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Aufzucht weiblicher Kälber und Jungrinder
in landwirtschaftlichen Unternehmen
Teil III
Normales Verhalten von Kälbern
in der mutterlosen, intensiven Aufzucht
(Rearing of female calves and young cattle in agricultural enterprises
Part III
Normal behaviour of calves in motherless, intensive rearing)
Anke Schuldt
Regina Dinse
Neubrandenburg, Juni 2023
Reihe I, Band 11
Impressum
Herausgeber: Prof. Dr. Gerd Teschke
Rektor der Hochschule Neubrandenburg
für die Reihe
Alle Rechte, auch die der Übersetzung, des Nachdrucks, der Wiedergabe, der Ver-
vielfältigung und der Übernahme auf Datenträger, auch auszugsweise, sind der
Hochschule Neubrandenburg vorbehalten.
Schuldt, Anke; Dinse, Regina:
Aufzucht weiblicher Kälber und Jungrinder in landwirtschaftlichen Unternehmen;
Teil III: Normales Verhalten von Kälbern in der mutterlosen, intensiven Aufzucht.
Anke Schuldt, Regina Dinse. [Hochschule Neubrandenburg: Hrsg.: Rektor der Hoch-
schule Neubrandenburg]. – Neubrandenburg: Hochschule Neubrandenburg, Rektor,
2023 (Schriftenreihe der Hochschule Neubrandenburg: Reihe I, Fachbereich Agrarwirt-
schaft und Lebensmittelwissenschaften; Bd. 11.)
ISBN 978-3-941968-83-7
Für den Inhalt der Beiträge sind die Verfasser selbst verantwortlich.
Einbandgestaltung: G e l l e r
Hochschule Neubrandenburg
Druck: Steffen Media, Friedland
http://www.steffen-media.de
Gedruckt auf 100 %-Recyclingpapier
Einband-Papier chlorfrei gebleicht
ISBN: 978-3-941968-83-7
Content
1Introduction 13
2Literature overview 15
2.1Definitions 15
2.2Ethology 21
2.3Behavioural observations 22
2.4Normal behaviour of dairy calves 23
2.4.1Resting behaviour 25
2.4.1.1Lying down and standing up 25
2.4.1.2Resting, lying, sleeping 25
2.4.2Feeding behaviour 26
2.4.2.1Drinking behaviour 27
2.4.2.2Intake of pasture and textured supplementary feed 28
2.4.2.3Behaviour associated with water-intake 29
2.4.3Elimination and comfort behaviours 29
2.4.4Social behaviour 30
2.4.4.1Dam–calf relationships 30
2.4.4.2Social contacts of the calves 33
2.4.5Locomotion, exploratory, inquisitive and play behaviours 35
2.4.5.1Locomotion behaviours 35
2.4.5.2Exploratory, inquisitive and play behaviours 36
2.4.6Behaviour in the circadian rhythm 40
2.5Off-cow rearing of dairy calves 41
2.6Behaviour and health 44
2.6.1Behavioural enrichment 44
2.6.2Ethopathies and behavioural disorders in calves 46
2.6.3Behaviour of sick calves 47
2.7The genetics of behaviour 49
2.8Animal welfare, animal friendliness and well-being 51
2.9Animal welfare and ethics 58
3Materials and methods 60
3.1Study farm, housing and feeding of the calves 60
3.2Data acquisition and processing 62
3.3Statistical analysis and presentation of the results 65
4Presentation of the results 67
4.1Circadian rhythm 67
4.2Resting behaviour 69
4.2.1Resting behaviour to day 49 69
4.2.2Resting behaviour in the weaning phase 73
4.3Food and water-intake behaviour 75
4.3.1Food and water-intake behaviour up to day 49 75
4.3.1.1Drinking behaviour 75
4.3.1.2Behaviour associated with supplementary feed intake 78
4.3.1.3Behaviour associated with water-intake 82
4.3.1.4Visits to the dispensing station without CMR-feed-intake (blind visits)
83
4.3.2Food and water-intake behaviour during the weaning phase 86
4.3.2.1Drinking behaviour 86
4.3.2.2Behaviour associated with supplementary feed-intake 89
4.3.2.3Behaviour associated with water-intake 94
4.3.2.4Visits to the dispensing station without CMR-feed-intake (blind visits)
95
4.4Social behaviour 98
4.4.1Social behaviour up to day 49 98
4.4.2Social behaviour in the weaning phase 100
4.5Other activities and play behaviour 100
4.5.1Other activities 100
4.5.2Play behaviour 105
4.6Sucking activities without visiting the dispensing station 111
4.6.1Cross-sucking on another calf 111
4.6.2Sucking on dummy teats 114
5Discussion 119
5.1Methodology used to observe and evaluate the behaviour 119
5.2Behaviour of calves in off-cow intensive rearing 121
5.2.1Circadian rhythm 121
5.2.2Resting behaviour 124
5.2.3Food and water-intake behaviour 127
5.2.3.1Drinking behaviour 127
5.2.3.2Behaviour associated with supplementary feed-intake 130
5.2.3.3Behaviour associated with water-intake 132
5.2.3.4Visits to the dispensing station without CMR-feed-intake (blind visits)
133
5.2.4Social behaviour 135
5.2.5Other activities and play behaviour 137
5.2.5.1Locomotion and exploratory behaviour 138
5.2.5.2Play behaviour 139
5.2.6Sucking activities without visiting the dispensing station 145
5.2.6.1Cross-sucking on another calf 145
5.2.6.2Sucking on dummy teats 147
5.3Normal behaviour of calves in intensive off-cow rearing 149
5.4Ethogram for calves in off-cow rearing 153
6Summary 155
Zusammenfassung 158
Acknowledgements 161
References 162
Attachment of tables 193
Figures
Figure 1: Lying positions of calves, groups of starting positions with bent front
limbs: square position, hind limbs .............................................................. 26
Figure 2: Natural suction drinking of adult cattle ........................................... 29
Figure 3: Play behaviour of calves ................................................................ 39
Figure 4: Animal welfare in relation to environmental challenges as shown by
the outdated concept-based homeostasis and the new concept based on the
inverted U curve of (di)stress. .................................................................... 45
Figure 5: Three concepts for animal welfare ................................................. 56
Figure 6: Calf group pen with lying and running area, hay rack, dispensing
station, feeding trough, toy (brush, chairs and ball) ................................... 61
Figure 7: Schematic of the calf group pen with lying and running area, hay rack,
dispensing station, feeding trough, toys (brush, chains and ball) and dummy
teats as well as location and field of view of the cameras.......................... 61
Figures 8: (a) – (d): Drinking and feeding facilities in the calf pen: (a) CMR-feed
dispensing station, (b) automatic drinker, (c) trough and (d) rack .............. 62
Figures 9: (a) – (c): Toys: (a) ball, (b) chains, (c) brush ................................ 63
Figures 10: (a), (b): (a) ‘Milk Bar’ calf feeder, (b) dummy teats ..................... 63
Figure 11: Scheme of coded behavioural categories .................................... 65
Figure 12: Mean duration of activities and rest time during the day from the
week 3 to 7, n = 9 calves ........................................................................... 67
Figure 13: Mean number of CMR-feed-intakes per animal and day from 4:00 to
6:00 a.m. as well as from 6:00 to 8:00 a.m. by seasons from week 3 to 7, n
= 9 calves, significance: 4:00 to 6:00 a.m. vs. 6:00 to 8:00 a.m., p = 0.01 68
Figure 14: Mean duration of activities and rest time in minutes during weaning
from day 50 to 105, n = 5 calves ................................................................ 69
Figure 15: Mean duration of total rest time in hours per day from week 3 to 7,
n = 9 calves ............................................................................................... 70
Figure 16: Mean duration in minutes and number of rest sessions per day from
week 3 to 7, n = 9 calves ........................................................................... 71
Figure 17: Mean duration of rest sessions in minutes over the course of the day
from week 3 to 7, n = 9 calves ................................................................... 71
Figure 18: Percentage of activities during 5 minutes before and after rest from
week 3 to 7, n = 9 calves ........................................................................... 72
Figure 19: Mean duration of total rest time in hours per day during weaning
from day 50 to 105, n = 5 calves ................................................................ 73
Figure 20: Mean duration of rest sessions in minutes and number of rest
sessions per day during weaning from day 50 to 105, n = 5 calves ........... 74
Figure 21: Percentage of activities during 5 minutes before and after rest during
weaning from day 50 to 105, n = 5 calves ................................................. 75
Figure 22: Mean duration in minutes and mean number of CMR-feed-intakes
per animal and day from week 3 to 7, n = 9 calves .................................... 76
Figure 23: Percentage of CMR-feed-intakes over the course of the day from
week 3 to 7, n = 9 calves ........................................................................... 76
Figure 24: Percentage of activities during 5 minutes before and after CMR-
feed-intake from week 3 to 7, n = 9 calves ................................................ 78
Figures 25: (a), (b): Mean duration in minutes and number of feed-intakes per
animal and day (a) total solid feed (b) hay and dry TMR (total mixed ration)
from week 3 to 7, n = 9 calves ................................................................... 79
Figure 26: Percentage of solid-feed-intakes over the course of the day from
week 3 to 7, n = 9 calves ........................................................................... 80
Figure 27: Percentage of solid-feed-intakes per animal by duration of intake
from week 3 to 7, n = 9 calves ................................................................... 80
Figure 28: Percentage of activities during 5 minutes before and after solid-feed-
intake from week 3 to 7, n = 9 calves ......................................................... 81
Figure 29: Mean duration in minutes and number of water-intakes per animal
and day from week 3 to 7, n = 9 calves ..................................................... 82
Figure 30: Percentage of water-intakes during the day from week 3 to 7, n = 9
calves......................................................................................................... 83
Figure 31: Mean duration in minutes and number of blind visits per animal and
day from week 3 to 7, n = 9 calves ............................................................ 84
Figure 32: Percentage of blind visits by mean duration in minutes from week 3
to 7, n = 9 calves ........................................................................................ 84
Figure 33: Percentage of blind visits over the course of the day from week 3 to
7, n = 9 calves............................................................................................ 85
Figure 34: Percentage of activities during 5 minutes before and after blind visits
from week 3 to 7, n = 9 calves ................................................................... 86
Figure 35: Mean duration in minutes and number of CMR-feed-intakes per
animal and day during weaning from day 50 to 105, n = 5 calves ............. 87
Figure 36: Percentage of CMR-feed-intake over the course of the day during
weaning from day 50 to 105, n = 5 calves ................................................. 88
Figure 37: Percentage of activities during 5 minutes before and after CMR-
feed-intake during weaning from day 50 to 105, n = 5 calves .................... 89
Figure 38: Mean duration of total solid-feed-intake in hours per day and number
of solid-feed-intakes per animal and day during weaning from day 50 to 105,
n = 5 calves ............................................................................................... 90
Figure 39: Percentage of solid-feed-intake over the course of the day during
weaning from day 50 to 105, n = 5 calves ................................................. 90
Figure 40: Percentage of solid-feed-intake by duration of intake during weaning
from day 50 to 105, n = 5 calves ................................................................ 91
Figures 41 (a), (b): Mean and maximum duration of intake of (a) hay and (b)
TMR (total mixed ration) during weaning from day 50 to 105, n = 5 calves 93
Figure 42: Percentage of activities during 5 minutes before solid-feed-intake
during weaning from day 50 to 105, n = 5 calves ...................................... 93
Figure 43: Mean duration in minutes and number of water-intakes per animal
and day during weaning from day 50 to 105, n = 5 calves ......................... 94
Figure 44: Percentage of water-intake over the course of the day during
weaning from day 50 to 105, n = 5 calves ................................................. 95
Figure 45: Mean duration in minutes and number of blind visits per animal and
day during weaning from day 50 to 105, n = 5 calves ................................ 96
Figure 46: Percentage of blind visits by mean duration in minutes during
weaning from day 50 to 105, n = 5 calves ................................................. 97
Figure 47: Percentage of blind visits over the course of the day during weaning
from day 50 to 105, n = 5 calves ................................................................ 97
Figure 48: Percentage of activities during 5 minutes before and after blind visits
during weaning from day 50 to 105, n = 5 calves ...................................... 98
Figure 49: Mean duration in minutes and number of calf–calf contacts from
week 3 to 7, n = 9 calves ........................................................................... 99
Figure 50: Percentage of calf–calf contacts over the course of the day from
week 3 to 7, n = 9 calves ........................................................................... 99
Figure 51: Mean duration in minutes and number of calf–calf contacts per
animal and day during weaning from day 50 to 105, n = 5 calves ........... 100
Figure 52: Mean duration of active time and other activities (locomotion,
exploration, play) in hours per animal and day and other activities as a
percentage of active time from week 3 to 7, n = 9 calves ........................ 101
Figure 53: Percentage of other activities over the course of the day from week
3 to 7, n = 9 calves ................................................................................... 101
