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Effect of hatching time on poultry

behaviour: the impact of incubation
environment during the last days of
embryonic development
On
Humboldt-University of Berlin
Institute of Biology, Perinatal Adaptation
Philippstrasse 13, 10115 Berlin, Germany

Introduction

Impact of environmental factors during

embryonic development

Barbara Tzschentke

Stage of development of physiological

functions during hatching time
n precocial birds including poultry, physiological
mechanisms are well developed during hatching
time, which represents the last days of embryonic
development (roughly from plateau phase
until hatching). Peripheral and central nervous
mechanisms, for instance, of the cardio-vascular-,
respiratory-, and thermoregulatory system show
characteristics, which are not very different from the
post-hatching one. Most of the regulatory systems
undergo a dramatic qualitative change from an open
loop system without feed back mechanisms into a
closed control system with feed back mechanisms
(Tzschentke and Plagemann, 2006 Tzschentke,
2007; Tzschentke and Rumpf, 2011). In chicken
embryos the hypothalamic-pitatury-thyroidal axis,
for instance, is functional on embryonic day (E)
19 (Muchow et al., 2005). In this species, also the
thermoregulatory system shows feed back reactions
on acute temperature influences during the last
days of embryonic development (Tzschentke and
Plagemann, 2006 Tzschentke, 2007; Tzschentke
and Rumpf, 2011). Sensory modalities in avian
embryos develop in the following sequence: (1)
nonvisual photic sensitivity, (2) tactile sensitivity,
(3) vestibular sensitivity, (4) proprioception, (5)
audition and (6) vision (after Gottlieb, 1968 and
Vince, 1974). Hence, the sensory capacity for
hearing and vision in bird embryos develops lastly,
but both are functional during hatching time.

During the last days before hatching the embryos

may also activate behavioural mechanisms. The
embryos are able to produce calls (Tuculescu and
Griswold, 1983; Rumpf and Nichelmann, 1993;
Brua, 2002). Embryonic motility is an important
component of development and may be a precursor
of post-hatching motor behaviour (Nechaeva et al.,
2010). In chicken embryos first movements occur
on E3.5, increase during development, and during
final incubation possibly transforms into targeted
hatching behaviour (Hamburger and Balaban, 1963;
Hamburger et al., 1965; Oppenheim, 1966; Bradley,
1999; Bekoff, 1992; 2001; Nechaeva et al., 2010).
Finally, the embryos have all prerequisites to react on
environmental influences on physiological, neuroendocrine and with limits on behavioural level (e.g.
acoustic communication, motility).

The major impact of prenatal and early postnatal

environmental conditions for later development and
performance in poultry is more and more accepted.
Especially during critical periods the effect of the
actual environment is long-lasting and may be
even passed on the succeeding generations in an
epigenetic fashion (Plagemann, 2004; Tzschentke
and Plagemann, 2006). A critical period is a strict
time window during which a certain experience
necessarily must occur to enable development
to proceed normally and permanently alters
performance (Baily et al., 2001; Hensch, 2005). But
it has to be considered that even within a species
different physiological functions and mechanisms
of the respective function may have different
critical periods, which may arise in an earlier
stage of embryonic development, during hatching
time or the early post-hatching development or
be overlapping (Harwerth et al., 1986; Tzschentke
and Plagemann, 2006). In this context imprinting
of body functions, which mostly occurs during
perinatal critical developmental periods, is a basic
mechanism and obviously a fundamental process
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Barbara Tzschentke

of life. In his classical studies on newly hatched

goslings, Konrad Lorenz analysed the development
of social binding, applying the term imprinting
to describe this process (Lorenz, 1935). One of his
major hypotheses was that imprinting occurs during
limited and severely restricted critical periods
in early life. Later, Gnter Drner, a pioneering
developmental neuroendocrinologist, developed
a general origination concept of the epigenetic
perinatal programming of the lifetime function of
fundamental regulatory systems (Drner, 1974,
1975). Hormones as well as neurotransmitters
and cytotokines (as immune cell hormones) play
a central role in this concept. They act as carriers
of environmental information to the genome.
Ultimately, they too are acting as epigenetic
factors with long-lasting effects particularly during
critical developmental periods (Fig. 1). As an
example, a critical period in the development of
regulatory systems is the development of feed back
mechanisms. In chick and Muscovy duck embryos
first feed back control of the thermoregulatory
system, for instance, was found during hatching
time (Tzschentke and Plagemann, 2006).

