Chronobiology International

The Journal of Biological and Medical Rhythm Research

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Phase response curve to 1 h light pulses for the

European rabbit (Oryctolagus cuniculus)
Gerard A. Kennedy & Robyn Hudson
To cite this article: Gerard A. Kennedy & Robyn Hudson (2016): Phase response curve to 1 h
light pulses for the European rabbit (Oryctolagus cuniculus), Chronobiology International, DOI:
10.1080/07420528.2016.1191506
To link to this article: http://dx.doi.org/10.1080/07420528.2016.1191506

Published online: 15 Jun 2016.

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CHRONOBIOLOGY INTERNATIONAL
http://dx.doi.org/10.1080/07420528.2016.1191506

Phase response curve to 1 h light pulses for the European rabbit (Oryctolagus
cuniculus)
Gerard A. Kennedya and Robyn Hudsonb
School of Psychology, Counselling and Psychotherapy, The Cairnmillar Institute, Victoria, Australia; bInstituto de Investigaciones Biomdicas,
Universidad Nacional Autnma de Mxico, Ciudad de Mexico, Mexico

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ABSTRACT

ARTICLE HISTORY

While much is known about the circadian systems of rodents, chronobiological studies of other
mammalian groups have been limited. One of the most extensively studied nonrodent species,
both in the laboratory and in the wild, is the European rabbit. The aim of this study was to extend
knowledge of the rabbit circadian system by examining its phasic response to light. Twelve DutchHimalayan cross rabbits of both sexes were allowed to free-run in constant darkness and then
administered 1 h light pulses (1000 lux) at multiple predetermined circadian times. Changes in the
phase of the rabbits circadian wheel-running rhythms were measured after each light pulse and
used to construct a phaseresponse curve (PRC). The rabbits PRC and free-running period ()
conformed to the empirical regularities reported for other predominantly nocturnal animals,
including rodents and predatory marsupials. The results of the study are thus consistent with
reports that the rabbit is essentially a nocturnal animal and show that it can entrain to light/dark
(LD) cycles via discrete phase shifts. Knowledge about the rabbits circadian range of entrainment
to LD cycles gained in this study will be useful for examining the putative circadian processes
believed to underlie the unusual rhythm of very brief, once-daily nest visits by nursing rabbit
mothers and other nursing lagomorphs.

Received 4 January 2016

Revised 15 May 2016
Accepted 16 May 2016

Introduction
For a circadian clock to be entrained by a daily 24
h light/dark (LD) cycle (zeitgeber), it must be reset
each day by an amount that corrects for the difference between its period tau () and the period
of the LD cycle (T). This is achieved by a daily
cycle of changing sensitivity within the organism
to light (Aschoff, 1960; Daan & Pittendrigh,
1976a). This daily rhythm of sensitivity to light
can be quantified in terms of a phaseresponse
curve (PRC) to short (1560 min) light pulses.
The characteristic features of PRCs in response to
short light pulses in both nocturnal and diurnal
mammals are phase delays during the late subjective day and the early subjective night, and phase
advances in the late subjective night and early
subjective day. In nocturnal rodents, subjective
night approximates circadian time (CT) 12 to
CT24, and subjective day CT24 to CT12. The
greater part of the subjective day (CT2 to 10) is
characterized by a dead zone, during which the
CONTACT Gerard A. Kennedy
gerard.kennedy@cairnmillar.edu.au
993 Burke Road, Victoria, 3124, Australia. Tel: + 61 (0)3 9813 3400.
2016 Taylor & Francis

KEYWORDS

Circadian rhythms;
free-running rhythm;
phaseresponse curve;
phase advance; phase delay;
rabbit; Oryctolagus cuniculus

circadian system is largely unresponsive to light

(Daan & Pittendrigh, 1976a).
Pittendrigh and Daan (1976a, 1976b, 1976c)
proposed that the entrainment of nocturnal rodent
circadian rhythms is based on the effects of nonparametric changes in light intensity on the animals circadian clock at different times of the day.
The nonparametric model of entrainment is based
on the premise that the influence of the zeitgeber
is brief, causing discrete phase shifts of the circadian rhythm, either advancing or delaying its
phase on a daily basis so that its period matches
that of the zeitgeber. They also described the process of entrainment that occurs when animals are
exposed to the full photoperiod rather than just
brief, discrete, bursts of light at dusk or dawn.
They proposed that in this situation the sum of
the discrete phase delay and advance shifts can be
transposed into a velocityresponse curve. Thus,
light causes a continuous series of delay and/or
advance (parametric) phase shifts, resulting in
either the slowing down or speeding up of the
School of Psychology, Counselling and Psychotherapy, The Cairnmillar Institute,