Figure 54: Mean duration of active time and locomotion behaviour per animal
and day from week 3 to 7, n = 9 calves ................................................... 102
Figure 55: Licking objects: mean duration in minutes per day, maximum
number per animal and day and mean duration in minutes from week 3 to 7,
n = 9 calves ............................................................................................. 103
Figure 56: Mean duration of active time and other activities (locomotion,
exploration, play) in hours per animal and day and other activities as a
percentage of active time during weaning from day 50 to 105, n = 5 calves
................................................................................................................. 104
Figure 57: Percentage of other activities over the course of the day during
weaning from day 50 to 105, n = 5 calves ............................................... 104
Figure 58: Mean total duration in minutes and number of games per animal and
day and mean duration of games per animal and day from week 3 to 7, n =
9 calves .................................................................................................... 105
Figure 59: Percentage of games over the course of the day from week 3 to 7,
n = 9 calves ............................................................................................. 106
Figures 60 (a), (b): Mean (a) duration and (b) number of games with ball, brush
and chains as well as headbuttings per animal and day from week 3 to 7, n
= 9 calves ................................................................................................ 107
Figure 61: Percentage of playtime with ball, brush or chains as well as
headbutting in the total playtime per animal and day from week 3 to 7, n = 9
calves....................................................................................................... 108
Figure 62: Mean total duration in minutes per animal and day, mean duration
in minutes and number of games per animal and day during weaning from
day 50 to 105, n = 5 calves ...................................................................... 109
Figure 63: Percentage games over the course of the day during weaning from
day 50 to 105, n = 5 calves ...................................................................... 109
Figure 64: Mean duration of games with ball, brush and chains as well as
headbutting per animal and day during weaning from day 50 to 105, n = 5
calves....................................................................................................... 110
Figure 65: Percentage of playtime with ball, brush or chains as well as
headbutting in the total playtime per day during weaning from day 50 to 105,
n = 5 calves ............................................................................................. 110
Figure 66: Mean duration in minutes, maximum number of sucking sessions
per day and number of days with sucking activities from week 3 to 7, n = 9
calves, 1 sucking calf ............................................................................... 112
Figure 67: Mean duration of a sucking session in minutes, maximum number
of sucking sessions per day and number of days with sucking activities during
weaning from day 50 to 105, n = 5 calves ............................................... 112
Figure 68: Percentage of activities during 1 minute before and after sucking
activities, from week 6 to 15, n = 5 calves ............................................... 113
Figure 69: Mean duration of sucking activities at the calf feeder, at the dummy
teats next to the dispensing station (du sta) and at the gate (du gate), week
4 to 14, n = 10 calves .............................................................................. 115
Figure 70: Percentage of activities in the 5 minutes before and after sucking
activities on a dummy teat, from week 4 to 15, n = 10 calves .................. 118
Figure 71: Headbutting calves .................................................................... 141
Figures 72: (a) – (c): Toys for calves (a) ball, (b) brush, (c) chains ............. 142
Figure 73: A calf suckles another calf on the pasture ................................. 147
Tables
Table 1: Ethogram of the natural behaviour of suckling calves in suckler cow
husbandry on pasture at 10 weeks, proportion of observed activities as a
percentage over 24 hours .......................................................................... 41
Table 2: Husbandry and feeding plan ........................................................... 60
Table 3: Period of feeding and quality of solid feed ....................................... 60
Table 4: Number of evaluated calves, days and hours per week .................. 64
Table 5: Behaviour categories and their coding ............................................ 64
Table 6: Statistical parameters of sucking on dummy teats, proportion of
sucking events at night and proportion of visits to the dispensing station with
and without CMR-feed-intake in all activities up to 5 minutes before sucking
events from week 6 to 15 ......................................................................... 116
Table 7: Number of sucking sessions and sucking activities on the dummy teats
by calves, n = 13 calves ........................................................................... 117
Table 8: Ethogram of feed-intake behaviour of male HF calves in group housing
with automatic feeders at the age of 8 weeks .......................................... 129
Table 9: Mean number and duration of play activities of calves within 18 hours
with a CMR-feed-allowance of 9 – 12 L milk per animal and day up to week
8 ............................................................................................................... 140
Table 10: Differences
in
the
behaviour
of
mother-bonded
and
off-cow
reared
calves....................................................................................................... 151
Abbreviations
a.m. ante meridiem
approx. approximately
bv visits without CMR-feed-intake, blind visits
c number of calves
CCC calf-calf-contacts
cf calf feeder
CF conventional feeding
cm centimeter
CMR calf milk replacer
CMR-feed calf milk replacer feed
CMRA calf milk replacer feed-allowance
d day
du gate suction dummies at the gate
du sta suction dummies next to the dispensing station
et. al et alii, and others
FM fresh matter
g gram
GPS Global Positioning System
h² heritability
IF intensified feeding
kg kilogram
l liter
m meter
m² square meter
max maximum
ME Metabolizable Energy
min minimum
MJ Megajoule
ml milliliters
n number
p probability of error
perc. percentage
p.m. post meridiem
p.
n. post natal
p.
p. post partum
QTL Quantity Trait Locus
R round
resp. respectively
s standard deviation
T dry matter
TMR total mixed ration
vs. versus
XP crude protein
Introduction 13
1 Introduction
Genetic selection has improved productive outputs from agricultural livestock
while simultaneously increasing the risk of behavioural changes and physiolog-
ical and immunological problems. If genetic selection leads to homeostatic im-
balance in farm animals, pathologies may develop that negatively impact ani-
mal welfare. (RAUW et al., 1998)
Livestock should be kept in a manner that is beneficial for them. Many across
society believe that modern livestock production does not entirely meet this
requirement. A 2015 report prepared by the scientific advisory committee for
agricultural policy in the German Federal Ministry of Food and Agriculture con-
cluded that the conditions under which animals are kept are not sustainable
across large areas of animal husbandry. (STEFANSKI, 2016)
A German survey by CHRISTOPH-SCHULZ (2019) revealed that consumers have
quite heterogeneous views on animal husbandry. Attitudes are not as uniformly
hostile as often presented in the media. In this study the group with favourable
attitudes towards current animal husbandry methods was in fact larger than
those with negative views. For beef and dairy cattle farming, of the 1419 people
surveyed 63.4% and 61.3% respectively saw a need for improvement, while for
poultry and pig farming it was over 70%. However, this survey also revealed
that consumers have a perception of agricultural animal husbandry that does
not necessarily align with reality. For example, images of cows behind feed rails
are familiar from the media and consumers therefore assume that all cattle are
permanently fixed in place. This study also indicated that cattle husbandry has
become more accepted in recent years because visible measures for animal-
friendly husbandry have been introduced.
It is nevertheless of interest to increase animal welfare, particularly in calf rear-
ing, given that the young animals represent the potential productive output of
future dairy cows. In the publications in the current series on ‘Rearing of female
calves and young cattle in agricultural enterprises’, Parts I and II, the require-
ments for husbandry and feeding of female offspring were presented. Based
on this, Part III of this series of publications investigates the behaviour of calves
intensively reared with a daily allowance of 12 L calf milk replacer feed up to
day 49. As explained in Part II of the above publication ‘Influence of milk re-
placer supply in calf rearing on health, performance and well-being’, increasing
the volume of milk replacer supplied to between 12 L and 14 L until day 49 is
Introduction 14
necessary and achievable in agricultural practice. Weaning is done gradually
so that the calves are completely weaned on day 106.
Part III of this series of publications presents an ethogram for breeding calves
throughout the entire off-cow group housing. This positively answers the ques-
tion of whether off-cow rearing of calves can be designed to improve their well-
being as much as possible. Such normal behaviour can and will be used as a
benchmark for animal-friendly housing and feeding of calves in off-cow rearing
and should be further adapted to meet the natural needs of the calves.
Literature overview 15
2 Literature overview
2.1 Definitions
A calf is the young of a cow aged up to six months (SAMBRAUS, 1997a).
Ethology, behavioural biology or comparative behavioural research is the sci-
ence of human and animal behaviour and its causes. The study of animal be-
haviour using biological methods is part of zoology. (SAMBRAUS ET AL., 1978;
IMMELMANN, 1982; JENSEN ET AL., 1986; WUKETITS, 1995; VON BORELL, 2009)
Applied ethology is an area of ethology that examines animal species that are
of ‘immediate’ interest for humans and therefore places greater emphasis than
basic ethology research on the possible opportunities for practical implementa-
tion and application. Based on the two types of animals of immediate interest
in this sense, zoo animals and domestic animals, applied ethology is divided
into captive animal biology and farm animal ethology. (KLINGHAMMER AND FOX,
1971; IMMELMANN, 1982; SAUERMOST AND FREUDIG, 1999; VON BORELL, 2009)
As a branch of agricultural sciences and veterinary medicine, applied farm
animal ethology defines the behavioural needs of livestock based on their
level of domestication and mode of use, which can be used to evaluate hus-
bandry systems in terms of animal welfare (VON BORELL, 2009).
Behaviour is defined as the integrated responses and actions of whole living
organisms (SCHEUNERT and TRAUTMANN, 1987). It includes externally noticeable
movements and postures, short-term and reversible colour and shape changes
in individual body parts, vocal expressions, and excretion of glandular products
and other products (SAMBRAUS et al., 1978; APFELBACH and DÖHL, 1980). Be-
haviour is an essential component of the animal’s mechanisms of adaptation
and an inseparable part of its physiological or morphological adaptations
(JENSEN et al., 1986). It is the result of continuous changes to musculature that
can be considered an uninterrupted sequence of physical postures and move-
ments and can be described by their environmental impacts or functions, allow-
ing a precise representation of an activity (HUNTINGFORD, 1984). Behaviour in-
cludes an animal’s movements, vocal expressions and physical postures as
well as any externally noticeable changes that are intended for reciprocal com-
munication and thus in turn trigger a behavioural response from the partner
(IMMELMANN, 1982). It is based on taking in, processing and expressing infor-
mation, has a high evolutionary significance and is encoded in the genetic ma-
terial of an animal (SCHEIBE and BILDT, 1987; VON BORELL, 2009). Behaviour is
expressed primarily as more or less pronounced motor activity comprising
Literature overview 16
changes in the position of individual body parts or the entire body; individual
behaviours can occur in different functional areas (SCHEUNERT and TRAUTMANN,
1987).
When behaviours are defined by certain rules or are directed at an appropriate
object, they form part of the normal behaviour of an animal (SAMBRAUS,
1997b). This is determined for individuals in a group of animals, each of a de-
fined genetic origin that can establish and maintain themselves, that is, they
can also avoid harm, without human help in an environment closely resembling
their natural one (ZEEB, 1989).
An ethogram is a comprehensive and differentiated catalogue of as many be-
haviours and behavioural patterns as possible that are exhibited by an animal
species and requires observation and recording of activities. An ethogram in-
cludes a detailed description of the behavioural characteristics of a specific
species and precise naming of individual behaviours in their spatial and tem-
poral sequence (SAMBRAUS and STEINEL, 1978; APFELBACH and DÖHL, 1980;
IMMELMANN, 1982; HÖRNING, 1993; EIBL-EIBESFELDT, 1999; FÖLSCH and KREMER,
2002; FRASER and BROOM, 2002). Behaviours of specific individuals are also
included in ethograms (SCHEUNERT and TRAUTMANN, 1987).