a later developmental stage. For instance, shortterm alterations in incubation temperature did not
change embryonic motor activity in an early stage
of incubation (E5 E14; Oppenheim and Levin,
1975; Nechaeva and Turpaev, 1991). But, a shortterm temperature increase or decrease modified
embryonic motility in older embryos (E15 E20;
Oppenheim and Levin, 1975). Similar age dependent
influence on motility was also found under hypoxia
(Nechaeva et al., 2010; Nechaeva, 2011). Whether
such short-term environmental influences on motor
activity during the last third of incubation have
also an effect on post-hatching motor behaviour
needs further investigations. Finally, there are
two possibilities regarding long-lasting effects of
environmental factors during incubation on later
development including behaviour. On one hand,
the incubation environment is a basis for perinatal
mal programming, which besides metabolic
disorders and cardiovascular diseases also may
cause behavioural disorders in birds during later life
(Schwabl, 1996, 1997; Bock et al., 2005). On the
other hand, knowledge and better understanding
of the imprinting mechanisms might be specifically
used to induce long-term
adaptation of an organism,
Imprinting of physiological conrol systems
for instance, to the postnatal
Developmental trajectory
climatic conditions (epigenetic
critical
temperature
adaptation;
period
Nichelmann et al., 1994, 1999;
Tzschentke and Basta, 2002),
which includes also behavioural
environmental influences
thermoregulation.
changes in hormone
concentration transmitters/
neuropeptides cytokines

developing embryo/fetus
pre-programmed by
genetics instructions

longlasting modification of
the pre-determined adult
phenotype via changes in
gene expression

Figure 1 - Imprinting of physiological control systems by environmental

factors during critical periods of early development (modified from
Tzschentke and Plagemann, 2006).

Importance of prenatal environmental

factors on behaviour
Behaviour is an interaction between the organism
(genes and physiological status/ homeostasis)
and the environment (Tembrock, 1978). Prenatal
environment may immediately influence the
embryonic behaviour und may also induce longlasting behavioural alterations after hatching.
Clear modifications of embryonic behaviour by
environmental influences were mostly found in

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Altogether, the long-lasting

influence of incubation factors
on post-hatching behaviour in
poultry is not much investigated.
The
following
presentation
is focused on the influence
of acoustic signals, light and
temperature during hatching time
on poultry behaviour.

Acoustic signals,
hatching synchronization and
post-hatching behaviour
Development
transmission

of

hearing,

sound

Jones et al., (2006) proposed two periods of

ontogeny of the hearing system in chick embryos.
The first period is the so-called prehearing period

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(about E12-E16). It is the period of endogenous

cochlea (inner ear) signalling. It is important for
the normal development of the central binaural
processing pathway. In the second period (about
E16-E19) the cochlea begins to detect and encode
sound. In the late chick embryos between E18 and E
21 auditory structures and functions show an adultlike development in most respects (for details, see
Rubel and Parks, 1988; Manley et al., 1991; Jones
and Jones 1995a,b; Jones et al., 2006). Also in
other poultry species exogenous acoustic signals are
detectable at the beginning of the plateau phase of
metabolism. In Muscovy ducks with an incubation
time of 33 days, for instance, the acoustic-sensorycardiac axis is functional from embryonic day 27 and
heart rate responses to acoustic stimulation were
observed (Hchel et al., 2002).