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G. A. KENNEDY & R. HUDSON

circadian clock (modulation of clock velocity). The

result of this process is that the circadian clock will
establish a unique phase relation to the LD cycle,
either phase leading or phase lagging depending
on the periods of and T.
Although the characteristics of the PRC are
similar in all nocturnal rodents studied to date,
there are consistent differences in the shape of
the PRCs. Daan and Pittendrigh (1976a) determined that there is an empirical relationship
between the shape of the PRC and . In nocturnal
species with a mean of <24 h, the ratio of the
area under the delay section to the area under the
advance section (D/A) of the PRC is >1, whereas
in species where is >24 h D/A is <1. The shorter
is, the greater the D/A ratio. Thus, species with
<24 h respond to light pulses with large phase
delays and small phase advances, while species
with >24 h primarily entrain via phase advances.
There are a number of parameters that can affect
the PRC shape, and these have been investigated in
mice (Mus musculus) by Comas, Beersma,
Spoelstra and Daan (2006, 2007) and Comas,
Beersma, Hut and Daan (2008). These parameters
include the duration and intensity of the light
pulse given in DD, whether PRCs are derived
from LD transitions, the point within the light
pulse where measurements are derived, the plotting of the onset versus the offset of the activity
rhythms and the phase response to light (modulation of clock velocity).
The circadian system of nocturnal rodents has
been extensively investigated, but little is known
about the characteristics of the circadian systems
of other mammalian taxonomic groups (review in
Johnson, 1999). PRCs for two marsupial species,
the stripe-faced dunnart (Sminthopsis macroura)
(Cassone, 1987) and the kowari (Dasyuroides
byrnei) (Kennedy et al., 1989), have been reported
and found to conform to the characteristics shown
by nocturnal rodents. The D/A ratio for these
species conforms to the empirical regularity suggested by Daan and Pittendrigh (1976a), and like
the rodent species investigated by these and other
researchers, considerable variation was found
between species in the waveform, amplitude and
location of the transition point from phase delays
to advances (cf. Cassone, 1987; Kennedy et al.,
1989). These marked differences in the waveform

and timing of species PRCs and the discovery of

empirical regularities suggest the utility of investigating and comparing the phasic response to light
in a variety of mammalian groups including diurnal species (Lee & Labvak, 1997; Slotten et al.,
2005).
The European rabbit (Oryctolagus cuniculus;
order Lagomorpha), the ancestor of all domestic
rabbit breeds, provides a particularly good opportunity to extend knowledge of the biological significance of and physiological processes
underlying mammalian circadian biology. Due to
the rabbits significance in animal production, as
an agricultural pest, and as a model species in
biomedical research, much is known about its
biology, including under natural and semi-natural
conditions. Specifically, diurnal and circadian processes have been studied in the rabbit in an unusually diverse range of laboratory and natural
conditions, and in a wider variety of biological
contexts than in most other mammalian species
(review in Gonzlez-Mariscal et al., 2016; Jilge &
Hudson, 2001; Schlolaut et al., 2013).
The wild rabbit is a burrow-dwelling, essentially
nocturnal animal, sheltering below ground during
the day and emerging primarily at dusk
(Mykytowycz & Rowley, 1958; Villafuerte et al.,
1993; Wallage-Drees, 1989; review in Jilge &
Hudson, 2001). Reports on the daily pattern of activity in the laboratory rabbit, however, are equivocal,
with some authors postulating an attenuated rhythm
(e.g. Horton et al., 1974), and others reporting that
the rabbit is diurnal (e.g. Katz et al., 1975), crepuscular (e.g. Nuboer et al., 1983) or nocturnal (e.g.
Pivik et al., 1986). As a result of a series of longterm investigations, Jilge (1980; 1991) reported characteristics of nocturnality in the laboratory rabbit.
He found that most activity occurred during the dark
period of a 12:12 LD cycle, and that the length of the
free-running period in constant light was >24 h,
decreased with decreasing light intensity and in a
manner characteristic of nocturnal species. He concluded that the laboratory rabbit is an endogenously
nocturnal animal, but that its rhythms are sensitive
to a number of other external factors. While the sex,
breed and age of rabbits do not significantly affect
the 24 h rhythm, external animal house noise and
restricted food access have a considerable effect and
may result in the inversion of the endogenously

