#Hoplosternum littorale$ (Hancock, 1828) shows a marked circadium rhythm of food demand. Feeding activity is mainly nocturnal, with two peaks, and is synchronized whith the diel light cycle. We tested the effect of infradian (period > 28h) light dark cycles on this rhythm in fish with demand-feeders : fish subjected to a 13.5 L/22.5 D light dark cycle for 8 days and a 25.5 L/10.5 D cycle for a further 8 days showed a strictly nocturnal feeding activity a response that only passively reflected the cycle alternation of light and dark. (Résumé d'auteur

Boujard, T.

Luquet, Pierre

Moreau, Yann

Horizon / Pleins textes

1 
Aqunt. Liring Resour., 199 I .  4, 22 1-22 j 
Entrainment of the circadian rhythm of food demand by 
infradian cycles of light-dark alternation 
in Hoplosternum littorale (Teleostei) 
Thierry Boujard ( I ) ,  >-ann Xloreau and Pierre Luquet 
Boujard T., Y .  Moreau. P. Luquet. . 4cpn r .  Liring Resow., I99 I ,  4. 221-225. 
Abstract Hoplosirrut?z littorde (Hancock. IS2Y) shows a markcd circadian rhythm of food demand. Feeding 
activity is mainly .nocturnal. with two peaks. and is synchronized with thc diel light cyclc. We tested 
the effect of infmdinn (period >?Y li) light,dark cycles on this rhythm in hsh with demand-feeders: 
fish subjected to a 13.5 L i 2 3  D light dark cycle For S days and a 75.5 L / ì 0 3  D cyclc t'or ;i further 
8 days showed a strictly nocrurnal feeding activity. a response that only passively rellected the cyclic 
alternation or  light and dark. 
Keywords : Siluriform. diel ch-cle, feeding rhythms. 
Résumé H~;plosierriirm liriorale (Hancock. IS'S) presente un rythme circadien marqué d'auto-distribution de 
nourriture. Ce rythme. d acrophase nocturne. se caracterise par deux pics. il est synchronisé par 
l'alternance jour/nuit de la lumiere. Nous avons mis l'épreuve le caractere circadien ( 2 4 5 4 h )  de ce 
rythme i l'aide de distributeurs d'aliments ((A la demanden. Les poissons soumis à un éclairage 
simulant la photopériode naturelle [ I 3 3  J!lO.j N) ont un rythme circadien d'auto-distribution d'ali- 
ments. Lorsqu'ils sont sujets i un cycle d'éclairage infradien (période >28 h) i longue scotophase 
(13.5J@.jN) durant Sjours. puis li longue photophase (25.5Ji10.5N) durant S autres jours. ils ont 
un cycle d'auto-distribution d'aliments qui est entrain6 sans délai par les changements successifs de 
cycles d'éclairage, et se nourrissent exclusivement durant les scotophases. 
Mots-clés : Siluriforme. rythmes circadiens, cycles nycthem6raus. alimentation. 
$ 
INTRODUCTION alternation of d a y  a n d  night, or dawn,  or dusk)  or 
temperature (Randolph  and Clemens, 1976). I t  may 
be endogenous, i. e. ,  persisting with its own period in 
constant environmental conditions, or exogenous if it 
only passively reflects the cyclic fluctuations of the 
environment. Circadian rhythms have been demon- 
A circadian rhythm (242~4 hrs.) is characterized by 
its amplitude. mean value (or Mesor) and acrophase 
(Gwinner, 1986; Halberg e r d . .  1977; Koukkari  et al., 
1987). It is generally synchronized by photoperiod (or 
Agircil. Lhing Rrsuur. 91/04 121 O5 s 2.50 8 IFREiMER-Gnurhier-Villars 
. . . . . ~ .. . . .  . . . _  _ _ _  +-.L, 
7 7 7  
I-" 
Four groups o f  1 1 1  Hop(c-~,\.i3,.!1!cr7i r!rorii/'c* tht-it hzid 