Sound production
(1) Vocalizations and its relevance for
acoustic communication
Beginning with penetrating the inner egg

Figure 2 - Prenatal vocal activity of single (white

columns) and paired embryos (black columns).
Each column represents the mean value of three
embryos (significant different, p<0.01 three days
and one day before hatching, not significant
different 2 days before hatching) from Rumpf
and Tzschentke (2010), according to Rumpf and
Nichelmann (1993).

Barbara Tzschentke

Whether an embryo can hear only its neighbour

or all the embryos in the clutch depends on
sound transmission between the eggs. Rumpf
and Tzschentke (2010) described measurements
of sound transmission in Muscovy duck embryos
at E32 before the outer eggshell was pipped (2-3
days before regular hatching) using an artificial
sound emitter (broad band noise signal in the range
of 50-5000 Hz, in the range of the embryos own
sound production frequencies). The artificial sound
was presented to the first egg and was measured
on each of the 9 following eggs in 10 lengthwise
or crosswise positioned eggs. Summarizing all
measured frequencies, sound absorption in
crosswise positioned eggs was by 5-10 dB greater
compared with that of lengthwise positioned
eggs (Lauch, 1989). Above 3 kHz no frequency
dependence of sound absorption was measured.
As sound absorption from the first to the 6th egg
amounted to 10 dB it can be assumed that embryos
have mutual acoustic contact with all the other
embryos in a clutch. Finally, sound transmission is
based on a sound-conducting medium. In a clutch,
sound is conducted via the eggshell, lengthwise and
crosswise. In industrial or commercial incubators
a metal grid or a metal setter tray may serve as a
sound-conducting medium. Even in the case when
embryos are connected via a metal grid, a nearly
perfect sound conductance was measured (Lauch,
1989). Finally, we suppose that most sound within
the prenatal period is bone-conducted and not airconducted (Rumpf and Tzschentke, 2010).

membrane a species-specific sound repertoire

develops gradually. During the final prenatal
period vocalization activity rises that as is especially
evident in those bird species having a long period
of prenatal vocal production. Fig. 2 shows the
vocalization activity of the Muscovy duck (Rumpf
and Nichelmann, 1993). The number of vocalization
increased within the 3 days before hatching as
studied in single and in paired embryos. Similar
results of increasing vocalization activity are
published for naturally incubated domestic chick
embryos (Tuculescu and Griswold, 1983).

Higher acoustic activity might be regarded as

non-specific mutual stimulation which is supported
generally by the birds` own vocalization or other
stimulation within the perinatal period. Vice versa,
a disruption of mutual vocal communication in
the pre-hatching period led to lower acoustic
postnatal activity and also damaged postnatal
acoustic communication (Lauch, 1989). Pekin
ducks, for instance, need embryonic experiences
of a wide range of repetition rates of their contactcontentment call to develop a preference for
maternal calls (Gottlieb, 1985). Also recent studies
in domestic chicken (Kauser et al., 2011) and the
bobwhite quail (Harshaw and Lickliter, 2011) have
shown that prenatal auditory stimulation with either
species-specific or complex rhythmic music sounds
bias postnatal responsiveness to social stimuli,
facilitates spatial learning and influences memory.
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Up to now it is not known if and how the embryos
receive the acoustic signals of the breeding parent.
But it is likely that the breeding parent receives the
acoustic signals of the embryos. Vocalizations of
the embryos may affect the behaviour of the parent
(Tschanz,1968; Lauch, 1989) for instance eggturning, nest building or the amount of time parents
spend on the nest.

Barbara Tzschentke

Embryonic vocalizations might also serve as caresoliciting signals concerning temperature regulation.
Cold induced vocalization (distress calls; Fig. 3) may
help to restore normal incubation temperature
(Evans, 1989; 1990; Brua et al., 1996; Nichelmann
and Tzschentke, 1997). Under natural conditions it
may be a signal of the offsprings need for warmth
from the incubating parents. For the full review to
this topic see Rumpf and Tzschentke (2010).

