Effect of Melatonin and Lighting Schedule on Energy

Metabolism in Broiler Chickens

E. J. APELDOORN,* J. W. SCHRAMA,*,1 M. M. MASHALY,† and H. K. PARMENTIER*

*Animal Health and Reproduction Group, Wageningen Institute of Animal Sciences, Wageningen Agricultural University,
P.O. Box 338, 6700 AH Wageningen, The Netherlands, and †Department of Poultry Science,
College of Agricultural Sciences, The Pennsylvania State University,
University Park, Pennsylvania 16802

ABSTRACT The effect of melatonin and lighting
schedule on energy metabolism in broiler chickens was
studied. Eight groups of six female broiler chickens each
were assigned to a continuous lighting schedule [23 h
light (L):1 h darkness (D)] or an intermittent lighting
schedule (1L:3D), and were fed a diet with or without
melatonin (40 ppm). At 21 d of age, the chickens were
placed in respiration chambers for 20 d. Energy and
nitrogen balances, heat production, and physical activity
were measured per group. The only effect of melatonin
on energy metabolism, was a decreasing effect on
activity-related heat production. The intermittent light-
ing schedule induced improved feed conversion, higher
metabolizability of the diet, and lower physical activity
compared to continuous lighting. No interactions be-
(Key words: broiler, melatonin, lighting schedule, heat production, circadian rhythm)
tween melatonin and lighting schedule were found on
energy metabolism traits. Lighting schedule strongly
affected daily heat production pattern (total, activity-
related, and nonactivity-related heat production).
Melatonin had a reducing effect on activity-related heat
production during the day, especially during light
periods. The present study demonstrated that reduced
energy expenditure for physical activity, caused by the
supplementation of melatonin to the diet, might be a
reason for the often observed improvement of feed
conversion. Furthermore, this study showed that feed
conversion was improved with an intermittent lighting
schedule, which was related to higher metabolizability
and lower energy expenditure on physical activity,
compared to continuous lighting.
1999 Poultry Science 78:223–229

INTRODUCTION Broiler chickens normally do not eat during darkness,

Melatonin is a hormone that is secreted from the
pineal gland. It regulates daily and seasonal physiologi-
cal rhythms, including the cardiopulmonary, reproduc-
tive, excretory, thermoregulatory, behavioral, im-
munomodulatory, and neuroendocrine systems (Pang et
al., 1996). Melatonin also plays an important role in the
antioxidant defense system (Barlow-Walden et al., 1995).
The pineal is photosensitive; melatonin in birds is
secreted mainly during darkness (Pang et al., 1996). Most
studies find that melatonin depresses feed intake and
improves feed conversion, but its effect on body weight
is not consistent among published papers (Forbes and
Injidi, 1979; Bermudez et al., 1983; Injidi and Forbes,
1983; Phetteplace and Nockels, 1985; Clark and Classen,
1995).
as long as this period does not extend for more than
about 12 h (Savory, 1979). Therefore, it is assumed that
feed intake, as well as growth, are maximal for broilers
that are reared in (nearly) continuous illumination.
However, several studies showed that alternative light-
ing schedules, such as increasing or intermittent lighting
schedules, improve body weight and feed conversion,
and reduce leg problems and mortality (Ketelaars et al.,
1986; Classen et al., 1991; Blair et al., 1993; Clarke et al.,
1993; Buyse et al., 1994a,b; Buyse et al., 1996). Neverthe-
less, published papers concerning intermittent lighting
schedules on broiler performance are inconsistent.
It is likely that broiler chickens that are exposed to
continuous lighting will be severely deficient in serum
melatonin. It is also likely that improved performance
and health in broilers exposed to intermittent lighting

Abbreviation Key: D = dark; ERTOT = total energy retention; ERp =

Received for publication March 15, 1998. energy retained as protein; ERf = energy retained as fat; GE = gross
Accepted for publication October 6, 1998. energy; HTOT = total heat production; HACT = activity-related heat
1To whom correspondence should be addressed: production; HNACT = nonactivity-related heat production; L = light; ME/
Johan.Schrama@GenR.VH.WAU.NL GE = metabolizability of GE;MEm = ME required for maintenance.