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CHRONOBIOLOGY INTERNATIONAL

nocturnal rhythm and in the manifestation of a more

diurnal pattern of activity (Jilge, 1991). Jilge (1980)
also examined the phase response of the rhythm of
caecotrophy in four rabbits and found it conformed
to the response typically shown by nocturnal species.
However, the limitations of this study were that it did
not examine the rabbits under free-running conditions, the caecotrophy rhythm was examined rather
than the general locomotory rhythm and only four
rabbits were assessed.
Given the interest in the rabbit by chronobiologists, it is surprising that a full PRC for this
species has still not been constructed (Johnson,
1999; but see Jilge, 1980). This information
would be useful both to compare the function of
the rabbits circadian system with that of other
(nocturnal) mammals and to assist in the appropriate design of future experiments. Therefore, the
aim of this study was to investigate the characteristics of the rabbits circadian system by examining
its phasic response to 1 h light pulses under freerunning conditions in continuous darkness.

available ad libitum and were replenished each

second day at random times between 07:00 and
24:00. Sets of infrared beams, connected to a logic
circuit sensitive to the sequence of interruptions,
were strategically placed outside the drinking,
feeding and nest-box compartments to determine
the duration of time spent in these areas. Runningwheel revolutions were used as a measure of locomotor activity. Each wheel revolution resulted in
the closure of a reed switch via a magnet attached
to the perimeter of the running wheel. Wheel
revolutions were monitored and recorded at 15
min intervals using a custom-built, data acquisition computer. Actograms were subsequently
plotted in a manner that simulated the actograms
produced by paper event recorders, using a convergent-technologies CT 640 computer (Kennedy
et al., 1989, 1994). Two 40 W fluorescent tubes
(Thorn, Australia) suspended from a metal tube
frame via adjustable chains were used to administer light pulses. During light pulses, the rabbits
were confined individually in their own nest
boxes covered with a transparent acrylic top to
allow light to enter.

Materials and methods

Animals and apparatus

Procedure

Four female and eight male Dutch-Himalayan

cross rabbits were included in the study. The rabbits were obtained from private breeders and were
68 months old at the commencement of the
study. Prior to the experiment, the rabbits were
maintained in a semi-outdoor enclosure and were
subject to the natural photoperiod of late summer
in Melbourne, Australia. During the study, the
rabbits were individually housed in large acrylic
boxes (120 63 43 cm) with a wire mesh top
(Kennedy et al., 1994). The cages were situated in a
light-tight, sound-attenuated and temperaturecontrolled (21 2C) laboratory. A running
wheel 57 cm in diameter and with a track width
of 17 cm was located at one end of each cage. The
layout of each cage was as follows: compartments
for feeding and drinking, a removable nest box
(25.5 32 23 cm), and an open area linking
these compartments to the running wheel. Rabbits
were fed a mixture of rye grass hay, lucerne hay,
carrots, and rabbit and guinea pig rations
(Barastock, Australia). Food and water were

Rabbits were maintained in constant darkness (DD)

until stable free-running wheel-running activity
rhythms were established (e.g. Figure 1). When these
were attained, 1 h light pulses (1000 lux) were administered at predetermined CTs. Subsequent light
pulses were not given until at least 14 days had elapsed
following the preceding light pulse. The animals were
administered light pulses in a room close to the
laboratory in which they were housed to minimize
disruption of their behavior. Rabbits were confined in
their own nest boxes to minimize the possibility of
phase shifts occurring as a result of environmental
change (cf. Mrosovsky, 1988). Temperature in the
nest boxes during light pulses was measured and
found to remain at 21 2C. Light pulses were administered to the rabbits at different CT in a random
order. To determine the phase of activity onsets,
regression lines were fitted to the onsets of at least
10 days of stable activity before and after each light
pulse. Phase shifts were measured on the day following the administration of light pulses and the magnitude determined as follows: the distance (mm)

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G. A. KENNEDY & R. HUDSON

Figure 1. Rabbit # 3 free-running in DD showing phase shifts in wheel-running activity resulting from light pulses given at CT21, 12,
3 and 24.

between the extrapolated pre- and post-light pulse

onsets of the activity rhythm was measured on the
double plotted activity records. This measurement
was divided by a factor of 5.125 and multiplied by
the mean of the pre- and post-phase shift /24 in each
case in order to convert it to circadian hours
(Kennedy et al., 1989). Phase advances were designated by a positive sign and phase delays by a negative
sign. Twelve rabbits were used in the study and the
majority received light pulses at each designated CT.
However, some rabbits did not receive light pulses at
all CTs due to logistic difficulties in accomplishing the
timing of this procedure (the n for each data point is
given on the PRCs in Figure 2). The mean phase shift
was calculated for each light pulse at each CT, and
PRCs were constructed using a computer-generated

line of best fit (Harvard Graphics, ver. 3.0) (Figure 2).