Figure 3 - Distress call rate of Muscovy duck

embryos before (min -5 to 1) and after a
cold stimulus of 20 to 22 C (min1 to 19) from
Nichelmann and Tzschentke (1997).
(2) Clicking sounds and
its relevance for hatching
synchronization
Accompanying with the
development of breathing the
embryos begin to regularly
produce so-called clicking
sounds (Vince and Salter, 1967),
clicks or clicking noises. Prenatal
clicks are accompanying noises
of respiration and not a real
vocalization, controlled by the
syrinx. In the Muscovy duck first
clicking sounds were observed
once the inner eggshell was
penetrated (Lauch et al., 1988).
The clicking rate (number of
clicks per minute) corresponded

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with the audible respiration frequency. The

development of the clicking rate of one Muscovy
duck embryo from the first clicks to the hatching is
shown in Fig. 4.
Investigations on the development of prenatal
clicking rate are only available for Muscovy ducks
(Lauch et al., 1988; Lauch, 1989) and quails (Vince
and Salter, 1967).
Since the investigations by Vince on
quails (1964a,b) it is known that the acoustic
communication by clicking sounds is an essential
factor for hatching synchronization. Vince
discovered that the development and hatching
time of quail embryos could be accelerated or
decelerated, by clicking sounds (Vince,1966a,b;
Vince 1968a,b; Vince and Cheng, 1970; Vince et
al., 1984). In the Muscovy duck synchronization
starts, when the second embryo within a clutch
begins to click. Within certain, species-specific
range embryos adapt their clicking rates to those
of others. The result is a common rhythm. Muscovy
duck embryos need about a minute to achieve this
common rhythm (or beat). Normally, embryos try to
synchronize their rhythms 1:1. Then the resulting
clicking rate is the same in both or in all of the
embryos. If the lung respiration is not sufficiently
developed and the embryo cannot follow the
given rhythm over a long time, the embryo initially
follows the clicking beat irregularly (with breaks).
Towards hatching, embryos click more and more
regularly. Usually, the Muscovy duck embryos in a
clutch were able to synchronize their clicking rates
(Fig. 5 A, B) if the surrounding sound level does
not exceed about 80 dB as measured in industrial
incubators (Nichelmann et al., 1990). However, if

Figure 4 - Prenatal clicking rate of a single embryo (measured every 30

minutes); method described by Lauch et al., (1988). A- External pipping;
B- Cutting (pipping while rotating); C- Hatch. Figure from Rumpf and
Tzschentke (2010), according to Lauch et al., (1988).

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the embryos had a big developmental difference to

the next sibling, this embryo hatched considerably
earlier or later (one example in Fig. 5 B). It can
be hypothesized that small differences between
the embryos as caused by, e.g., temperature can
be compensated. For hatching synchronization
in the Muscovy duck the mutual acoustic contact
between the embryos was necessary during the
whole prenatal clicking period (Lauch, 1989). As
already given before, clicking sound communication
is different from communication by vocalization
(Rumpf and Tzschentke, 2010). In laboratory
experiments using white noise in combination with
low pass and high pass filters respectively, it was
possible to eliminate clicking sound communication
and allow communication by vocalization on the one
hand, and to allow clicking sound communication
but exclude communication by vocalization on the
other hand (Lauch et al., 1988). Clicking rates and
hatching synchronization were measured under
different sound levels (steps of 5 dB). Over a sound
level of 80 dB (white noise low pass filter) clicking
sound communication was disturbed and Muscovy
duck embryos hatched asynchronously (Fig. 5 C, D).
Hatching interval under desynchronizing conditions

(80 dB) was significantly, on average 4 times longer,

than under synchronizing conditions (Lauch et al.,
1988). Commercial incubation conditions seem
to be more similar to desynchronized laboratory
conditions (Nichelmann et al., 1990).
Summarizing all findings there is evidence
that prenatal clicking sounds in contrast to other
acoustic signals play an important role for hatching
synchronization at least in quails and Muscovy
ducks. For the full review to this topic see Rumpf
and Tzschentke (2010).