223
224 APELDOORN ET AL.

are at least partly due to melatonin (Classen et al., 1991). TABLE 1. Composition of the diet1
However, very few studies have been done to examine
Ingredient Percentage
the effect of melatonin, in dependency of lighting
schedule, on performance. The current experiment was Wheat (11.9% CP) 35.0
Soybean meal (45.4% CP) 15.0
conducted to gain an insight into the ways that Maize (8.7% CP) 10.0
melatonin and lighting schedule influence energy Peas (20.7% CP) 10.0
metabolism and performance of broilers. Soybean, toasted 5.0
Sunflower meal (34% CP) 5.0
Meat and bone meal (58.4% CP) 5.0
MATERIALS AND METHODS Tapioca (65% starch)
Lard
3.94
3.0
Soybean oil/maize oil 3.1
Feather meal (hydrolyzed) 1.5
Animals, Housing, and Vitamin and mineral mix2 1.0
Experimental Design NaCl 0.26
Limestone (CaCO3) 0.88
Monocalcium phosphate 0.92
In total, eight groups of female Ross broiler chickens L-Lysine HCl 0.20
were used. Each group consisted of six chickens. Each DL-Methionine 0.20
group was considered an experimental replicate and was Calculated contents
randomly assigned to an experimental treatment accord- CP 22.0
ing to a 2 · 2 factorial design. The first factor was the ME, kcal/kg
Ca
2,866
0.86
lighting schedule; either a near-continuous lighting P 0.71
schedule [23 light (L):1 dark (D); lights on from 0100 to Lysine 1.254
Methionine + cystine 0.917
2400 h] or an intermittent lighting schedule (1L:3D; lights
1As-fed
on from 0100 to 0200, 0500 to 0600, 0900 to 1000, 1300 to basis.
2Provided the following amounts of vitamins and minerals per
1400, 1700 to 1800, and 2100 to 2200 h). The second factor
kilogram of complete diet: vitamin A, 10,000 IU; cholecalciferol, 2,000 IU;
was the addition of melatonin to the diet at a level of 0 or vitamin E, 15 mg; vitamin B12, 15 mg; vitamin K, 5 mg; riboflavin, 5 mg;
40 ppm. niacinamide, 40 mg; d-pantothenic acid, 12 mg; choline chloride, 500 mg;
The experiment lasted 20 d. This experimental period folic acid, 0.75 mg; biotin, 0.1 mg; Fe, 60.3 mg (FeSO4·7H2O); Zn, 34.1 mg
(ZnSO4·H2O); Mn, 63.1 mg (MnO2); Co, 0.21 mg (CoSO4·7H2O); Cu, 15.2
consisted of three consecutive balance periods (one of 6 d mg (CuSO4·7H2O); Se, 0.15 mg (Na2SeO3·5H2O); ethoxyquin, 100 mg.
followed by two of 7 d). At the start of the experiment
chickens were 21 d old and the mean BW was 0.601 kg and
ranged from 0.526 to 0.713 kg (SEM = 0.041). In the present
study, the long-term effect of melatonin and lighting
per group were measured during the three successive
schedule on energy metabolism was assessed in broilers
between 21 and 41 d of age. Therefore, the chickens were balance periods. Excreta and dust production were
already exposed to the experimental treatments from 1 to collected quantitatively per group and sampled for energy
21 d of age. From 1 to 21 d of age, the lighting schedule was and nitrogen analysis per balance period. Gross energy
according to the experimental protocol and chickens had (GE) values of feed and excreta were determined with
free access to one of the experimental diets (Table 1), and adiabatic bomb calometry and N contents by Kjeldahl. The
water. Prior to the experiment, chickens were housed in ME intake per group was derived from the energy
battery cages (length by width by height = 0.60 · 0.73 · contents of the feed and the excreta. Total heat production
0.40 m, one group per cage). (HTOT) for each group was determined every 9 min from
At the start of the experiment, each group was placed in the measurement of exchange of CO2 and O2 (Verstegen et
one of two identical, open circuit, indirect climatic al., 1987), and calculated according to the method of
respiration chambers (Verstegen et al., 1987). The respira- Romijn and Lokhorst (1961). Heat production was
tion chambers measured 1.0 · 0.8 · 0.97 m (length by measured throughout the experiment, excluding the days
width · height). Light intensity was 79 to 83 lx at chick on which the respiration chambers were opened for
level, provided by two light bulbs of 25 W per chamber. collecting excreta and dust (Days 0, 6, 13, and 20). Total
The temperature was kept in the thermoneutral zone at 22 energy retention (ERTOT) was calculated by subtracting
C, with the relative humidity maintained at 67.5 – 2.5%. HTOT from ME intake per balance period. The retention of
Air velocity was < 0.20 m/s. The chickens were allowed ad N was estimated from N in feed, in excreta, in aerial NH3,
libitum access to feed and water. The experiment was and in NH4+ of water that condensed on the heat
conducted in accordance with Dutch law regarding the exchanger. Energy retention as protein (ERp) was calcu-
use of experimental animals. lated as 5.7 · 6.25 · N retention, where 5.7 kcal/g is the
energetic value of protein. The energy retention as fat
Measurements (ERf) was derived as the difference of the ERTOT and the
ERp. The ME required for maintenance (MEm) was
Individual BW was measured at the start of the calculated as:
experiment period (Day 0) and on the final day of each
balance period (Days 6, 13, and 20). Energy and N balances MEm = ME – (ERp/0.54) – (ERf/0.74) [1]
 EFFECT OF MELATONIN AND LIGHTING SCHEDULE ON ENERGY METABOLISM 225
TABLE 2. Energy intake, heat production, and energy retention in broilers (during 21 to 41 d of age)
as affected by lighting schedule1 and melatonin