The experimental protocol was approved by the
Animal Ethics Committee, Department of
Psychology, La Trobe University, Melbourne,
Australia, where the study was performed. The study
also conforms to the ethical standards and methods
for the use of animals in biological rhythm research
(Portaluppi et al., 2010).
Results
In DD, rabbits
with ranging
rabbits free-ran
free-ran with
ran with very

showed free-running rhythms

from 23. 20 to 24.69 h. Six
with <24 h, four animals
>24 h and two animals freeclose to 24 h. The mean free-

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CHRONOBIOLOGY INTERNATIONAL

Figure 2. (A) The phaseresponse curve to 1 h light pulses administered in DD for the rabbit (n = 12). (B) The phaseresponse curves
for short and long groups of rabbits. The asterisks represent the
short (23.87) and the diamonds represent the long (24.16)
animals (numbers in brackets = the number of rabbits administered
light pulses at each circadian time).

running in DD was 23.97, SE 0.29 h, for the

12 rabbits. The mean free-running in DD for
females (n = 4) was 23.94, SE 0.06 and for
males (n = 8) it was 23.91, SE 0.08. This difference in length was not significant (t(10) =
0.12, p = 0.91). Figure 1 shows an example of
the free-running rhythm of wheel-running
activity of a rabbit administered light pulses at
selected intervals. The effects of light pulses on
the phase of the rabbits rhythms are summarized in Figure 2(A) and (B).
Light pulses administered during the greater
part of the subjective day (CT3 to CT12) and the
greater part of the subjective night (CT12 to
CT19.5) delayed activity onset (Figure 2A). The
maximum phase delay was 1.6 h. Light pulses

administered during the late subjective night

(CT19.5 to CT24) advanced activity onsets. The
maximum phase advance was 0.4 h. The early part
of the subjective day (CT24 to CT3) was characterized by a dead zone, where light pulses had
little or no effect on the phase of the activity
rhythm. The transition point from delays to
advances occurred at CT19.5. The D/A ratio was
7.47. No transient cycles that were either longer or
shorter than the usual daily cycles were evident
following either phase delays or advances. An estimate of the rabbits possible range of entrainment
was derived by subtracting the maximum phase
advance from the rabbits mean free-running
for the lower boundary and adding the maximum
phase delay to the mean for the upper boundary.
The estimated range of entrainment is from a
lower boundary of 23.57 h to an upper boundary
of 25.57 h.
There was a significant although weak correlation between the pre-light pulse and the magnitude of phase shifts (r = 0.30, p = 0.05). Light
pulses administered during the early subjective
day (CT24 to CT4.5 and CT10.5 to CT12) and
during the subjective night (CT12 to CT24) caused
a shortening of , whereas light pulses administered during the greater part of the subjective day
(CT4.5 to CT10.5) resulted in a lengthening of .
These changes, however, were small in magnitude
( lengthening up to 0.04 h and shortening up to
0.15 h) and the standard errors were large. These
changes in length reflect the effects of light
slowing down and speeding up the circadian
clock (modulation of clock velocity) (Pittendrigh
& Daan, 1976b, 1976c).
For a more detailed analysis, the rabbits were
divided into two groups according to their individual
mean free-running . A short group included rabbits
with shorter than 24 h (n = 6; four males, two
females), and a long group included rabbits with
equal to or longer than 24 h (n = 6; four males, two
females). The mean for the short and long groups
were 23.87 h, SE 0.1 h and 24.16, SE 0.18 h, respectively. The mean for females (n = 2) in the short
was 23.91 h, SE 0.08 and for females (n = 2) in the long
group it was 24.06 h, SE 0.02. The mean for males
(n = 4) in the short was 23.89 h, SE 0.07 and for
males (n = 4) in the long group it was 24.12 h, SE
0.09. Figure 2(B) shows the PRCs constructed for the