Light, brain development

and post-hatching behaviour
In precocial birds light stimulation during
embryogenesis is very important for normal brain
development. Especially the specialization of the
avian hemispheres (brain lateralization) depends on
light exposure of the embryo during hatching time.
The influence of light during the last days before
hatching on the brain lateralization is related to
Barbara Tzschentke

Figure 5 - Synchronization of prenatal clicking rates of 8 (A) and 12 (B) Muscovy duck embryos under a
sound pressure level in the laboratory incubator of 50 dB and desynchronization of prenatal clicking rates
of 7 (C) and 12 (D) Muscovy duck embryos under a sound pressure level in the laboratory incubator of 80
dB, from Rumpf and Tzschentke (2010), according to Lauch 1989). (broken line=eggshell pipped, mutual
eggshell contact , white noise low-pass-filter<2kHz ).
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XXIV
the body position of the embryo in the egg; the
left eye is covered by the body and the right eye
is near the light-transmissive egg shell, which was
already observed in chick embryos by Kuo (1932).
Lateralization of the brain is fundamental for
the post-hatching behaviour (Rogers, 2011). For
investigations of brain lateralization and behaviour,
it is important to know, that the visual information
from each eye is processed to the opposite
hemisphere (left eye to right hemisphere, right eye
to left hemisphere).

Barbara Tzschentke

The left eye-right hemisphere preferentially

controls behaviour, which is important under
emergency or stressful conditions (e.g. attention to
novel objects, predators, fear response). In chicken
the right hemisphere is also involved in copulation
behaviour (Roger et al., 1985) and different aspects
of social learning (Daisley et al., 2009), which
includes, e.g., the maintenance of social hierarchy
(Hogue et al., 1996) and recognition of face-like
stimuli (Daisley et al., 2009).
The main functions, which are preferentially
controlled by the right eye-left hemisphere, are learnt
and routine behaviour under nonstressful situations.
The ability to learn to distinguish between different
objects (e.g. between food grains and pebbles,
Mench and Andrew, 1986), for instance, is a feature
of the left hemisphere. Another example, which is
controlled by the left hemisphere, is the ability to
focus attention on training cues in an experiment
(Roger et al., 2008).
In dark incubated chicken and other bird species
no or weakly developed anatomical and functional
brain asymmetries were found (Dharmaretnam and
Rogers, 2005; Rogers, 2011). During the first posthatching weeks limitations or losses in behavioural
abilities (e.g. learning, social behaviour) are typical.
Whether these behavioural changes are persistent
needs further validation. In comparison with light
incubated chicks, dark incubated chicks develop,
for instance, less stable social hierarchies (Roger and
Workman, 1989). They are unable to discriminate
between different conditions (Chiandetti and
Vallortigara, 2009) and produce more distress calls,
followed by a higher fearfulness (Dharmaretnam
and Rogers, 2005). Finally, dark incubated birds
are more vulnerable to post-hatching stress. To
reduce this negative effect, bird embryos need light
exposure, which can be an important contribution
to life-long welfare (Rogers, 2011).

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Incubation temperature
and behaviour
Investigations on the influence of incubation
temperature on post-hatching behaviour in birds
are rare. In Wood ducks (Aix sponsa) it was shown
that already mild changes in incubation temperature
have a significant influence on locomotion in the
offsprings (Hopkins et al., 2011). In Wood ducks the
natural incubation temperature varies between 34.8
and 37.8 C, so that also the incubation length differs
from 30 to 37 days (Hepp et al., 2006). Locomotor
activity of Wood ducks incubated at three different
ecologically relevant temperatures (35 C, 35.9
C and 37 C) was tested at 15 and 20 days posthatching. Ducklings, which were incubated at the
lowest temperature, showed a significantly reduced
aquatic swim velocity compared with the ducklings,
which were incubated at both higher temperatures
(Hopkins et al., 2011). The authors suggest that the
lower aquatic swim velocity at the lowest incubation
temperature is a sign for reduced fitness, because
locomotor activity might be important for survival
under natural conditions.
In poultry chronic and short-term changes in
incubation temperature during the last days until
hatch have different effects on the post-hatching
performance and adaptability to the environment
(Tzschentke and Halle, 2009; Halle and Tzschentke,
2011). The impact of these changes in incubation
temperature on the post-hatching behaviour is not
so much investigated. Some examples are given in
the next paragraphs.