Lighting schedule (L)

23L:1D 1L:3D P Value
Variable Control Melatonin (M) Control Melatonin SEM L M L · M
Initial body weight, kg 0.659 0.581 0.550 0.573 0.041 0.231 0.540 0.291
Growth, kg/d 0.069 0.065 0.067 0.066 0.003 0.972 0.375 0.604
Feed intake, kg/d 0.134 0.121 0.121 0.120 0.006 0.319 0.266 0.361
Feed conversion 1.93 1.84 1.78 1.79 0.03 0.015 0.196 0.127
GE intake, kcal·kg–0.75·d–1 492.5 475.1 482.9 473.4 10.94 0.630 0.287 0.735
ME intake, kcal·kg–0.75·d–1 339.2 332.5 342.9 336.7 7.22 0.611 0.287 0.735
ME/GE 0.689 0.700 0.710 0.711 0.001 0.071 0.414 0.457
Heat production, kcal·kg–0.75·d–1
Total, HTOT 204.5 200.7 208.7 204.4 4.99 0.475 0.468 0.960
Activity-related, HACT 26.2 22.0 22.6 20.6 0.95 0.058 0.032 0.318
Nonactivity-related, HNACT 178.3 178.7 186.1 183.8 4.26 0.204 0.835 0.762
Retention, kcal·kg–0.75·d–1
Energy, ERTOT 134.7 131.8 134.2 132.4 6.65 0.998 0.733 0.933
Protein, ERp 67.6 67.9 69.3 67.4 1.20 0.653 0.540 0.407
Fat, ERf 67.1 63.8 64.9 65.0 6.20 0.932 0.806 0.797
ME for maintenance
(MEm, kcal·kg–0.75·d–1) 123.5 120.4 126.8 124.1 5.82 0.564 0.661 0.994
123L:1D: continuous lighting schedule (23 h light followed by 1 h darkness), 1L:3D: intermittent lighting schedule (1 h light followed by 3 h
darkness).

where 0.54 and 0.74 were the values used as the efficiency
of utilization of ME for protein and the fat retention,
respectively. Energy balance data, and the data on heat
production and energy retention were expressed in
kilocalories per metabolic kilogram per day.
Physical activity was continuously monitored by
Doppler-radar activity meters2 (Wenk and van Es, 1976;
Verstegen et al., 1987), but was recorded in the same
intervals as HTOT. Per group and per day, the
9-min data on HTOT were related to activity according to
the following equation:
HTOT:ij = m + Di + b1 · Xj + eij
where HTOT:ij = heat production during day period i and
9-min period j; m = overall mean; Di = fixed effect of day
period i (i = 1,2); X j = activity counts during
9-min period j; b1 = regression coefficient of heat
production on activity counts; eij = error term. Circadian
rhythms of heat production (Aschoff et al., 1974) can only
partially be explained by physical activity (van der Hel et
al., 1984; Henken et al., 1993; Schrama et al., 1994).
Therefore, two levels for the fixed effect of day period
were included in Equation [2]. The day was divided into a
day period (lights on) and a night period (lights off) (see
experimental design). Heat production related to activity
(HACT) was calculated for each 9-min period as follows:
where HACT:j = activity-related heat production during
9-min period j; Xj = activity counts of 9-min period j; b1 =
the estimated regression coefficient of HTOT on activity
from Equation [2]. The heat production not related to
physical activity (HNACT) was derived by subtracting
HACT from HTOT. As with HTOT, HACT and HNACT were
determined continuously every 9 min throughout the
experiment, except on days the chambers were opened.
Statistical Analyses
Mean values over the experiment of energy balance
[2] traits, growth, feed intake, and feed conversion were
tested for the effects of melatonin and lighting schedule
using a two-way ANOVA. To asses the effect of melatonin
and lighting schedule on heat production (HTOT, HACT,
and HNACT), the hourly means of HTOT, HACT, and
HNACT were calculated. Preliminary analyses showed
that heat production patterns during the day were
strongly affected by lighting schedule. Therefore the effect
of melatonin, on the hourly means of HTOT, HACT, and
HNACT was tested separately, within each lighting
schedule, using one-way ANOVA. All statistics were done
using SAS (1986). P < 0.10 was accepted for statistical
significance.
RESULTS AND DISCUSSION