been hatched in the lahorutor:, aiid reared in pand.; 
[mean ue:sht  4qgi  in French Guianri a t  the Saci)u- 
mou Farm I 5c5.53c31 ' \ f~~  \\.ere dijtributed into 3W-! 
tanks. on the 1Sth of Januar:. IQQft .  The  tanks \vere 
continuousl:. supplied \sith reclycied i\ %iter [ temperat- 
ure 38i:3-C. oïygen at  saTuration1 dnd held in 3 
light-tight room ui th  artificihl lighting pro\.ided b) 
four 50 v+:jtt neon tubes contrillied b! ;in electric 
timer. For  a period of 13 da!s. the fish ivere wtt1ecteld 
to a 24hrs .  lighr c)cle that simulated the Katurai 
phoroperiild (13.5 L Ill.5 D I .  The  light c)ck \\:i$ 
T. Boujard C t  al. 
thereafter changed to a 36 hrs. light cycle \vith ;i 
longer scotophase for a period of 8 days from the 30th 
c71'lanuary 1990 [ 13.5 L 21.5 DI.  and subsequently to 
;i q c l e  ( i f36 hrj. \ \ . i r l i  a longer photophase for another  
period of S d a ~ s .  frani the 7th I-?Ï Februar! 19W 
(25.5 L l f i .5 DI.  
Each taiik \vas fitted wi th  3. metal rod \vhich closed 
;i 13 Y circuit \\hen a fish moved the rod. .A "70 l' 
feeder drli:ered 1 of food \.ia a rcld!.. This &\ice 
\ u s  cciuplrd t o  :I computer. Lvliich recorded the hour. 
date and tank number of each demand for food. In 
order to restrict unintentional triggering of the feeder 
b! the fish. the rod was loczted 2 cm below the \sater 
w r i x e .  i. <'.. far Îram the usual zone of activit! of 
this bottom d\veller tish. Furthermore. the thank 
ccmtant iy  drained from the thanks \\;itti the \vater 
$3 vert7 i:, w. \!.e there fo re k 11 o?\ t h 3 t the cc7 n j uni p t i o I: 
of fond csi:ld take place oni l  iiithin n Ìcl.\ minutes 
after denund-feedins  Operators entered the exper- 
imental room ont! e:-er: -3-4 dt-iya <incl h rncr  the tish 
n e r e  dizturhcd ;er>. little during the ri.perimen:. 
uL-t$ii 4- : ... \ ias sach :!ì,.ìt ünitjfijiiiiied i ü ~ d  pellefi ~ k 2 . 1 ' 2  
R ES L, LTS 
1 
Feeding rhythm in Hoplosterniini littorale 223 
Series 
Figure 2. - Quantity of Food self-distributed in each series expres- 
sed in of live weight per 1.5 hrs. during: A) 24hrs. cycle: 
I3.5LllO.5D; tanks I .  2, 3 ,  4. B)36hrs. cycle: 13.5L!13.5D: i 
1 
B 
22/01 23/01 24/01 25/01 26/01 27/01 28/01 29/01 
n I C O a * - œ  u 
~ m m œ - m o r  
1 1 1 1 3 L !  m n i - ) - - n O p a - * m  i BIL 
BELU ZII!3- 7 
Iu j-c 
2 KIM W I B  Ism m U 
I I  
06/02 
3 
30/01 31/01 01/02 02/02 03/02 04/02 05/02 
no data Y" -IC- 4r 'P 2 1 1 0  m l l l 7 1 1 I I 1 U -  ; 
4 IIlr m m  -E 
I l  1 07/02 08/M 09/02 10102 lli02 12/02 13/02 14/02 I 'P o m  :* o =F I m. = - I  m j  4 -I .-*I - œ - - L m : T  9+--- nodata . -.i . :k . .  I i .  I 
O 0.5 1 1.5 2 2.5 3 3.5 4 4.5 5 5.5 6 5 5 7 7 5  8 
Time (days) 
Et 
[ 
Figure 1. - Timing of the sclf-ciibtribution of food during thc dit'frrent lighting rcglmcs iincl ILr Ihr .  J gruup, 01' IO lish ~ t t 1 ~ 1 1 u J .  Bluck 
squares ind ia te  hours during which more than I g w;is demanded. tbhite squ:ires indicate hours during ivhich ulllh I g u'iis tlrmnntlcd. 
Scotophase corresponds to the hatched parts ur the graph. 
1 5  r ' Tank2 
Tank 3 
I = =  ; 
' -  
.A 
. .  .I. 1_1 i!, 
__I._ 
2 
I 
I 
1.1 1 
I 
I 1 .- 
0.5 
O 
Table 1. - Menn aniounis of self-distribuicJ food crprcsscd ;IS '",, 
of live weicht D ~ Y  unit time. 
