used here therefore permits us
to state that the mean duration time of ultradtan activity periods and
their behavioral organization varies according to the experimental sit-
uation. The mice also modify exploratory frequency and the duration
of certain washing actions as a function of the size of the cage, but
maintain the same occurrence of most other behaviors. Interpretation
of these results indicates a constant expenditure of energy by the ani-
mal within the time period considered. Further accounts of variables
linked to energetic metabolism could permit a better understanding of
the underlying mechanisms.
.REFERENCES
1. Stier, T.J., Jow. Gen. Psvchol. .4, 67 (1930).
2. Behney, W.H., Jour._Mommalogy U. 232 (1936)•
3. Hatfield, D.M., Jour. laloov U. 261 (1940).
4. French, R.L., ,oVM. Mmmalomr U. ?4 (1955).
5. Kavanau, J. L. 	 of 1M U, 161 (1962).
6. Kavanau, J.L., Eccolacv 449 95 (1963).
7. Guillot, A., C. R. Some. _ igI . 32_1 9 543 (1977) •
8. Guillot, A., "Contribution 11'itude d'effets sociaux che2 la
Souris on cage expirimentale," These III e cycle, Lyon-
9. Grant, E.C., B2hav= 21, 261 (1963).
10. Achelis and Northdurft in Calhoun, J.B., Z. of Tenessee Acad, of
Science_  12, 300 (1944).


Behavioral study of ultradian activity periods of mice enclosed in experimental cages of different dimensions

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