HACT:j = b1 · Xj [3] Energy Balance

Data on energy intake, heat production, and energy
retention, as affected by lighting schedule and melatonin,
are summarized in Table 2. At the start of the experiment,
2Radar MD5, Suther, Vierpool, Amsterdam, The Netherlands. there were no differences (P > 0.10) in BW between
226 APELDOORN ET AL.
treatments. Energy balance traits, averaged over the
experiment (21 to 41 d of age), were not affected by the
addition of melatonin to the diet (Table 2), except for
HACT; supplementation of melatonin to the diet reduced
physical activity from 24.4 to 21.3 kcal·kg–0.75·d–1 (P <
0.05). Feed intake and feed conversion, averaged over the
20-d experimental period, were not affected by the
addition of melatonin (Table 2). However, when the data
were analyzed for each week of the experimental period
separately, supplementation of melatonin had (P < 0.10) a
lowering effect on both GE intake (480.6 vs 448.6
kcal·kg –0.75 ·d –1 ) and ME intake (334.1 vs 311.7
kcal·kg–0.75·d–1) during the 1st wk of the experimental
period (data not shown). Feed conversion was not affected
by melatonin during the 1st wk of the experimental period
(P > 0.10). However, during the last week of the
experimental period, supplementation of melatonin to the
diet resulted in a lower feed conversion (2.04 vs 1.95, P <
0.05).
Growth, feed intake, and GE intake were not affected
by lighting (P > 0.10, Table 2). Feed conversion was
influenced by lighting schedule (P < 0.05); feed conversion
was 1.79 for the intermittent lighting and 1.89 for the
continuous lighting (Table 2). The ME/GE was higher for
the intermittent lighting schedule (0.711 vs 0.695, P< 0.10,
Table 2). Activity-related heat production was 2.5
kcal·kg–0.75·d–1 lower (P < 0.10, Table 2) for the intermit-
tent lighting than for the continuous lighting; HACT,
expressed as a percentage of HTOT, was lower for the
intermittent lighting schedule (0.12 vs 0.10, P < 0.01, data
not shown). Lighting schedule had no effect on ER (133.3
kcal·kg–0.75·d–1), nor on ERp or ERf (P > 0.10, Table 2). No
significant interactions between melatonin and lighting
schedule were found on energy metabolism traits (Table
2).
Feed intake and feed conversion were not affected by
the supplementation of melatonin to the diet (Table 2).
This result is in contrast to reports of Forbes and Injidi
(1979), Bermudez et al. (1983), Injidi and Forbes (1983),
Phetteplace and Nockels (1985), Osei et al. (1989), and
Clark and Classen (1995). The absence of an effect in the
present study might be related to a change in housing
conditions at the start of the experiment. Clark and
Classen (1995) found that melatonin had a small effect on
weight gain and feed consumption only during the first 2
wk of their experiment. This short-term effect of melato-
nin is in agreement with our findings. During the last
week of the experiment, melatonin had a beneficial effect
on feed conversion.
The present study demonstrated that HACT was
reduced when melatonin was supplemented to the diet.
This reduced HACT might be related to the sleep-inducing
capacity of melatonin, as demonstrated by Forbes and
Injidi (1979) and Bermudez et al. (1983). The observed
decline in HACT might be one of the explanations for an
improved feed conversion often found in other studies
when melatonin was added to the diet (Phetteplace and
Nockels, 1985; Clark and Classen, 1995).
Intermittent lighting in this study improved feed
conversion. This finding is in agreement with those of
Cherry et al. (1978), Cave (1980), Malone et al. (1980),
Simmons (1982), Ketelaars et al. (1986), Buyse et al. (1994a),
and Buyse et al. (1996). The lower feed conversion in the
present study with intermittent lighting was related to
reduced feed intake and not to altered growth rate.
However, Cherry et al. (1978), Simmons (1982), Ketelaars
et al. (1986), and Buyse et al. (1994a) found lower feed
conversion together with a higher growth rate under
intermittent lighting.
The present study showed that the improved feed
conversion under intermittent lighting was related to a
higher ME/GE as well as a lower HACT. The difference in
metabolizability, in general, could be due to a difference in