Functional areas describe the type of relationship between an organism and
its environment. First described by VON UEXKÜLL (1921) as a ‘self-contained
chain of effects in every animal action that links subject and object to each
other’. Subjects are the organisms, and their environment is the object that is
the source of stimuli (VON UEXKÜLL, 1921; GATTERMANN, 1993). The most im-
portant of these according to VON UEXKÜLL (1921) are prey, enemies and sexual
partners as well as the medium. The term ‘functional area’ today refers to the
action classes of behaviour, which differ in terms of their purpose, motivation
or reference object (GATTERMANN, 1993). In modern farm animal ethology, the
term ‘functional area’ is understood to be the sum of behaviours that can be
allocated to individual body functions (BOGNER and GRAUVOGL, 1984). FRASER
(1983) differentiates eight primary behavioural systems and many specific be-
haviours that are seen under natural free-ranging conditions. For cattle, there
are 44 specific behaviours described for the following functional areas: behav-
iours related to rest, social interactions, locomotion, food intake, elimination,
play and exploratory behaviour, and reproduction; deviations from these are
defined as behavioural disorders (WINCKLER, 2009).
Well-being is more than freedom from suffering, pain or harm; it is freedom
from hunger, thirst and malnutrition, access to comfort and protection, freedom
from diseases and injuries, the opportunity to express natural patterns of
Literature overview 17
behaviour, and freedom from stress and fear (TEUTSCH, 1987; FAWC, 1993).
Along with health, well-being includes an undisturbed, behaviourally appropri-
ate progression of the life processes appropriate for a species, freedom from
negative sensations and greater needs in an environment that does not exceed
the ability of the animal to adapt (SAMBRAUS and STEINEL, 1978; STUBER, 1980;
VAN PUTTEN, 1982; GRAUVOGL, 1995, 1997; SAMBRAUS, 1997b). Germany’s An-
imal Welfare Act considers well-being to be the undisturbed, species-appropri-
ate and behaviourally appropriate sequence of life processes. A more favoura-
ble environment leads to higher well-being (JENSEN and SANDØE, 1997). PUPPE
et al. (2012) define well-being as ‘the state of physical and mental health that
results – against the backdrop of individual, also cognitive requirements and
abilities – from the process of ethological and physiological adaptation resulting
from coping with challenges present in the environment and the subjective ex-
periences and emotional evaluations made throughout’.
Any phenomenon that can be perceived through the senses and described can
be used as a characteristic (TSCHANZ, 1985).
Stimuli are measurable physical, chemical, optical, acoustic, electrical or mag-
netic states or changes in state in the environment or in the organism that lead
to animals maintaining or changing behaviours and which can trigger reactions
(APFELBACH and DÖHL, 1980; GATTERMANN, 1993).
SCHEUNERT and TRAUTMANN (1987) view motivation (also a readiness to act or
drive) as resulting from an inner state that triggers a behaviour without its key
stimuli having to be present. Movement sequences without an identifiable ex-
ternal stimulus, known as vacuum activities, are triggered if drive or motiva-
tion are blocked or if the threshold is lowered (VON BORELL, 2009).
The term environment expresses the totality of the sensory capacity of an an-
imal, whereby each species has its own characteristic environment (FRASER
and BROOM, 2002). The entire surroundings or external world of an animal is
circumscribed and not differentiated into factors that are significant or mean-
ingless for the animal; in other words, the environment is the totality of those
factors in the surroundings or surrounding conditions that in fact exert some
effect on the organism or which the organism affects (IMMELMANN, 1982;
SCHEIBE and BILDT, 1987).
Intensive animal husbandry is characterised by rapid animal turnover, high
animal stock density, a high degree of mechanisation, low workforce require-
ments and efficient conversion of feed into marketable products (PRESTON,
1961, cited in HARRISON (1968)). For BOGNER and GRAUVOGL (1984), this pro-
cess is characterised by concentrating animal stocks in a relatively small area
Literature overview 18
and year-round housing systems. The United Nations Food and Agriculture Or-
ganization defines intensive livestock production as systems in which less than
10% of the dry matter fed to animals is farm produced and in which the stocking
rate exceeds ten livestock units per hectare of agricultural land (SERÉ and
STEINFELD, 1996). In the European Union, intensive livestock production sys-
tems are defined as animal husbandry methods in which animals are kept in
such a number in such a small area, under such conditions or at such a pro-
duction level that their health and welfare depend on frequent controls by hu-
mans (92/583/EEC, Article 1(1) clause 3).
Extensive animal husbandry is defined by BOGNER and GRAUVOGL (1984) and
BAUMGARTNER (1997) as the opposite to intensive livestock production and also
requires more intensive monitoring by farm workers. For ERNST (1996) an ani-
mal husbandry method is extensive if at least one production factor (work, soil,
capital) is used extensively, that is, only minimally. The aim of extensive live-
stock farming is to cultivate usually unprofitable agricultural areas in a manner
that is still economically viable; in densely populated arable areas, preservation
of cultivated landscapes is also included in the interests of conservation
(ACHILLES and MARTEN, 1997).
Imprinting is an obligatory learning process in early development in which the
newborn becomes familiar with features of its environment (LORENZ, 1978;
SCHEUNERT and TRAUTMANN, 1987). For KOLB (1987) imprinting refers to the
strong bond between a young animal and its mother. With imprinting, specifi-
cally social imprinting, the animal becomes attached to the species it first has
contact with (SAMBRAUS et al., 1978).
Emotions refer to affective states in humans and mammals controlled by the
brain using physiological and/or neurochemical processes and associated with
a subjective perception, the feeling. Environmental stimuli are interpreted
based on these emotional states. Examples of feelings include pain, malaise,
tiredness, hunger, thirst, thermal discomfort, fear, grief, frustration, boredom,
loneliness, general suffering, enjoyment, jealousy, anger, sexual pleasure.
(BROOM, 1998; KÄSTNER, 2020)
For MASON (1984) temperament combines the basic attitude of an individual
towards environmental change and its challenges. This definition is later limited
to behavioural reactions of the animal in dealings with humans and can be in-
cluded as a measure of human–animal interactions (BURROW, 1997; GAULY et
al., 2001; SCHUTZ and PAJOR, 2001). Differences in temperament are seen be-
tween breeds (GONYOU, 2019) and between individuals in a breed (LE NEINDRE
et al., 1995).
Literature overview 19
Pain is a personal experience that is influenced to varying degrees by biologi-
cal, psychological and social factors (MERSKEY and BOGDUK, 2012). It varies in
intensity, duration and quality and is described as stabbing, burning, tearing,
dull (LOEFFLER, 1990; SAMBRAUS, 1997b). It cannot be perceived as a subjective
sensation in animals but leads to aggressive reactions or flight behaviour
(BERNATZKY, 1997). The inability to communicate does not negate the possibil-
ity of animals experiencing pain (MERSKEY and BOGDUK, 2012; PRUNIER et al.,
2012). In animals the degree of pain is determined using nonverbal methods,
such as motor (e.g. flight) or autonomic reflexes (e.g. heart and respiratory
rates), and endocrine reactions (e.g. cortisol level) (OTTO, 1997). Acute pain
and suffering can lead to behavioural changes that can be detected by direct
observation (GÄRTNER and MILITZER, 1993).
Suffering in an animal is a sustained, subjective perception of impaired physi-
cal functions and behaviour (SCHEIBE, 1997). Avoidable events are described
as suffering, meaning that natural feelings such as the fear of a prey animal
when detected by predators are not included (FRASER and BROOM, 2002). One
criterion to estimate suffering is the extent to which it is no longer possible for
an animal to behave in a species-specific way (DAWKINS, 1982). Animals living
in captivity suffer in situations in which they are prevented from doing some-
thing that they are highly motivated to do (DAWKINS, 1990). Feelings, which also
include suffering, can only be comprehended by drawing analogies (SAMBRAUS,
1982). The most important instrument to evaluate pain and suffering, particu-
larly under field conditions, is observing the animal’s behaviour, with additional
information provided by physiological measurements such as heart rate, res-
piratory rate and body temperature (BARNETT, 1997). DAWKINS (2008) defines
suffering as a series of negative emotions such as fear, pain and boredom that
can be recognised operationally as a state caused by negative reinforcers.
Fear is a particular type of suffering, an unpleasant emotional state with the
expectation of a highly negative event (SAMBRAUS et al., 1978; SAMBRAUS,
1997b). Often no external or internal injuries are apparent, but fear neverthe-
less causes sweating, accelerated heart function or huddling together in a
group; fear-induced urination, pale skin, shaking, raised hackles and widely
open eyes or mouth have also been observed as fear reactions as is ‘panic-
stricken falling down’ (GRAUVOGL, 1983). The intensity of fear reactions is influ-
enced by genetics, meaning that domesticated animals show a weakened flight
response as a result of selective breeding (GRANDIN and DEESING, 1998). Flight
or fight responses are controlled by the amygdala, a paired almond-shaped
Literature overview 20
region of the brain that is part of the limbic system (SETEKLEIV et al., 1961;
MATHESON et al., 1971; ALM et al., 2005).
Ethopathies or behavioural disorders are behaviours that deviate from a
species-specific locomotor coordination in terms of the frequency of occurrence
or their sequence and that are induced by unfavourable husbandry conditions
(APFELBACH and DÖHL, 1980). Behavioural disorders caused by organic pathol-
ogy have developed during domestication and may be caused by genetics or
exogenous factors (IMMELMANN, 1982; BUCHHOLTZ, 1993). They include any be-
haviour that significantly and persistently deviates from the species-specific
norm for an individual animal or a group in terms of its modality, intensity or
frequency and that occurs repeatedly or persists, can be inherited or acquired,
and is likely to harm the organism or another member of the same species
(MEYER, 1984; SAMBRAUS, 1997b). Behaviour that deviates from the species-
specific norm exceeds the ability of an individual to adapt to certain environ-
mental conditions and generally results in harm to the organism, meaning that
outputs cannot be achieved that are necessary for developing and maintaining
the physical and behavioural characteristics that correspond to the norm, either
for the individual itself or other members of the same species, or for preserving
the reproductive community (ZEEB, 1989; TSCHANZ, 1995). The occurrence of
abnormal behaviour indicates that the farm animals are not coping with the en-
vironment created for them by humans, their fundamental environmental re-
quirements are not adequately considered and their ability to adapt is exceeded
(SCHEIBE, 1987; HÖRNING, 1993). SAMBRAUS (1997b) assigns ethopathies to the
following categories:
1) actions directed towards an inadequate object, e.g., without an ob-
ject or directed towards inanimate and animate objects
2) altered sequences of behaviour
3) behaviour with a frequency deviating significantly from the norm
4) stereotypies
5) apathy
Stereotypies are invariant, repeated, relatively unchanged sequences of
movements, almost identical, repetitive behavioural patterns or vocal expres-
sions that do not have an obvious purpose and are performed with no detecta-
ble function (IMMELMANN, 1982; MASON, 1991; LEBELT, 1998; FRASER and
BROOM, 2002; VON BORELL, 2009). They persist even when the causative fac-
tors are not present (SAMBRAUS et al., 1978). They are not necessarily indica-
tions of reduced welfare (BRUMMER, 1977). WECHSLER (1995) suspects that
Literature overview 21
stereotypies become established to reduce stress if it is no longer possible to
adapt to the environment.
SELYE (1973) defines ‘stress’ as a non-specific reaction by the body to any
demand placed on it to adapt, regardless of whether this demand generates
pain or pleasure. The absence of chronic stress is a prerequisite for the welfare
of animals, and environmental stimuli that lead to imbalance in the homeostasis
(according to CANNON (1932): ‘balance of the physiological mechanisms’) are
defined as ‘stressors’ and the corresponding defensive response of an animal
is described as a ‘stress response’ (MÖSTL and PALME, 2002).
Technopathies are injuries to individual animals’ bodies that occur because of
housing environments that are not behaviourally appropriate (ZEEB, 1989). This
includes injuries to the extremities (claw injuries, claw/joint inflammation, lame-
ness), infectious diseases (e.g. pneumonia, endometritis), injury caused by
other members of the species (e.g. biting of ears or tails) and behavioural dis-
orders (e.g. stereotypies) that develop as a result of technical facilities
(TROEGER, 1996).
2.2 Ethology
Ethology, the biology of behaviour, has gradually gained a foothold in the nat-
ural sciences and become firmly established across several disciplines. Initially,
the behaviour of animals was investigated more as a model for the behaviour
of humans, however. (KLINGHAMMER and FOX, 1971)
Konrad Lorenz founded the science of ethology in the German-speaking coun-
tries in the 1930s and was awarded the Nobel Prize for his research. Since its
inception, ethologists emphasised the importance of the applied aspects of this
scientific discipline. In the late 1950s the first publications in applied ethology
appeared, with behavioural observations of domesticated animals available
only from this time on. (SAMBRAUS, 1999)
Humans have only a limited ability to see a situation rationally and emotionally
from an animal’s perspective (DAWKINS, 1982; SCHEIBE, 1997).