Epigenetic temperature adaptation and

post-hatching thermoregulatory behaviour
Birds
and
mammals
attain
effective
thermoregulation by physiological as well as
behavioural mechanisms (Schmidt, 1984). Precocial
birds are able to use thermoregulatory behaviour, like
the innate ability for temperature preference, shortly
after hatching. During the first days post-hatching
behavioural temperature regulation is essential
to maintain a stable body temperature, because
the physiological mechanisms are not completely
developed. Further, behavioural thermoregulation
is not so much energy consuming like physiological
thermoregulation. Finally, to regulate body
temperature by behavioural mechanisms is an
effective energy saving mechanism. During the
first 10 days of life we found for different poultry
species, for instance, a close relationship between
the preferred ambient temperature for resting

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behaviour and thermoneutral temperature (Fig.

6). Resting under thermoneutral condititons is
an effective energy saving mechanism. In this
way, high-energy costly thermoregulatory heat
production is to a large degree not necessary to
keep body temperature constant (Tzschentke, 2003;
Tzschentke and Nichelmann, 2003).

At the 10th day post-hatching cold experience

during the last days until hatching elevated the total
neuronal hypothalamic warm-sensitivity through
an increased proportion of warm-sensitive neurons
and a reduced proportion of cold-sensitive neurons
in comparison with the control group. Exposure
to the warmer temperature during the prenatal
period induced the opposite effect. This change
in neuronal hypothalamic thermosensitivity after
exposure to various incubation temperatures may
be caused through a down or upward shift in the
range in which the respective neurons are sensitive
to temperature changes.

Figure 6 - Preferred ambient temperature (PT)

for resting behaviour (R) and thermo-neutral
temperature (TNT) in 9 days old turkeys (from
Tzschentke & Nichelmann, 2003).

These findings were supported by our study on

the influence of prenatal temperature experiences
on neuronal activity of the thermoregulatory centre
in the hypothalamus (Tzschentke and Basta, 2002).

Figure 7 - Influence of incubation temperature (34.5

C, 37.5 C) on preferred ambient temperature in
1-, 5- and 10-d-old Muscovy ducklings (n=15 in
each experimental group).

Barbara Tzschentke

Temperature experiences during hatching-time

significantly influenced postnatal temperature
preference. We carried out experiments in Muscovy
ducks and turkey, which were chronically incubated
at lower or higher temperatures than usual (37.5 C)
during the last days until hatching. Preferred ambient
temperature was determined in a temperature
gradient tunnel temperated between 10 and 45 C.
During this period, every day the chosen ambient
temperatures were observed for 9 hours every 10
min (Tzschentke and Nichelmann, 1999). With
increasing age Muscovy ducklings incubated at a
low temperature (34.5 C) preferred a significant
lower temperature than birds incubated at the
normal incubation temperature (37.5 C) during the
first 10 days post-hatching (Fig. 7). This supports
the hypothesis that avian prenatal cold experience
leads to a downward shift of the thermoregulatory
set-point (Tzschentke and Nichelmann, 1999). On
the other hand, the preferred ambient temperature
in 1- to 10-day-old turkeys (Fig. 8) is higher after a
prenatal heat load (38.5 C) than in birds incubated
under normal temperature (37.5 C). This indicates
an elevation of the thermoregulatory set-point after
prenatal heat load (Tzschentke and Nichelmann,
1999).