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G. A. KENNEDY & R. HUDSON

short and long groups. The PRC for the short

group showed delays during the greater part of the
subjective day and the subjective night (CT2 to
CT20), and advances in the late subjective night
(CT20 to CT24). The early subjective day (CT24 to
CT3) was characterized by a dead zone. The PRC
for the long group showed delays during the greater
part of the subjective day and early subjective night
(CT4 to CT18), and advances during the late subjective night and early subjective day (CT18 to CT24).
The early subjective day (CT24 to CT4) was characterized by a dead zone. The transition point from
delays to advances occurred at CT20 and CT18 for the
short and long groups, respectively. In addition, the
maximum phase delays were greater in the short
(2 h) relative to the long group (0.75 h), whereas the
long group displayed larger maximum phase
advances (0.75 h compared to 0.25 h). The D/A ratio
was calculated for both groups and found to be 27.14
in the short group and 0.98 in the long group.
Discussion
The characteristics of entrainment to LD cycles in
the rabbit are consistent with those outlined for
nocturnal rodents in the introduction (Aschoff,
1960; Daan & Pittendrigh, 1976a). Under natural
conditions where rabbits typically spend the daylight hours in the safety of their burrows
(Mykytowycz & Rowley, 1958; Villafuerte et al.,
1993; Wallage-Drees, 1989; review Jilge &
Hudson. 2001), entrainment most likely occurs
via the nonparametric phase-shifting effects of
light. Whereas phase delays are affected by light
pulses falling during the latter part of the subjective day and early subjective night, phase advances
are affected by light pulses falling in the late subjective night and early subjective morning (cf.
DeCoursey, 1960, 1961). In the present study, the
rabbits free-ran in DD with a mean of 23.97 h,
although some individuals had s that were >24 h.
The PRC indicates that the rabbits range of
entrainment can extend from approximately
23.02 h to 25.57 h, which is in close agreement
with the observed range of actual s (23.20
24.69 h). The observed phase delays of up to
1.6 h and phase advances of up to 0.95 h appear
to be of sufficient magnitude to reset free-running
rhythms within this range of s to 24 h, and thus

to ensure entrainment to the 24 h day. When

divided into two groups according to their individual mean , rabbits with s shorter than 24 h
exhibited phase delays of up to 2.4 h and phase
advances of up to 0.6 h. Rabbits in the long
group showed phase delays of only up to 0.8 h
and phase advances of up to 1.1 h. These phase
shifts were also of sufficient magnitude to entrain
rabbits with in the range observed (23.7023.98 h
and 24.0224.39 h for the short and long groups,
respectively).
The correlation, although weak, between prelight pulse and phase shifts suggests that the
longer the before the phase shift, the more likely
it is that the phase shift will be an advance. This
has functional significance in that animals with
long free-running rhythms require an overall
phase advance of the rhythm to entrain to a 24 h
LD cycle. However, as the correlation was only
small, it is likely that the modulation of has no
significant role in entrainment to LD cycles in the
rabbit in the wild. In addition, an analysis was
performed subsequent to each phase shift to determine whether any systematic change in period
modulation occurred. In animals with a short ,
it was found that when a light pulse caused a delay
of the rhythm the period shortened in half of the
cases and lengthened in the other half. A similar
pattern emerged for all other possible combinations. Thus, error due to over- or underestimation
of the phase shift as a result of period change
balanced out and no systematic bias was apparent.
This is consistent with the results of Daan and
Pittendrigh (1976a), who reported that the golden
hamster (Mesocricetus auratus) and white-footed
deer mouse (Peromyscus maniculatus) also showed
small changes in following either delay or
advance phase shifts. Similarly, no effects of light
pulses on were found for either the stripe-faced
dunnart (Cassone, 1987) or kowari (Kennedy
et al., 1989). By contrast, in the laboratory rat
(Rattus norvegicus), length appears to be highly
susceptible to modification by light pulses
(Summer et al., 1984). These authors postulated
that the circadian system of nocturnal rodents can
respond to changes in the presence of environmental light by either acutely (nonparametrically)
phase-shifting the entire rhythm or by continuously (parametrically) altering the frequency of