Questions related to animal behaviour can only be answered by ethological
considerations. The most important principle of ethology, the scientific, objec-
tive study of animal behaviour, is a recognition that the fundamental behaviours
of animals are innate and specific for a species. Understanding the require-
ments domestic animals have of their housing environment needs awareness
of the behaviour of the original or wild forms. Basic biological functions form
mechanisms that have developed in the species and persist to the present in
Literature overview 22
domesticated animals. Domestication started 2,500 years before recorded his-
tory, meaning that naturally the behaviour of the original wild forms of today’s
domesticated animals can no longer be observed. For this reason, we look at
primitive breeds or wild species with life rhythms that are not determined by
feeding or milking times such as, for cattle, the semi-wild Camargue cattle that
are tended by herders, Chillingham wild cattle in Northumberland or bison.
Suckler cow husbandry on pasture also provides an opportunity to observe the
natural behaviour of domesticated cattle for which the relationship to humans
and changes in behaviour due to domestication and breeding in particular play
a role. (SCHLOETH, 1961; THORPE, 1965; KOCH, 1968; ZEEB and MACK, 1970;
SCHLICHTING and SMIDT, 1986; SCHEUNERT and TRAUTMANN, 1987; BROOM,
1991; TSCHANZ, 1995)
An essential prerequisite for better understanding how animals perceive and
understand their environment and anticipate future events is an awareness of
the physico-cognitive capacities of farm animals. This knowledge can help to
reduce stress in housing and managing the animals, to design novel types of
enrichment and thus to improve animal welfare. (NAWROTH et al., 2017)
2.3 Behavioural observations
Behaviour can be perceived and interpreted, and its causes and consequences
can be identified. Humans have long been familiar with the behaviour of do-
mesticated and wild animals. The prehistoric hunter needed such knowledge
to capture prey and avoid dangerous animals. The initial step for animal keep-
ers in the domestication process is understanding of the needs of their animals
to avoid any dangers and prevent animals escaping. (SAMBRAUS, 2002)
Behaviour enables animals to interact with their environment and adapt to
changing environmental conditions. It develops over the evolutionary history of
a species and satisfies specific biological functions that enable growth, self-
maintenance and reproduction. (SCHRADER, 2007)
The classification of the behaviour of farm animals is based on oral or written
descriptions of an activity in which the animal is involved. Quantifying the be-
haviour of animals for research requires that a person identifies and encodes
the behaviour of the animal being studied. The classification of behaviours may
be subjective and differs between observers. (JIANG AND DAILEY, 1996;
MORROW-TESCH ET AL., 1998)
Ethologists concentrate as a rule on describing the form of behavioural patterns
(HUNTINGFORD, 1984).
Literature overview 23
WEBSTER and SAVILLE (1982) noted, however, that slight differences between
observers did not distort comparisons of calf behaviour between systems. Be-
havioural observations by trained persons are appropriate for evaluating animal
husbandry systems.
In the early days of ethology, the behaviour of animals was directly observed
on site. In studies by WALKER (1962) two observers alternated every two hours
(h) over the day. At night behaviour was observed every four hours under illu-
mination with torches and noted. GRAF et al. (1976) observed the lying behav-
iour of fattening calves over 24 h, with four observers alternating in a 6-hourly
rhythm.
Since camera technology has become available, the behaviour of animals has
been recorded and subsequently assessed. This method has the advantage of
enabling images to be viewed again at any time and minimising the personnel
required. The animals are also not disturbed by the presence of the observers
and display their natural behaviour. (WEBSTER AND SAVILLE, 1982; BRUMMER,
2004)
Behaviour is also directly observed on site using camera recordings so that the
results can be applied to the real world of agriculture or the wild habitat of an
animal (DAWKINS, 2008).
Most studies use standard digital colour video cameras for capturing data.
Large scale phenotyping of animal behaviour characteristics is time-consum-
ing. Small sample sizes are usually selected for training and validating the ob-
servations. (WURTZ et al., 2019)
It is helpful to record only a few characteristics so as to not unnecessarily in-
crease costs (SCHMIDT, 1998). Only individual focus animals should be contin-
uously or periodically observed and assessed over a specific representative
period in terms of previously defined behavioural parameters (HOY, 2009a).
2.4 Normal behaviour of dairy calves
TSCHANZ (1985) assumes that all manifestations of a characteristic that lie
within the 95% limit of a normal distribution can be described as ‘normal’. De-
viations from this indicate that the environment does not meet the requirements
of the animals or legislators and should be examined more closely. The type of
individual that has arisen in domesticated animals as a result of domestication
and breeding and that fundamentally differs from the wild form must be consid-
ered, such as the characteristics of a breed. Modifications in behaviour have
thus developed as adaptations that enable a domesticated animal to behave
Literature overview 24
according to the norm. If this is not possible, behavioural changes appear that
must be defined as harmful and their correlations can be used as indicators.
Domestication involves an evolutionary process that is based on switching as
much as possible from natural selection to artificial selection by humans. Wild
and domesticated animals therefore require different conditions and resources
to achieve a high degree of welfare, even though biologically they still belong
to the same species. High welfare is thus not achieved simply by providing the
animals with the requirements for which their wild predecessors were selected.
Many expectations for domesticated animals (e.g. constant availability of food,
controlled environmental conditions) refer to the artificial environment created
for them by humans. Furthermore, their physiological and behavioural systems
are designed so that they are adapted to the specific conditions of the human
environment. At the same time, the ability to adapt may be overwhelmed if the
animals are challenged by ‘natural’ stressors. However, domesticated animals
require some essential factors for which their wild predecessors were selected.
The reference system to identify what ensures a high degree of welfare for do-
mesticated animals is therefore a generous, animal-friendly husbandry system
(e.g. suckler cow husbandry on pasture) rather than the ecological niche of the
wild predecessors. (BROOM and JOHNSON, 1993; SACHSER, 2001)
Changes to the frequencies of behaviours may indicate that an animal is suf-
fering. Changes in temperament, comfort behaviours, play or the circadian
rhythm as well as stereotypies and changes to food intake behaviours also in-
dicate negative impacts on animals’ welfare. (FRASER, 1984)
Coping behaviour in animals is a reaction to adverse situations in an unnatural
environment. If this behaviour is unsuccessful, the animals respond with abnor-
mal behaviour. (WECHSLER, 1995)
Intensive breeding has contributed to changes in behaviours. The separation
of a dairy cow from her calf and its rearing has led to a reduced influence on
temperament and adaptability. (LE NEINDRE, 1989)
Husbandry-related restrictions of certain aspects of normal behaviour do not
always mean that the husbandry method is not animal friendly. Welfare is not
impaired in the absence, for example, of stimuli that trigger flight or defensive
behaviour, even though this behaviour is part of the natural spectrum. An ani-
mal cannot avoid other behaviours such as sucking. Their absence triggers
frustration or even behavioural disorders. An understanding and consideration
of the resting and active behaviour of farm animals is thus an essential prereq-
uisite for animal-friendly husbandry. (SCHRADER, 2007)
Literature overview 25
‘Reactive disorders’ are the result of unsuitable husbandry conditions. Most be-
long to the functional areas of feeding behaviour and locomotion because these
are the two aspects that are most constrained in agricultural animal husbandry.
Adaptation of natural behaviours can be observed, such as cattle standing like
horses when there is insufficient space available. (SAMBRAUS, 2002)
2.4.1 Resting behaviour
2.4.1.1 Lying down and standing up
Calves lie down and stand up very soon after birth in a way that is also observed
in adult cattle (SAMBRAUS, 1985).
Calves normally start the lying down process by first bending the forelegs and
lowering down onto the carpal joints. Then they allow the bent hind legs to drop
down onto the upper legs and then the lying position is taken with the forelegs
folded backwards. Young cattle often lie with equal frequency on the right or
left side. The normal procedure for standing up begins with the front legs, with
the animals pushing themselves forwards with the hindquarters, kneeling on
the carpal joints and then raising the hindquarters with a head swing, which is
why they need plenty of space around their heads. They then extend their fore-
legs. In calves, this often happens very quickly and they almost appear to jump
upright. (SAMBRAUS, 1971; KILEY-WORTHINGTON and PLAIN, 1983; SAMBRAUS,
1985)
2.4.1.2 Resting, lying, sleeping
Calves need plenty of rest and seek out protected spaces to rest. Before lying
down, they intensively sniff the ground and often lie down where other calves
or they themselves have previously rested. There have been 40 different lying
positions described with Figure 1 showing a selection of the most common.
With the normal posture, the calf lies on its breastbone, the body is rotated on
itself, and the rear of the body lies on one leg with the hind legs extended away
from the body. Calves also often sleep lying flat on their sides, presumably be-
cause of accumulated gases in the as-yet underdeveloped rumen. The resting
and sleeping duration change only slightly in the first six months of life.
(SCHEURMANN, 1971; SÜSS and ANDREAE, 1974; KILEY-WORTHINGTON and PLAIN,
1983; SAMBRAUS, 1985, 1997a)
Literature overview 26
Figure 1: Lying positions of calves,
groups of starting positions with bent
front limbs: square position, hind limbs
Source: SCHEURMANN, 1971
WEBSTER and SAVILLE (1982) identified
only minimal differences between vari-
ous husbandry systems regarding the
time the calves spend lying down. Re-
gardless of the housing method, calves
aged 14 weeks spent more time lying
down than all older animals.
FERRANTE et al. (1998) identified differ-
ences in the resting behaviour depend-
ing on the husbandry system: Calves
that were housed in larger calf pens
rested during the day for significantly longer than calves of the same age in
small pens. Resting disorders between the calves increase with the size of the
group, which is why a group should not contain more than 12 to 15 animals
(LIEBENBERG, 1965).
BRADE (2001, 2002) provides guidelines for the resting behaviourof calves in
the first five weeks, suggesting 20–22 hours rest per day.
For young calves, standing up is strenuous, which is why they need a thick dry
mat of straw so that they do not slip while standing up. Slippery floor materials
lead to animals preferring not to lie down and stand up. The consequence of
this would be lying down for too long on a hard surface, leading to stiffening of
the limbs. (SAMBRAUS, 1985)
This is also shown in a comparison of postures of fattening calves. On deep
litter, calves rest in total for considerably longer than on fully slatted floors,
stand up more often and also rest briefly during the day. The litter is also used
by the animals as something to occupy themselves with. (GRAF et al., 1976)
2.4.2 Feeding behaviour
Feeding behaviour is very complex. Animals are highly curious, which enables
them to learn what it is in their surroundings that they must be aware of.
(THORPE, 1965)
Literature overview 27
An understanding of species-specific feeding behaviour and animals’ nutritional
needs is important for evaluating their health, protection and welfare (NIELSEN
et al., 2016).
2.4.2.1 Drinking behaviour
The sucking reflex is one of the innate motor responses of the newborn mam-
mal. This complex, dominant, unconditional reflex has clear arousal and inhibi-
tion phases (KRUM and CUSKOW, 1958). It is strongest and most persistent from
day 15 to 60. A calf seeks out the udder with vertical bumping movements about
40 to 100 cm above the ground and finds a small, compact udder more quickly
than a large pendulous one, with a preference for the underside of the cow’s
abdomen and particularly the area around the axils and knee folds. If the calf
bumps the back of its nose against the cow’s body, it starts making licking
movements until it finds a teat and only then does it begin sucking. The calf
sucks on each teat for five to 30 seconds and then it switches to one of the
other teats. The calf encloses the teat with its tongue, creating an air-tight seal,
and generates a vacuum so that the milk can flow. Towards the end of the
suckling period, the calf switches teats more frequently and bumps with its
head. Newborns generally do not suckle more often than six times a day while
at age three months they suckle three to at most five times a day for 5 to 10
minutes (WALKER, D.E., 1962; PORZIG, 1964; SCHEURMANN, 1974a; HAFEZ and
BOUISSOU, 1975; RIESE et al., 1977; SAMBRAUS et al., 1978; SOMERVILLE and
LOWMAN, 1979;KILEY-WORTHINGTON and PLAIN, 1983; SAMBRAUS, 1985; PORZIG,
1987).
From these observations, it is apparent that calves seek out a ‘feature’ that is
warm, with hairs, located about 40 to 100 cm above the ground and with an
upper margin. The odour of the milk also attracts the calves and helps in their
search for the source (KILEY-WORTHINGTON and PLAIN, 1983).