Figure 8 - Influence of incubation temperature

(IT: 38.5C, 37.5C) on preferred ambient
temperature in 1- to 10-days-old turkeys (n=15
in each experimental group) from Tzschentke and
Nichelmann (1997).
In the last days until hatching, body temperature
of chicken and Muscovy duck embryos was strong
influenced by chronic changes in incubation
temperature (Fig. 9; Loh et al., 2004). We assume
a critical period in the development of the
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normal incubated group (Halle and

Tzschentke, 2011).

Barbara Tzschentke

Prenatal temperature training,

robustness and behaviour
In contrast to chronic warm
incubation,
short-term
warm
stimulation (temperature training)
during hatching time induces
different effects on post-hatching
performance
under
standard
growing conditions (Tzschentke and
Halle, 2009; Halle and Tzschentke,
2011). In comparison to the
usual
incubation
temperature,
chronic warm incubation (1 C
over standard) had no effects on
hatchability and later performance.
But, short-term warm stimulation
Figure 9 - Temperature of allantoic fluid as measure for embryonic improves hatchability, feed intake,
body temperature in warm (ww, 38.5 C), cold (cc, 34.5 C) and feed conversion, and body weight
normal (nn, 37.5 C) incubated chicken embryos on different days of at slaughter, especially in male
incubation (* and o are extreme values). For each incubation group, broilers (Tzschentke and Halle,
the first letter indicates incubation temperature during the last 2009). Further, in this study the ratio
days until hatching and the second letter refers to the temperature of hatched female to male chicks
conditions during 1h of measurement (in this experiment similar with shifted in favour of the male chicks.
the respective incubation temperature) from Loh et al., (2004).
During temperature stimulation
the embryos show a decrease in
heat production and, finally, a lower embryonic
thermoregulatory system at the end of the embryonic
temperature befor hatching compared with the
development. Therefore the lower embryonic
non-stimulated group. The idea is that shorttemperatures under cold incubation condition and
term temperature training during the last days of
the higher one under warm incubation condition are
embryonic development might improve robustness
prerequisites for changes in the thermoregulatory
of the broiler chickens. Thus, changes in behaviour
set-point.
during later life might result. In our first study on
the influence of short-term warm stimulation on
The
above-described
changes
in
the
post-hatching locomotion we found no difference
thermoregulatory behaviour were observed during
between the warm-stimulated and the normal
the first 10 days post-hatching. We assume that
these changes are persistent, because we found
incubated group. However, this topic is still under
long-lasting changes in different physiological
investigation.
parameters.
Heat
induced
c-fos-expression
of hypothalamic neurons, for instance, was
significantly different between chronic warm- and
cold-incubated chickens even after 8 weeks posthatching (Janke and Tzschentke, 2010). Warm
For optimal development poultry embryos need
growing conditions (32 C) induce decrease in feed
environmental stimulation during the last days of
incubation. During this time-window physiological
intake and body weight. We found that under this
growing condition chronic warm incubated male
mechanisms are well developed. Its further
broiler chicks had a much lower feed consumption
maturation will be improved by environmental
than the normal incubated group. Moreover, the
inputs, like temperature variations, light-dark
cycle, and acoustic signals. Light, for instance is
final body weight (d 35) in this group was significant
lower, too. The stronger reduction of feed intake
very important for the normal brain development,
especially the brain lateralization. Besides
in chronic warm incubated male chickens lead
to a stronger decrease in body heat production,
improvement of robustness and performance,
environmental influences could also have a long
which significantly minimized the warm induced
thermal strain of these birds in comparison to the
lasting influence on post-hatching behaviour. Dark

Conclusion

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incubated poultry embryos, for instance, show

reduced cognitive abilities.
Commercial incubation programmes should
include environmental stimulation, with temperature
and light as the foremost and more silent environment
to realize mutual acoustic communication between
the embryos. Such conditions are near to the nature
and will address also the animal welfare aspect.

References
BAILY, D.B., BRUER, J.T., SYMONS, F.J. and LICHTMAN,
J.W. (2001) Critical thinking about critical periods. P H
Brookes Publishing Co.
BEKOFF, A. (1992) Neuroethological approaches to the
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