Total duration of the light dark cycle (hrs.J 24.0 36.0 36.11 
Food-deninnd per hours during thc scoto- 
Fod-dCmt\IId..l-+ hrs. 3.9 J.0 7 -1 
Lcngth OF the scotophasc (hrs.) 10.5 10.5 37.5 
phase 11.37 0.57 0.24 
The high variability in food demand betwcen tanks 
and days was caused by moisture in the device, LvIiich 
happened in t a n k 4  on 25th and 26th January,  in 
tank 2 on 2nd and  3rd February, and  i n  tank 4 
between 7th and 14th February. During these periods; 
the fish were not able to obtain Food on demand.  
The observed differences of food demand between 
tanks were high, especially during the Inst phuse of 
the experiment. This \vas probably also caused by 
moisture in the device. making i t  more o r  less difficult 
for the fish of the different tanks to close the 1 2 V  
Tine. However, if these experimental failures restrict 
the significance of the results in terms of amoun t  of 
food demand,  they d o  not affect at  all the significance 
of the temporal variations of the demand. 
DISCUSSION 
The endogenous aspect of a diel cycle is generally 
tested with isolated individuals in a constant environ- 
ment (Millet a n d  Manachere, 1983; Koukkari  et al.. 
1957). In such a "free-running" situation, endo, menous 
rhythms are  revealed by their specific periodicity. each 
individual within a given species tending to exhibit a 
slightly different periodicity. In  the case of Hoploster- 
~ I W Z  lirrornle, individuals isolated from their con- 
geners did not use the rod to  demand food and  
223 T. Boujard cr al. 
starved (unpublished data 1 .  h.Ioreo\.er. sociril intercic- 
tions appear to be important in C ~ J ¡ ¡ ~ L - / I ~ J ~ ~ ~ ~ C ~  ( Krlinier 
and Graham. 197br. I n  H o p l o s ~ c r J ~ ~ i 1 7 1  i i i ~ o r d e  reared 
in ponds. one c;?n see air-breathing fish moving i n  
shoa!j of I O  t o  3 indiiiduals. swimming close to the 
bot tam.  and periodicall) mo\ , ing up to tlie surfLice 
one i f ter  the other (unpublished darai .  
I n  our experiment. in  the case of a n  e n d o p o u s  
rh!chin ;i decrease in self-feeding u n s  ekpected to be 
i]bser\.ed during late scotophase \\.hen the latter 15 
prolonged. or  d n  ûmet  of self-feeding \vas expected 
during late photophase \.vhen that phase is prolangecl 
Sei ther  of these t u o  responses \\ere obtnined. dnd 
the circadian rhythm of food intake \\;is synchronized 
I)! a n external :ìlid periodical stimulation, the light. 
A s  ;I result. in Hoplus;er1711111 / i ( tordc .  feeding activity 
inight Se J n  exopenous rhythm. and  if not. i t  i s  niil) 
ne~ikl!  coupied to :in endogenous oscillator. Lvith a 
inasking effect of tlie photophase.  The  strong relation- 
<hip berueen scotophase and feeding emphasize the 
ÏJet that th i s  species I S  truly nocturnal. and in relirins 
conditions. feeding t h t  fish by night might improve 
suri.iva1 or growth. hut this Iiypotl~rsis remains tû Se 
tested in ponds. Severtheless. c ~ u r  results show that  
:ircifíci:illy 1ncreasir.g tlie sso tophaje  length doez not 
iiicre:i';e dail! food coiisui-nption. 
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Entrainment of the circadian rythm of food demand by infradian cycles of light-dark alternation in Hoplosternum littorale (Teleostei)

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