When suckling, calves usually stand with their limbs extended and the front
limbs spread apart, sometimes bent, with a lowered back and the head
stretched forward, in a reverse parallel position to the mother, raising their hind-
quarters and wagging their tails hard. The waggling movements of the tail indi-
cate peaceful and calm sucking. During the suckling process the calf often
bunts the cow’s udder with its head to stimulate the flow of milk. The cow almost
always remains inactive when suckling or sometimes ruminates but rarely eats.
(BROWNLEE, 1950; RITTER and WALSER, 1965; KOCH, 1968; HÜNERMUND, 1969;
SCHEURMANN, 1974a; SAMBRAUS et al., 1978; DERENBACH, 1981; BOGNER and
GRAUVOGL, 1984; SCHÄFFER et al., 1999; DE PASSILLÉ, 2001)
Literature overview 28
Most dams remain standing when suckling, while some check the calf by sniff-
ing (SCHÄFFER et al., 1999). More than 20% of suckling events are performed
through the back legs with the calf standing behind the cow (SCHÄFFER et al.,
1999), often when calves rob milk from other cows (DERENBACH, 1981). This
suckling position helps when robbing milk because the cows are unable to iden-
tify the calves by their smell and fend them off (SAMBRAUS et al., 1978; KILEY-
WORTHINGTON and PLAIN, 1983).
DE PAULA VIEIRA et al. (2008) did not note any change in suckling behaviour
between two and nine weeks for calves that could suck from their own mothers.
The ability to suck was not affected either by milk availability or feelings of hun-
ger.
In contrast, SCHEURMANN (1974b) observed that milkability and milk output of
the dams, but also the frequency with which a calf was allowed to go to its
mother, affected the duration of the suckling periods.
2.4.2.2 Intake of pasture and textured supplementary feed
The prerequisite for timely intake of pasture and textured feed is the ability to
ruminate and digest. Ruminating starts from day 7, at the latest from week 3.
From week 8, it increases rapidly and then only rises moderately with individual
differences developing. (PORZIG, 1987)
Calves start to eat supplementary feed from day five to eight. In the first weeks
of life, the calves already start to nibble on stalks of grass or hay. At three
months they have mastered the movement pattern of eating and adapt approx-
imately to the grazing cycle of the herd. Suckling calves graze most intensively
in the mornings and evenings and after suckling over the day. With species-
appropriate feeding, the calves start to ruminate at the latest from week 3.
(SAMBRAUS et al., 1978; BOGNER and GRAUVOGL, 1984; SAMBRAUS, 1985,
1997a)
Cattle have a highly mobile tongue that extends out of the side of the mouth
when eating to wrap around the feed and pull into the mouth. This movement
pattern is innate but is learned more quickly in the presence of older, experi-
enced animals. (SAMBRAUS et al., 1978; BOGNER and GRAUVOGL, 1984)
According to SAMBRAUS et al. (1978), there is a positive correlation between
eating duration and the quantity of feed ingested. The duration of the feed in-
take depends on the quality of the feed.
Literature overview 29
Calves prefer textured supplementary feed, that is, pellets instead of floury con-
centrate or muesli and coarse hay instead of finely ground hay (MONTORO et
al., 2013; FREITAG et al., 2018; PAZOKI et al., 2019).
2.4.2.3 Behaviour associated with water-intake
Cattle are suction drinkers (Figure 2), that is, they suck up the water. To drink,
the upper and lower jaws are separated and the tongue is pulled back so that
it acts like the piston of a mechanical suction pump. The muzzle is immersed
2–5 cm into the water source. Cattle tend to seek out water after eating.
(SAMBRAUS et al., 1978; BOGNER and GRAUVOGL, 1984; BARTUSSEK, 2008)
Source: BARTUSSEK (2008)
Figure 2: Natural suction drinking of adult cattle
The water should be drunk in sips into the rumen, which is why it must be of-
fered to calves from a bucket, drinking trough or an automatic drinker.
(RADEMACHER and LORCH, 2003; JILG and BRÄNDLE, 2007; SPRENG, 2011)
2.4.3 Elimination and comfort behaviours
Excretion (defecation and urination) in cattle is barely expressed in space and
time and is not accompanied by any specific behaviour. Faeces and urine are
often eliminated after lying down, while night sleep is also interrupted for elimi-
nation. Excrement is typically eliminated when standing, and cattle also defe-
cate while moving and rarely when lying down. When defecating, the hind legs
are slightly straddled and somewhat forward, the back is slightly bent and the
tail is slowly raised. When urinating cattle raise their tail higher than when def-
ecating and the back is curved more. The consistency and frequency of faeces
are closely related to the quantity of liquid and supplementary feed ingested.
(SAMBRAUS, 1971; SAMBRAUS et al., 1978; BOGNER and GRAUVOGL, 1984)
Calves that are isolated from their mothers eliminate their first faeces and urine
on the first day later than calves that have been intensively licked by the moth-
ers. Subsequently, the isolated calves lick themselves more, which METZ and
Literature overview 30
METZ (1985) consider to be an expression of a behavioural need, in this case,
the licking by the mother.
Comfort behaviour is expressed in calves primarily by licking body parts and
scratching and rubbing against objects or with their hind legs. Calves can al-
ready lick themselves on the shoulders and feet in the first hours of life, with
this initially done while lying down, and older calves groom themselves for
about 1.3 minutes every hour. (KILEY-WORTHINGTON and PLAIN, 1983)
Calves scratch themselves 28 times a day, in total for almost an hour. Mutual
licking or scratching is performed both for grooming and as part of social con-
tacts. (FRASER and BROOM, 2002)
Cattle preferably scratch those parts of their bodies where their tongues and
hooves cannot reach, particularly the head and neck region (SÜSS and
ANDREAE, 1974). Camargue calves on pasture always seek out particular spots
with bushes and trees (SCHLOETH, 1961).
2.4.4 Social behaviour
2.4.4.1 Dam–calf relationships
The social contacts between the calf and its mother and other calves are im-
portant for the development of social behaviour, which is essential for living in
the herd. These contacts are also important for the development of organs and
essential behaviours. (ZEEB and MACK, 1970)
Mother–child contacts in cattle are auditory (calls), tactile (sucking, rubbing and
licking), olfactory (smelling) and gustatory (tasting and licking) (KILEY-
WORTHINGTON and PLAIN, 1983). They enable the dam to bond with her calf, to
protect and feed it and ultimately to end this bond when weaning (KEYSERLINGK
and WEARY, 2007).
Intensively bred breeds such as Friesian have a genetically-induced weaker
mother–child bond than more extensive breeds such as Salers (LE NEINDRE,
1989).
Birth of the calf
A few hours before the birth, cows become restless and seek out a quiet, dry
place away from the herd. Shortly before and during calving, the cow vocalises
with a deep, rumbling moo that is later used to call the calf. In the first hour after
the birth there is intensive mother–child contact such as sniffing and licking of
the newborn calf by the dam and the rumbling vocalisations. After just five or
ten minutes, the calf will stand up, 20 to 60 minutes postnatal (p.n.) it can stand
Literature overview 31
up and seek out the udder, another five minutes to an hour later it can suck and
drink the colostrum. Individual needs then become more important. For the calf,
locomotion activities and seeking out the udder decrease, and it rests and
sleeps deeply. In the first three hours postpartum (p.p.), the imprinting of the
dam on her calf creates a stable bond and the mother very quickly recognises
her calf visually and olfactorily, licks it intensively and encourages it to suck if it
does not do so by itself. This licking dry of the newborn by the mother is a
reliable indicator of the start of a stable mother–child relationship. The cow
starts to eat close to the calf, within hearing range, two hours p.p. and rests
extensively in the first 72 hours. The calves rest for about 80% of daylight hours
in the first five days. The imprinting of the calf on the dam takes place in the
first four days of life. Initially, the calf recognises its mother only by her voice
and follows every large moving object unless the mother intervenes and drives
off other animals, placing herself between her calf and other animals. From the
fourth day, the calf can recognise its mother by her voice. The cow will rigor-
ously fend off any foreign calves until mutual imprinting of calf and dam is com-
plete. The calf approaches a cow with an extended head and only when it is
recognised and tolerated will it seek out the udder. (HAFEZ and BOUISSOU, 1975;
SAMBRAUS and BRUMMER, 1978; REINHARDT, 1980; DERENBACH, 1981; KILEY-
WORTHINGTON and PLAIN, 1983; KOLB, 1987; SAMBRAUS, 1992; BAUMGARTNER,
1997; MÖRCHEN, 1997; LANGBEIN et al., 1998; TOST and HÖRNING, 2001; UNGAR,
2006; CHENOWETH et al., 2014; SPENGLER NEFF et al., 2017)
Among Heck cattle kept in semi-wild conditions, PERREY et al. (1996) observed
two variations in the behaviour shortly before and after the birth. Some cows
separated themselves by more than 30 m from the herd and left their calves
lying secluded from the herd for up to four days p.p. when they went to eat. The
isolation from the herd over three days correlates with the need to imprint the
calf on its mother. Other cows calved in the middle of the herd.
HUDSON and MULLORD (1977) observed that in multiparous Jersey or Jersey–
Friesian cross cows, 5 minutes’ contact with a calf immediately after the birth
was sufficient to create a strong, specific maternal bond to her calf. This bond
also persisted when the calf was removed for 12 hours from the dam and then
was returned to her.
Calves are ‘hiders’, that is, they lie down in a protected position away from the
pasture where high vertical structures (tall grass, trees and bushes or walls and
fences) trigger the hiding behaviour. In many cases, the young animal actively
removes itself from the mother so that she does not know the precise hiding
location of the calf. This behaviour appears to have evolved so that the mother
Literature overview 32
does not reveal the location of the young animal to predators by her behaviour.
(LANGBEIN et al., 1998)
Similar observations are also described by VITALE et al. (1986) in semi-wild,
free-ranging Maremmana cattle in Tuscany. In the first two to three days of life,
the calves hid themselves in dense undergrowth while the cows grazed nearby.
As the calves got older, they spent more time at a greater distance from their
mothers.
Immediately after the birth, the dams spend a great deal of time caring for the
calf. As METZ and METZ (1985) observed, this time rapidly decreased over the
course of the first day and was only briefly seen on the third day p.p.
A bond between cow and calf also persists if the calves are only suckled and
no further contacts are possible (JOHNSEN et al., 2015).
Because the imprinting of the calf on its mother only takes place during the first
two days p.n., it possibly does not mean any stress for the newborn to be re-
moved from the mother (KILEY-WORTHINGTON and PLAIN, 1983).
Suckling period
LANGBEIN et al. (1998) observed lying cow–calf pairs in the first five days of life.
On the first day, the percentage of direct contact lying (distance < 3 m) was
65%. From the second day on, most cows distanced themselves from their
calves with the mean distance being 9 to 17 m with a maximum of more than
20 m measured. Only 20% of the observed pairs lay together in direct contact.
From the fourth day of life on, the distance between all studied pairs decreased
again with cows and their calves then spending 55% of the lying time close to
each other.
In the first two weeks, the calf rests a great deal, and the mother leaves it alone
to eat nearby. Gradually, the cow and calf return to the herd. Up to week 8,
calves band together in groups in which they rest, play and frolic about. The
mother is then only sought out for suckling, usually with the calf being prompted
for food. Contacts are also observed in which the young animal is only licked.
Calves call their mother when they are hungry. The mother usually responds
and goes to the calf. (SPENGLER NEFF et al., 2017)
At least once a day, calves are thoroughly licked by the mother in a suckler
herd (SAMBRAUS and STEINEL, 1978). When playing, calves bunt against the
willingly lowered head of the mother, which has been observed on pasture but
also in mother-bonded rearing in the barn with allocated suckling times
(SCHLEYER, 1998; WAIBLINGER et al., 2013).
Literature overview 33
Most interactions between the cows and their calves are initiated by the mother
(0.4 calls per hour from the cows vs. 0.02 calls per hour from the calves) with
the social contacts of the calves being directed increasingly towards calves of
the same age rather than their mothers. Mothers and their calves have more
contact with each other on average than other members of the herd but both
cows and calves have more contact with the animals of the same age than with
other members of the herd. (KILEY-WORTHINGTON and PLAIN, 1983)
Weaning
From four months of age, the calves reduce the number of times they suckle
during the day by about one per month, so that at eight or nine months they
wean themselves from their mothers (SCHEURMANN, 1974b; KUNZ, 2017).