digestion or a difference in energy losses with urine. The
latter is probably not affected in this study, because
energy ERp was not affected by lighting schedule (Table
2). It is, however, unclear how lighting schedule might
affect digestibility. One explanation could be due to
greater waste of feed under the continuous lighting
schedule; however, in the present study, no feed spilling
was observed. The higher HACT in broilers exposed to
continuous lighting might be related to a lower en-
dogenous melatonin level compared to broilers exposed
to intermittent lighting.
In the present study, lighting schedule had no signifi-
cant effect on ERp or ERf. This result is in contrast to those
of Malone et al. (1980), Robbins et al. (1984), and Buyse et al.
(1994b), who reported less fat deposition on intermittent
light; however, Ketelaars et al. (1986) found that intermit-
tent lighting increased fat deposition. The reason for these
contradictions is not clear.
Because of the light-dependency of the pineal gland
(Pang et al., 1996), a lighting schedule by melatonin
interaction could be expected, which Phetteplace and
Nockels (1985) observed for weight gain in an experiment
with cockerels. However, in the present experiment, as in
the report of Osei et al. (1989), no significant interactions
were found.
Daily Heat Production Patterns
Lighting schedule had a great impact on the heat
production during the day. Almost every hour, lighting
schedule had a significant effect (P < 0.05) on HTOT,
HNACT, and HACT. There were also a number of
interactions between lighting schedule and melatonin
(0000 h, P < 0.01; 0400 h, P < 0.10; 1100 h, P < 0.05; 1500 h, P
< 0.05, and 2300 h, P < 0.10) on HACT during the day (data
not shown). To show the effect of melatonin on daily heat
production patterns, lighting schedule was eliminated as a
main effect by separating the two lighting schedules. The
effects of melatonin on HTOT, HNACT, and HACT during
the day and for each lighting schedule are depicted in
Figure 1 (continuous light) and Figure 2 (intermittent
light).
 EFFECT OF MELATONIN AND LIGHTING SCHEDULE ON ENERGY METABOLISM
On both lighting schedules, a distinct peak in heat
production occurred just after every dark period.
However, the difference in total heat production between
dark and light periods was much greater for the
intermittent lighting schedule (approximately 1.97
kcal·kg–0.75·h–1), than for the continuous lighting schedule
(approximately 1.19 kcal·kg–0.75·h–1). The peaks in HTOT
for the intermittent lighting schedule were partially
related to the increased activity, which was indicated by
the peaks in HACT. Overall, the daily patterns in heat
production were flatter for the continuous lighting
schedule and more variable for the intermittent lighting
schedule.
FIGURE 1. Hourly means of total heat production (HTOT), activity-
related heat production (HACT), and non-activity-related heat produc-
tion (HNACT) of broilers (21 to 41 d of age), exposed to nearly continuous
light, fed a diet without (o) or with ( ) melatonin (# P < 0.10; * P < 0.05;
** P < 0.01).
227
Melatonin had no effect on the daily pattern in HTOT or
HNACT (P > 0.10) within each lighting schedule. Melatonin
had a lowering effect on HACT during the day (Figure 1
and 2); however, the lowering effect of melatonin on HACT
during the day was much more obvious in the continuous
lighting schedule.
Mean data on heat production, during dark (lights off)
periods and during light (lights on) periods, as affected by
lighting schedule and melatonin, are summarized in Table
3. Melatonin reduced HACT, especially during light
periods (HACT was 0.17 kcal·kg–0.75·h–1 lower when
melatonin was supplemented to the diet, P < 0.10, Table 3).
During dark periods there were only slight differences in
FIGURE 2. Hourly means of total heat production (HTOT), activity-
related heat production (HACT), and non-activity-related heat produc-
tion (HNACT) of broilers (21 to 41 d of age), exposed to intermittent light,
fed a diet without ( ) or with ( ) melatonin (# P < 0.10; * P < 0.05).
228 APELDOORN ET AL.
TABLE 3. Heat production in broilers (during 21 to 41 d of age) during the dark and
light periods of the day, as affected by lighting schedule1 and melatonin