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CHRONOBIOLOGY INTERNATIONAL

the circadian clock. The present findings, together

with what is known about the behavior of rabbits
in the wild, suggest that although the circadian
system of the rabbit can be modulated parametrically, under natural conditions it predominantly
utilizes the nonparametric mechanism.
As reported above, wild rabbits spend most of
the day underground shielded from the light,
emerge around dusk, graze throughout the night
and return to the burrow in the early hours of the
morning. Findings from experiments employing
simulated den systems in squirrels (DeCoursey,
1960, 1961, 1986a, 1986b) support this view.
DeCoursey reported that when flying squirrels
(Glaucomys volans) with free-running of <24 h
were provided with light-proof nest boxes, they
made extremely few trips into the light and often
saw no light for several consecutive days.
Entrainment consisted of a free-running state
with discrete instantaneous phase delays when
light exposure occurred every few days and
resulted in a saw-toothed pattern of activity onset.
A comparison of the PRCs for the four species
of nocturnal rodents studied by Daan and
Pittendrigh (1976a) showed that there was considerable variation among species in waveform,
amplitude and the location of the transition
point from phase delays to phase advances. In
addition, two marsupial species, the stripe-faced
dunnart and the kowari, exhibited marked differences in the form and timing of their PRCs
(Cassone, 1987; Kennedy et al., 1989). While the
rabbit PRC exhibits essentially the same characteristics as those derived for rodent species, some
differences from these in the three parameters
mentioned above are also apparent. The rabbit
PRC obtained in the present study appears to be
similar to the PRC documented for the whitefooted deer mouse, a nocturnal rodent with a
free-running period similar to that of the rabbit
(23.99 compared to 23.97 for the rabbit). The
amplitude for both delays and advances are also
similar in the two species, although the maximum
delays for the rabbit are of a slightly larger magnitude. The transition point from delays to
advances also occurs at approximately the same
time in both species. The rabbit has a smaller
advance area than the deer mouse and tends to
show phase delays earlier in the subjective day

than any of the other three rodent species or the

two marsupials species. A comparison of the rabbit PRC to those for two diurnal rodent species
Octodon degus (Lee & Labyak, 1997) and
Arvicanthis ansorgei (Slotten et al., 2005) showed
that it was more similar in form to the nocturnal
species described above for various rodents and
marsupials, with a bigger and earlier delay area.
A relationship between PRC shape and has
been reported in four nocturnal rodents (Daan &
Pittendrigh, 1976a). The longer the species mean
, the smaller was the D/A ratio. This empirical
relationship has subsequently been found in a
number of other species including flying squirrels (DeCoursey, 1986a, 1986b), stripe-faced dunnarts (Cassone, 1987) and kowaris (Kennedy
et al., 1989). However, the laboratory rat appears
to constitute an exception to this relationship.
Honma et al. (1985) reported a D/A ratio of
3.9, higher than would be expected for animals
with a mean of 24.43. In addition, when the
rats were divided into a short (24.33 h) and a
long group (24.53 h), a larger D/A ratio was
observed in the long group (4.7 versus 2.5).
However, the results of the present study are
consistent with the empirical relationship
reported by Daan and Pittendrigh (1976a).
Given the rabbits mean of 23.97 in DD, it
was expected that it would have a ratio of D/A
>1.0. The D/A ratio was found to be 7.47. When
separate PRCs were constructed for the short
(23.87) and for the long (24.16) groups, the
D/A ratios were found to be 27.14 and 0.98,
respectively.
In conclusion, the results of the present study
are consistent with reports that the rabbit is essentially a nocturnal animal, which can entrain to LD
cycles via discrete phase shifts in a manner broadly
similar to the nocturnal rodents and nocturnal
marsupial species studied to date (see also Jilge,
1980). Knowledge gained from the present study
in relation to the range of entrainment of the
rabbit may be a useful tool for studying putative
circadian processes underlying, for example, the
unusual pattern among lagomorphs of brief,
once-daily nursing visits by the mother
(Gonzlez-Mariscal et al., 2016; Hudson et al.,
1995; Jilge & Hudson, 2001; Ninomiya-Alarcn
et al., 2004; Rdel et al., 2012).

G. A. KENNEDY & R. HUDSON

Acknowledgements
We thank Mark Hamalainen, John Wright, Ian Moore, David
Johns, Mark Brooks, Nick Maerz, David Walsh and Carolina
Rojas for excellent technical support, and the following psychology students for their valuable contribution to data collection: Catriona Ross, Maria Galati, Linda Marson, Connie
Salamone and Clara Stojanovska.

Declaration of interest
The authors declare no conflicts of interest for this report.

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