Other authors consider weaning not to be actively driven by the calves but in-
stead the mothers wean their calves. Up to an age of one month, calves suckle
on the pasture as often as they want. As they become older, the dams limit
suckling to a suckling period in the mornings and the evenings. Calves are
weaned from their mother in that she interrupts the suckling and later no longer
allows suckling, that is, she fends off the calf. (WAGNON, 1963; HÜNERMUND,
1969; SAMBRAUS et al., 1978; REINHARDT and REINHARDT, 1981; KAPHENGST,
1984; RIST and SCHRAGEL, 1992; ŠPINKA, 1992; KEYSERLINGK and WEARY, 2007;
SPENGLER NEFF et al., 2017)
The weaning process by the mother animal is relatively abrupt in that she en-
ergetically boxes away the calf whenever it attempts to suckle and threatens it
so that within one to two weeks it abandons any attempt to suckle and is thus
weaned (RIST and SCHRAGEL, 1992).
The mother–child bond weakens considerably after weaning put does never-
theless persist, in some cases for years. In some cases, dams spend more time
with the yearling calves than the average for the herd and continue to lick them,
even when they are already caring for a younger calf. A long time after weaning,
cows and their calves recognise each other by olfactory, visual and acoustic
signals. (THORPE, 1965; SAMBRAUS et al., 1978; REINHARDT, 1980; KILEY-
WORTHINGTON and PLAIN, 1983; VEISSIER et al., 1990; SPENGLER NEFF et al.,
2017)
2.4.4.2 Social contacts of the calves
Suckling calves have a strong inclination to making social contacts and con-
gregate in their own sub-herds, known as ‘creche or kindergarten herds’ when
they are introduced by their mothers to the herd. When the cows distance
Literature overview 34
themselves, at least one cow supervises these groups of young animals. The
groups of calves play together and come together when fleeing. (SCHLOETH,
1961; SAMBRAUS and STEINEL, 1978; KOLB, 1987; CHENOWETH et al., 2014)
In studies by VITALE et al. (1986), the longest daily averages for the time the
calves spent with each other were recorded between day 11 and 40.
In the first eight weeks of life, SAMBRAUS and STEINEL (1978) and SCHLEYER
(1998) observed social contacts predominantly between the calves and their
mothers with fewer contacts with members of the herd of the same age. Playful
contacts between calves referred mostly to locomotor games and playful
headbutting. KILEY-WORTHINGTON and PLAIN (1983) report increasing contacts
between the calves up to three months of age, particularly mutual licking and
rubbing against one another.
RAUSSI et al. (2008) concluded from their observations that permanent relation-
ships between calves of the same age that they formed up to an age of 14 days
reduce aggressive behaviour and enable the calves to better handle new and
potentially stressful situations. Occasionally, relatively stable pair formations of
calves are observed in the choice of pasture and resting places (REINHARDT et
al., 1978a).
Pair rearing in the first weeks of life has benefits in terms of early socialisation
and the biological outputs of the calves. The calves are grouped from the first
day of life after receiving optimal colostrum supply and housed in pairs in large
hutches (about 3 m2 or larger). (KOCH, 2021)
Calves kept individually responded more strongly to environmental and social
novelties than those kept in pairs and spent more time exploring and in social
interactions with an unknown calf. Calves that were kept in groups with an older
companion responded more strongly to new surroundings than calves that
were kept in groups of the same age. (DE PAULA VIEIRA et al., 2012)
BUČKOVÁ et al. (2021) did not identify any negative effects of paired housing of
calves on their health, feed intake or growth compared to calves housed indi-
vidually. Their locomotion-related behaviours (activity, research) confirmed the
observations made by the authors cited above.
Calves are outside the hierarchy but are sometimes driven away by older cattle.
Until sexual maturity, the calves only have hints of dominance relationships
among each other. A hierarchy forms at the earliest between three to six
months. (SCHLOETH, 1961; SAMBRAUS et al., 1978; SAMBRAUS, 1985)
Literature overview 35
SCHLOETH (1961) observed a strong need among cattle for tactile stimulation,
particularly on the horns, head, neck and shoulders. Mutual licking is closely
related to comfort behaviour.
In a suckler herd, 23.3% of playful interactions of suckling calves ended with
submission of a calf. Mutual licking was only rarely noted. The most common
partner-related behaviours were dominance gestures followed by jumping, and
considerably less social licking and cross-sucking. (REINHARDT et al., 1978b;
SAMBRAUS and STEINEL, 1978)
2.4.5 Locomotion, exploratory, inquisitive and play behaviours
2.4.5.1 Locomotion behaviours
FRASER and DUNCAN (1998) differentiate between positive and negative moti-
vations that lead to different reactions. Negative motivations (e.g. thirst, fear)
are used for survival or reproduction and should be minimised as much as pos-
sible by the livestock manager. Beneficial activities such as play and explora-
tion are based on positive motivations and help with fitness and well-being.
They should be encouraged where possible.
The proportion of time not spent lying down depends in part on the age of the
animals but also on the substrate; with increasing age, the calves’ urge to move
decreases (SAMBRAUS, 1985). This was also suggested by observations by
RIESE et al. (1977) in which calves in a suckler herd moved a total distance of
550 m a day in the first month of life, a little more than 400 m in the second
month and less than 400 m in the third month.
METZ and METZ (1985) reviewed studies in which the behaviour of suckling
calves that were kept with their mothers was compared to that of calves sepa-
rated from their mothers and kept isolated. On the first day p.n., the suckling
calves were supported by the mother during the first attempts to stand up,
which meant that they stood up earlier and for longer. On the third day there
were no longer any differences recorded in the total time spent standing.
Results from KRACHUN and RUSHEN (2010) showed that locomotor play (more
than 3 s rapid forwards movements such as running, galloping, jumping) by
calves is reduced by a small volume of milk (6 L vs. 12 L full milk per calf and
day) and early weaning from milk (7 vs. 13 weeks). Because more energy is
needed for locomotor play and it occurs earlier in the development of calves
than social play, it can be used as a measure of the effect of the drinking regime
on the welfare of calves.
Literature overview 36
Mother-bonded calves (m-calves) played alone more often than off-cow calves
(o-calves) in a study by WAIBLINGER et al. (2013) However, the m-calves had
an extended cow barn available while the o-calves had considerably less
space. The m-calves therefore displayed intensive locomotor play in the cow
barn and less in the smaller, separate calf barn. In contrast, the calves without
mother contact often played together. In these group games, there were no
differences detected between mother-bonded and off-cow reared calves.
2.4.5.2 Exploratory, inquisitive and play behaviours
Exploratory and inquisitive behaviour is an urge to seek out and explore new
situations and to experiment with objects in the environment (EIBL-EIBESFELDT,
1963). Animals can learn, that is, they can individually adapt their behaviour
based on previous experiences (SCHEUNERT and TRAUTMANN, 1987).
Exploratory behaviour in higher vertebrates is a specific form of obtaining infor-
mation about their environment or about objects within their environment. In
more highly evolved animals, active exploration is a primary motivation, hence
it is referred to as ‘inquisitive behaviour’, which enables latent learning. It is
triggered by the novelty of stimuli. If a stimulus occurs, the animal explores
intensively, such activities subsequently flatten out and then cease. In this time
the new information is compared with existing information in the central nervous
system and new information is saved. When the same stimulus occurs again,
the animal responds with a weaker reaction. The presence of the mother pro-
vides security and the calves play and explore their environment without re-
serve. If the mother’s protection is absent, the young animals show fear of new
objects. A latent possible evasion behaviour is always present, which enables
the animals to quickly flee if necessary. (JOHST, 1975; WINCKLER, 2009)
The herd is both a haven and a constant resource for exploratory behaviour
during the development of the calves, the limits of which are only exceeded
with the onset of sexual maturity (MURPHEY; MOURA DUARTE; et al., 1981;
KAPHENGST, 1984).
An extreme form of evasion behaviour is flight. Domesticated cattle have a par-
adoxical tendency to both approach and simultaneously avoid humans. The
type of environment – familiar or new – of the animals certainly does play a
role. (MURPHEY; MOURA DUARTE; et al., 1981)
Cattle explore primarily by smelling and licking, that is, with nose and mouth
(VAN PUTTEN and ELSHOF, 1982) so that they approach with a lowered and ex-
tended head. Exploratory behaviour very closely resembles play behaviour and
can also transition into play behaviour. It appears to be pleasant and rewarding
Literature overview 37
because at times it goes beyond simply obtaining information. (WINCKLER,
2009)
Calves begin to explore their environment at a very early age, immediately after
birth even, and appear to learn very early and quickly. They are more curious
than adult animals and explore intensively on average for 3.2 minutes per hour.
When doing so, they smell the ground to identify potential food and their envi-
ronment to find protection. (KILEY-WORTHINGTON and PLAIN, 1983; KOLB, 1987)
Curiosity and play do not have a direct biological or social objective, do not
usually proceed in an ordered sequence of actions and are not completed with
a final action, such as eating after capturing prey. During play, behavioural el-
ements and motor patterns from across all functional areas are displayed and
freely combined with each other. (HINDE, 1970; SCHEUNERT and TRAUTMANN,
1987; SAMBRAUS, 1997b)
Play often gives an impression of a lack of seriousness because aggressive
movements are executed without the intention of injuring the playmate. The
motivation that triggers play differs fundamentally from the behaviours that are
being mimicked. Nevertheless, play is not merely the result of the absence of
highly intense motivational factors for another behaviour. (HINDE, 1970)
Play behaviour does not relate functionally to the current situation but occurs
spontaneously, is voluntary and appears to give the animal pleasure. Normal
sequences of behaviour are exaggerated and are frequently incompletely exe-
cuted and with harmless intention. Individual behavioural elements are fre-
quently repeated but not in the manner of a stereotypy. Animals only play in
stress-free, apparently safe situations, when they are healthy and in good con-
dition. (BURGHARDT, 2001)
Play sequences are often interrupted to start a higher priority behaviour. The
sequence of play also deviates from the mimicked purposeful behaviour, is ex-
aggerated and often not executed to completion. (FAGEN, 1981)
Play is well developed in cattle and occupies considerable scope. It appears to
be innate in terms of motivation, emotions, triggers and objectives. Play is a
behaviour that is only executed in and of itself and does not serve a purpose
but prepares animals for unpredictable situations. Healthy animals in an
adapted environment and in good weather play more often and for longer than
sick, hungry or cold animals. (SCHLOETH, 1961; THORPE, 1965; SCHEUNERT and
TRAUTMANN, 1987; PHILLIPS, 1993; ŠPINKA et al., 2001)
From a very early age, calves play with objects on the pasture, such as plants
and clumps of earth and rarely with moving objects such as stones and loose
Literature overview 38
pieces of branch (SCHLOETH, 1961). Play and exploratory and inquisitive behav-
iour are particularly important for gradual integration of the calves into the herd
(KAPHENGST, 1984).
MORGAN (1900) suspected that the animals familiarise themselves with the en-
vironment through play and that play is therefore of great importance for their
future well-being.
The euphemism ‘mock activity’ to describe the play of cattle makes it clear that
young animals execute playful patterns of behaviours, such as fighting, de-
fence, flight or sexual activities, which are used for survival but that differ from
these behaviours fundamentally. Thus, play components can be exaggerated
or repeated versions of these behaviours. Play reinforces social bonds and fa-
cilitates the cohesion of the herd. (BROWNLEE, 1954; PHILLIPS, 1993)
A calf tries to familiarise itself with all the details of its environment. Play activ-
ities are triggered by stimulus objects from the environment but also by the
inner drive of the young animals. On pasture, play behaviour is only apparent
at certain times of the day, particularly after feeding. (PORZIG, 1964)
Cattle also play as adult animals but this occurs more often in younger animals
(BROWNLEE, 1954; HÜNERMUND, 1969; SÜSS and ANDREAE, 1974; SAMBRAUS et
al., 1978; SCHEUNERT and TRAUTMANN, 1987; PHILLIPS, 1993).