Lighting schedule (L)

23L:1D 1L:3D P Value
Variable Control Melatonin (M) Control Melatonin SEM L M L · M
Mean heat production during dark periods, kcal·kg–0.75·h–1
Total (HTOT) 7.40 7.24 8.19 8.03 0.445 0.151 0.746 0.997
Activity-related (HACT) 0.46 0.30 0.57 0.51 0.052 0.035 0.108 0.347
Nonactivity-related (HNACT) 6.94 6.95 7.62 7.52 0.418 0.208 0.915 0.898
Mean heat production during light periods, kcal·kg–0.75·h–1
Total (HTOT) 8.58 8.44 10.15 10.01 0.279 0.005 0.646 0.996
Activity-related (HACT) 1.12 0.96 2.10 1.92 0.069 0.0001 0.075 0.885
Nonactivity-related (HNACT) 7.46 7.47 8.05 8.09 0.224 0.054 0.914 0.959
123L:1D: continuous lighting schedule (23 h light followed by 1 h darkness), 1L:3D: intermittent lighting schedule (1 h light followed by 3 h
darkness).

HACT due to melatonin (0.11 kcal·kg–0.75·h–1, P > 0.10,
Table 3). During light periods, lighting schedule had a
strong effect on heat production; HTOT, HNACT, and HACT
were higher for the intermittent lighting schedule (1.57
kcal·kg–0.75·h–1 [P < 0.01], 0.61 kcal·kg–0.75·h–1 [P < 0.10],
and 0.97 kcal·kg–0.75·h–1 [P < 0.001], respectively), com-
pared to continuous lighting schedule (Table 3). During
dark periods, lighting schedule had only an effect on
HACT (0.54 kcal·kg–0.75·h–1 for the intermittent lighting
schedule vs. 0.38 kcal·kg–0.75–h–1 for the continuous
lighting schedule, P < 0.05, Table 3). For neither light nor
dark periods, an interaction between lighting schedule
and melatonin was found (Table 3).
The reduced activity over the whole day at the
intermittent lighting schedule (Table 2) seems to be
contradictory to the higher activity during both the dark
and light periods under intermittent lighting (Table 3);
however, the lower HACT over the whole day under
intermittent light is due to the difference in total hours of
darkness during the day between the lighting schedules
(18 vs 1 h). The peaks in HTOT during the day for the
intermittent lighting schedule are not only caused by an
increased activity level, but also by an increase in HNACT
(Figure 2). After this peaks, HNACT decreased more slowly
than HTOT and HACT (Figure 2), which is probably due to
high feed intake during the light period, followed by a
period of digestion.
Melatonin had an effect on HACT during the day. This
effect was much more obvious for the continuous lighting
schedule than for the intermittent lighting schedule
(Figures 1 and 2). The reason for this could be that
chickens that are reared under a continuous lighting
schedule, and therefore are severely deficient in melato-
nin, are more sensitive to exogenous melatonin than
chickens that are exposed to intermittent light.
In the present study, the long term effect of melatonin
and lighting schedule on energy metabolism was assessed
in broilers between 21 and 41 d of age. The chickens had
been already exposed to the experimental treatments from
1 to 21 d of age, in order to have the chicks in a steady state
regarding their response to the treatments during the
experiment (21 to 41 d of age). The present study
demonstrated that a reduced energy expenditure for
physical activity, caused by the supplementation of
melatonin to the diet, might be a reason for the often
observed improvement in feed conversion. Furthermore,
the present study showed that lighting schedule also
influenced feed conversion. Improvement in feed conver-
sion with intermittent compared to continuous lighting
schedule was due to a ME/GE and a lower energy
expenditure for physical activity.
ACKNOWLEDGMENTS
The assistance of M.J.W. Heetkamp and J. M. van der
Linden is gratefully acknowledged.
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