Play behaviour is associated with positive emotions and boosts fitness in the
long term. It is a luxury behaviour with low priority in the hierarchy of behaviours
and is only executed if all other primary needs are met, which is why it is often
considered an indicator of well-being. A state of psychological and physical
well-being is a prerequisite for play. (BROWNLEE, 1954; SAMBRAUS, 1985;
PHILLIPS, 1993; MANTEUFFEL, 2006; WINCKLER, 2009)
SAMBRAUS (1985) emphasises the importance of play and warns against view-
ing it as a luxury that can be forgone. SCHLOETH (1961) sees a soothing function
in the play of adult cattle.
Play acts as a reward in animals and is possibly accompanied by the sensual
emotional state that is associated with the consumption of rewards. As a rule,
play is spontaneous because its output often appears independent of any ex-
ternal stimulation, it carries with it its own reward or serves its own purposes.
This means that it is displayed as an end in itself rather than to achieve an
ultimate objective, with the reward for the animal lying in the behaviour itself.
Play also seems to ‘infect’ other members of the group. (HELD and ŠPINKA,
2011)
Literature overview 39
The innate drive of calves to move is expressed in the herd in running and
chasing games, frolicking and galloping, combined with sudden stops and
changes of direction (Figure 3). Initially, calves play alone or with their mother.
Later, calves box and butt their heads against other calves (headbutting), rarely
against inanimate objects, and mounting or riding each other is also part of
play. One activity can be succeeded by another in rapid sequence and high-
lights the calves’ drive to move. Interactions between the calves do not funda-
mentally involve any threatening gestures and do not end with a playmate being
defeated or driven away, rather those involved frequently switch roles. Usually
calves play in the group and mutually encourage one another to play. Activities
of an individual animal rarely emerge in a group of calves. Although very young
calves initially play with their mother, later they prefer to play with other calves
of the same age. (SCHLOETH, 1961; PORZIG, 1964; KOCH, 1968; HÜNERMUND,
1969; SÜSS and ANDREAE, 1974; SANTHA, 1977; SAMBRAUS et al., 1978;
REINHARDT, 1980; KILEY-WORTHINGTON and PLAIN, 1983; SAMBRAUS, 1985;
PHILLIPS, 1993; WINCKLER, 2009; NEJA et al., 2017)
Figure 3: Play behaviour of calves
When jumping, the forelegs spring for-
ward and the hind legs follow with a
buck, and the animals often kick out
sideways when doing so. With increas-
ing age, the calves jump less often. In a
group, calves encourage others to play
by leaping about repeatedly in front of
each other. (SÜSS and ANDREAE, 1974;
SAMBRAUS et al., 1978; KILEY-
WORTHINGTON and PLAIN, 1983)
SCHLOETH (1961) observed young
calves playing ‘mother and child’ in Ca-
margue cattle kept in semi-wild condi-
tions. One calf approached another in
the posture it would use to approach its mother and started, with bunting move-
ments, to playfully suck in a relaxed manner between stomach and hindquar-
ters. In older young animals, the suckling posture occurred at the end of games
with sexual overtones. In isolated cases in suckler cow husbandry, sucking of
a hungry calf on another calf was observed (KILEY-WORTHINGTON and PLAIN,
1983). It can therefore be assumed that calves also cross-suck in mother-
bonded rearing but it is only rarely observed and described.
Literature overview 40
2.4.6 Behaviour in the circadian rhythm
Active and resting phases in animals reveal rhythms that are very important for
the temporal orientation of the animals. Certain expressions of animal life are
subject to a circadian rhythm with a period length of about 24 h. Examples in-
clude changing from a sleep to an awake state, the search for and intake of
food and a range of physiological functional factors (e.g. core body tempera-
ture, hormone secretion) (SÜSS and ANDREAE, 1974; SCHEUNERT and
TRAUTMANN, 1987).
The genetic rhythm of behaviour is synchronised by zeitgebers temporal exter-
nal or environmental signals, such as the light–dark cycle (SCHEUNERT and
TRAUTMANN, 1987). In cattle SAMBRAUS (1971) and SAMBRAUS and BRUMMER
(1978) observed a circadian rhythm that is determined by sunrise and sunset.
A fixed sucking time at daybreak was also observed by PORZIG et al. (1969),
WALKER (1962) and SÜSS and ANDREAE (1974) in 80%–100% of calves that
suckled on their mothers. After suckling, the calves grazed intensively (SÜSS
and ANDREAE, 1974).
In suckler cow husbandry, there are firm bonds between dam and calf. Every
calf in a herd of 20 Simmental cows and their young suckled predominantly
between 9:00 and 11:00 a.m. and 2:00 and 4:00 p.m. (RITTER and WALSER,
1965)
SAMBRAUS and STEINEL (1978) also noticed a circadian rhythm in drinking be-
haviour that is determined by the overall activity of the herd.
In the long days in May and June on pasture, KOCH (1968) observed that eating
times during the day were interrupted by four lying periods. All animals lay down
uninterrupted from nightfall until daybreak. When the day length decreased, the
adult animals gradually dropped one lying period. The calves, however, main-
tained the four lying periods for quite a while.
Calves played extensively on pasture in the summer evenings and occasionally
the cows also participated (KILEY-WORTHINGTON and PLAIN, 1983). Maremmana
calves in Tuscany also played extensively in the early morning, play activities
decreased around midday and increased again in the afternoon, with play oc-
curring mostly after grazing (VITALE et al., 1986).
An
ethogram
for
calves in
suckler cow husbandry on pasture is shown in
Table 1
(WEBSTER and SAVILLE, 1982). The behaviour of the calves was recorded with
cameras over 24 h and subsequently analysed. The suckling calves were ob-
served only at night when ruminating.
Literature overview 41
For observations of 10-week-old calves in a suckler cow herd, WEBSTER and
SAVILLE (1982) noted 13.1% lying and 7.4% locomotion behaviours, that is, ac-
tivity, over 4 h during the day. In 24 h they spent 46.4% or 11.1 h lying and
4.7% or 1.8 h engaged in activities without eating food.
Table 1: Ethogram of the natural behaviour of suckling calves in suckler cow
husbandry on pasture at 10 weeks, proportion of observed activities as a per-
centage over 24 hours
Activity Percentage ownership (%)
Lying 46.4
Rumination 12.4
Sleeping 19.5
Movement, Game, Headbutts 8.5
Source: according to WEBSTER and SAVILLE (1982)
2.5 Off-cow rearing of dairy calves
Rearing without cow contact affects the social competence of calves. Threat-
ening behaviour by a cow causes calves that have been reared with cow con-
tact to behave submissively more often than calves without cow contact. How-
ever, this does not cause hormonal stress responses (e.g. salivary cortisol con-
centration) in the motherless calves. There were also no indications in the be-
haviour of the calves that indicated emotionality. (BUCHLI et al., 2016; CANTOR
et al., 2019; NOEMI et al., 2020)
If calves can have contact with their mothers, they react to separation with a
lower milk intake and play less because of the reduced energy intake but make
increased vocalisations, which RUSHEN et al. (2016) interpreted as a sign of an
emotional response to the separation.
KENT and KELLY (1987) established that calves that had been separated for
three or more days from their mothers, having spent the first 24 h p.n. with
them, accepted any other cow as a foster mother, while cows took any other
calf into their care when contact between cow and calf was enabled immedi-
ately after the birth. The search for their own calf stopped when a cow became
foster mother to a foreign calf.
When discussing rearing of dairy calves with mother contact, it should be noted
that mother and calf must also be separated in this type of husbandry. This
version of calf rearing therefore cannot be advocated uncritically, particularly
as additional aspects must be considered.
Literature overview 42
Early separation of dam and calf, which must happen in milk production, leads
to stress that can, however, be reduced by half-day contacts so that the animals
gradually become accustomed to the separation, experience positive interac-
tions with humans, and the calves can learn to use the automatic feeders, which
prevents growth setbacks after weaning. Suckling cows produce less saleable
milk, can show problems with milkability and have milk with a lower fat content,
and the issue of disease transmission must also be considered. The udder
health of the cow can be positively affected by suckling, however. (JOHNSEN et
al., 2016)
HILLMANN et al. (2019) conducted surveys of farmers in Switzerland and identi-
fied no risk for the health of the calf as a result of cow contact. The authors
even see an opportunity for improving calf health in cow-bonded rearing.
Cows that do not suckle are possibly not strongly motivated to seek out contact
with the calf from which they were separated shortly after calving, which likely
can be attributed to the lack of excretion of oxytocin and endogenous opiates
that are released during suckling (WENKER et al., 2020). If this assumption is
confirmed by further studies, it can be concluded that the level of stress expe-
rienced by cows due to the separation from the calf immediately after the birth
is considered acceptable.
CANTOR et al. (2019) discuss in their article the need to investigate the long-
term effects of artificial rearing systems (individual, group, mother-bonded rear-
ing) on the behaviour, perceptions, outputs and health parameters of dairy
calves. Among the short-term effects of conventional rearing, the authors in-
clude the negative impacts on social behaviour, reduced feed intake, increased
activities in a novel environment and signs of malnutrition that are associated
with limited milk intake and poor growth during the weaning phase. Long-term
effects of conventional rearing may include reduced social submissiveness, in-
creased reactivity to new environments and output losses (milk and reproduc-
tive output).
The most common husbandry method for suckling calves following individual
housing in hutches or pens from two to three weeks of age is group housing in
deep-litter pens with exposure to the outdoor climate and facing south-east.
The pens must ensure optimal air exchange in accordance with the German
Ordinance on the Protection of Animals Kept for Farming Purposes (Ti-
erSchNutztV, 2002) and must be free of draughts, moisture and harmful gases.
To create a micro-climate, it is helpful to ensure that the rear and side walls are
closed to a height of 150 to 200 cm and to provide wind-proof panels on top.
The lying area should be 2–3 m2 per calf with a fixed standing and running area
Literature overview 43
that is 220–250 cm long. Overcrowding must be viewed critically, particularly
considering the microbial load, and must be avoided because it does not cor-
respond to optimal rearing. An animal to eating place ratio of 1:1 should be
complied with so that the calves can all eat at the same time at the trough from
week 8. (GLATZ, 2016; MÖCKLINGHOFF-WICKE, 2018)
A suckling calf is provided with milk at the right time, with the correct tempera-
ture and almost free of microbes, which can also be achieved using automatic
feeders. The role of the cow as a social partner for the calf cannot be provided
in off-cow rearing, however. Cows that suckle their calves after milking show
reduced alveolar milk ejection during automatic milking and the milk fat content
is reduced, which in turn affects the milk price. This study was not able to
demonstrate any effect of suckling on udder health. (BARTH et al., 2007)
A 5-minute contact was sufficient to create a strong, specific maternal bond
between the cow and her calf. After two hours’ separation, the cow still shows
signs of stress but can no longer recognise her own calf. If contact between
dam and calf is not permitted for up to 5 hours after the birth, the maternal bond
is no longer formed in 50% of the animals. (HUDSON and MULLORD, 1977)
The presence of the calf in the first few hours p.p. affects the behaviour of the
dam. The presence of the calf for cows that had calved with assistance led to
longer lying times. (HOUWING et al., 1990)
In the wild, a cow responds to separation from her calf with an increase in vo-
calisations. This also occurs if the cows are kept together with their calves only
for a few days and there is time for a bond to form. (KEYSERLINGK and WEARY,
2007)
Added to this is that in group calving, a newborn affects the multiparous cows
that are present, and calves can be licked by foreign dams because the multip-
arous cows are attracted to the smell of the placenta clinging to the calf
(HUDSON and MULLORD, 1977; PINHEIRO MACHADO et al., 1997; WILLIAMS et al.,
2001; UNGAR, 2006; KEYSERLINGK and WEARY, 2007).
If a calf is born in the group, lower-ranking animals are fended off by the mother
and higher-ranking cows are permitted unimpeded contact with the newborn
(PERREY et al., 1996). Adoption of a foreign calf and subsequent rejection of
her own calf has also been observed in isolated cases (HÜNERMUND, 1969;
PINHEIRO MACHADO et al., 1997).
RUSHEN et al. (2016) investigated stress factors caused by the separation of
calves from their mothers. The vocalisations of the calves and their play behav-
iour were evaluated as criteria for the reaction to the separation. Calves that
Literature overview 44
suckled from their mothers up to week 6 showed significantly less play behav-
iour (measured here by the frequency of jumping) in the first three days after
the separation and a significantly higher vocalisation than calves of the same
age that were fed by automatic feeders during the day and suckled from their
mothers at night and another group that was bottle-fed exclusively. Five or more
days after the separation, there were no longer any differences in behaviour
observed between the calves. The authors attribute the reduced play behaviour
to a stress-related reduction in energy intake by the suckling calves.
Heifers that were reared physically separated from each other sniffed and
licked their calves in the first three hours after the birth in the same way as
heifers reared in groups. However, they responded less to changes in the ac-
tivity of the calf and showed reduced maternal behaviour. (BROOM and LEAVER,
1977)
From the cited literature, the following conclusions can be drawn: because the
bond with the calf requires three days, the calf does not suffer any stress when
separated from its mother in the first few hours p.n., which is reflected in feed
intake and socialisation and consequently behaviour in the first weeks of life.
On the other hand, later separation causes stress, even if the calves are only
given occasional access to their mothers.
2.6 Behaviour and health
2.6.1 Behavioural enrichment
All agricultural livestock is capable of complex cognitive processes that play a
critical role in the emotional assessment of their husbandry environment. Mean-
ingful integration of species-specific and appropriate cognitive environmental
enrichment (behavioural enrichment) in the husbandry environment therefore
has the potential to achieve sustainable improvements in behaviour and the
welfare of the animals. (PUPPE et al., 2012)
Environmental conditions (enriched vs. low stimulus) have the greatest influ-
ence on the nervous system of young animals. They enable the animal to have
experiences that affect behaviour acquired by conventional learning or by
changing the reactivity of the nervous system. (GRANDIN and DEESING, 2014a)
Negative emotions arise with well-developed cognitive abilities with simultane-
ously reduced environmental challenge, e.g., boredom, frustration and behav-
ioural disorders. The meaningful integration of cognitive challenges (cognitive
enrichment) in the normal husbandry environment of livestock can achieve
Literature overview 45
sustained improvements in behaviour, well-being and animal welfare of the
husbandry system. (MEYER et al., 2010; PUPPE et al., 2012)
At the start of the 20th century, YERKES and DODSON (1908) demonstrated that
stimulation of moderate intensity has the most beneficial effect on learning suc-
cess.
This was followed by the allostasis concept developed by KORTE et al. (2007)
in which animal welfare must always be considered in relation to the degree of
existing environmental challenges. Well-being plotted against environment ef-
fect forms a reverse U curve (Figure 4). Capability is optimal for moderate stim-
ulation and deteriorates with both hypostimulation and hyperstimulation. After
repeated exposure to the same stress, habituation develops. Because of gen-
otype–environment interactions, the organism can show a hypersensitive
stress response, no habituation, a prolonged stress response or a hyposensi-
tive response. This model makes it clear that sustained hypostimulation or in-
adequate allostatic load due to low stimulus can lead to many stress-related
pathologies just as a high allostatic load can. (KORTE et al., 2007; MEYER et al.,
2010; PUPPE et al., 2012)
Figure 4: Animal welfare in relation to environmental challenges as shown by
the outdated concept-based homeostasis and the new concept based on the
inverted U curve of (di)stress.
Source: OESTERWIND et al. (2010), according to KORTE et al. (2007)
Literature overview 46
Various studies in pigs but also poultry, fattening cattle and calves demon-
strated the extent to which environmental enrichment leads to increased loco-
motor activity, due to the satisfaction of behavioural needs, but is also able to
induce sleep in animals or minimise behavioural anomalies, which in turn has
positive effects on animal outputs (MILLET et al., 2005; ISHIWATA et al., 2006;
PUPPE et al., 2007; WEERD and DAY, 2009; NINOMIYA, 2014).
It has long been acknowledged that animals have emotions. However, they can
only be experienced by an individual and can only be identified by scientific
investigative methods using symptoms in the realm of the probable (TSCHANZ,
1987). Such symptoms include body movements or facial expressions that can
be recorded and interpreted in experimental behavioural research using learn-
ing algorithms (SINGH et al., 2020).
Higher animals are aware of their emotions, which is why it can be assumed
that they seek to trigger positive emotions, which does, however, require the
presence of positive stimuli (MANTEUFFEL, 2006). For play behaviour that trig-
gers positive emotions (Chapter 2.4.5.2), environmental enrichment with toys
represents just such a positive stimulus.
JENSEN (1999) demonstrated in an experiment using the example of locomotor
behaviour in large spaces that young cattle use the opportunity to express
learnt play behaviour. If, however, food is offered, this draws them away from
the play opportunity.
Positive emotions of animals include satiety, vitality, reward, satisfaction, curi-
osity and playfulness while negative emotions include anxiety and boredom. If
positive emotions clearly dominate, it can be concluded that welfare is high.
(GREEN and MELLOR, 2011)
WEBB et al. (2019) refers to this state as animal happiness.
2.6.2 Ethopathies and behavioural disorders in calves
Behavioural disorders are indicators that a husbandry system is not animal
friendly, and it can therefore be expected that the animal will suffer harm in the
longer term (TSCHANZ, 1995). Studies of impaired behaviour are thus an im-
portant approach for revealing information about the welfare of livestock
(SCHEIBE, 1987).
Van PUTTEN and ELSHOF (1982) developed a list of behavioural patterns in
calves that express ‘abnormal behaviour’ that is caused by husbandry systems.
For calves kept in narrow pens, this includes sucking on objects and other
calves, tongue playing or rolling, intense licking of forelegs and shoulders,
Literature overview 47
nibbling on objects and the tails of other calves, dog-like sitting, vacuum activ-
ities, stereotypies such as hitting the head against the pen walls, stamping on
the spot or rolling the eyes. The authors reasonably ask why a remedy is not
sought when these behavioural patterns are apparent. SAMBRAUS (1985) also
summarises in short order that although technopathies and ethopathies are
known indicators, in many cases an appropriately designed husbandry system
was not created. Limitations cannot be overcome for livestock with every fence
representing a restriction on their freedom to move. Overstraining the ability of
the animals to adapt must, however, be prevented.
SCHEIBE (1987) pooled groups of causes of behavioural disorders and
measures together to deal with them. Congenital anomalies or behaviour insta-
bility (internal causes) can only be eliminated in individuals by culling and in a
population, selection is an option to breed out such problems. Behavioural dis-
orders due to external causes such as infections, metabolic disorders or injuries
can be treated using veterinary therapies or prevented using prophylaxis. Fur-
thermore, external causes are cited that negatively affect behaviour (e.g. low
stimulus, lack of space), offering inappropriate or no targets (agonistic behav-
iour) or through which a normal behaviour leads to harm (e.g. injuries due to
crowding, flight, panic). For the latter, behavioural disorders can only be pre-
vented by optimising the husbandry method and facilities.
SAMBRAUS, H.-H. (1993) classifies four different categories of behavioural dis-
orders: central nervous, deficiency related, endogenous and reactive; the au-
thor includes cross-sucking among the latter. For every single behavioural dis-
order, it should be checked whether there is a reaction to adverse husbandry
conditions.
2.6.3 Behaviour of sick calves
The behaviour of sick calves differs considerably from normal behaviour. Sick
animals are often thirsty, their appetite drops off, their external appearance
changes and they can no longer adjust to their surroundings. (NEJA et al., 2017)
Behavioural observations can therefore help to estimate the progression of
pneumonia and diarrhoea, for instance, because sick calves display changed
behaviour not only when the disease is clinically manifest but also beforehand
and during convalescence. Drinking and eating behaviour and synchronicity of
behaviour in the group in particular are important for early diagnosis. (BÜNGER
et al., 1988)
Literature overview 48
Animals feel pain but occasionally this is difficult to identify and difficult to check
by visual observation (BRINKMANN, 2016). Pain leads to physiological changes
such as a change in heart rate, blood pressure, body temperature, weight or
food intake but also to changes in behaviour or responses to interactions with
humans. Recognisable signs of pain vary across species and individuals and
must always be considered as complex and viewed in context. Cattle can vo-
calise (grunting, bellowing), grind teeth, move hesitantly, change the expres-
sion of the eyes and head posture and show a decrease in outputs.
(UNDERWOOD, 2002)
The overall impression in pain states can be recognised by a rigid and lowered
head–neck posture, a tensed back or protection of extremities when standing
(SÜSS and ANDREAE, 1974).
GRAUVOGL (1983) cites shrill cries or howling, almost soundless groaning,
pressing the mouth shut, grinding teeth and sweating as expressions of pain.
Respiratory and heart rate are clearly accelerated and unmotivated turning and
bending of the body, lameness, attempts to obliterate the pain, such as by kick-
ing against the abdomen or kicking out with the affected limbs, are also typical.
Additional signs are apathy and sensory disorders, expressed as sitting with a
lowered head and hanging ears = ‘mourning’. All these signs can be objectively
recorded to varying degrees.
RADEMACHER (2003) adds that cattle with moderate to severe colic-related pain
tend to display increased movements such as frequent, violent beating of the
hind legs on the abdomen when standing and lying, repeatedly lying down and
standing up, thrashing and rolling.
Although in some cases a single indicator, normally a behavioural indicator, is
sufficient to identify pain, the combination of different types of indicators in-
creases the sensitivity and specificity of the pain assessment (PRUNIER et al.,
2012).
To reduce the negative impacts of diseases, early detection would be useful.
BELAID et al. (2019) developed a predictive model in which the behaviour of
male fattening calves aged from 30 to 85 days were assessed up to 10 days
before an illness. The most reliable prediction of a diagnosis was provided by
a multivariate logistic regression model of the visits to the feed trough, number
and duration of lying periods and number of steps. Sick calves took fewer steps
and visited the feed trough less often. There were no differences identified in
the lying time between sick and healthy calves.
Literature overview 49
Observations of the behaviour of calf pairs published by BERTELSEN and JENSEN
(2019) showed a relationship between the health of a calf and frontal pushing.
A healthier calf initiated more social play than the partner with worse health,
which is why the authors believe that differentiating between play elements can
be important when using calf play behaviour to evaluate animal welfare.
2.7 The genetics of behaviour
Behavioural sequences are made up of coordinated motor patterns that are
based predominantly on genetically determined connections in the central nerv-
ous system. These locomotor elements are often species-specific (SCHEUNERT
and TRAUTMANN, 1987) and according to LORENZ (1978) are classified as fixed
action patterns.
It is undisputed that behaviours can have an inherited, usually polygenetic com-
ponent that is studied in behavioural genetics. What is important when estimat-
ing the genetic effects are the genotype–environment interactions. (HÖRNING,
2008)
The behavioural phenotype is determined by the genotype–environment inter-
actions that an individual is exposed to during its development. Selection not
only affects outputs such as earlier sexual maturity, it also changes behaviour,
such as aggression, which ultimately differentiates domesticated animals from
their wild forms. However, the environment (separation by sex, individual hous-
ing, small social distance) and handling (feeding, dehorning, castration, artificial
insemination) also cause changes in behaviour that become established ge-
netically. (KLINGHAMMER and FOX, 1971)
In addition to its potential contribution to improving animal welfare, examining
the genetics of behaviour in cattle can provide further general insights into the
genetic control of such complex characteristics (GUTIERREZ-GIL et al., 2008).
Behavioural characteristics have been selectively manipulated since the begin-
ning of domestication and such manipulation provides good opportunities for
specifically influencing the welfare of animals (BUCHENAUER, 1999; WIENER,
1999; SCHUTZ and PAJOR, 2001; BRADE, 2002), which is particularly apparent in
refined dairy and beef cattle breeding. For example, along with many other
characteristics, the milking behaviour of dairy cows is also systematically con-
sidered in dairy cattle breeding. The periodic feeding and suckling behaviour of
calves, social behaviour within the hierarchy and the dam–calf bond, resting
and reproductive behaviours as well as temperament are genetically deter-
mined. Behaviour always results from a genome–environment interaction,
Literature overview 50
which has led to an increase in the variability of behaviour within a species over
the course of domestication. (BRADE, 2003; BRADE and BRADE, 2017b, a)
Behavioural control is multifactorial and, along with the genetic predisposition,
environmental factors, cognitive capabilities, and age and sex are also involved
in triggering and controlling complex behavioural characteristics. Breeding
therefore cannot resolve all behavioural problems that emerge. (BRADE, 2003)
The genetically determined willingness of animals to act allows the emergence
of behaviours in their repertoire, even if these behaviours are not appropriate
under the given husbandry conditions (SCHEIBE and BILDT, 1987).
The temperament of an animal is subject to complex genotype–environment
interactions and is influenced by genetics and the environment but also by
learning. Changes to a characteristic can have unexpected effects on other,
apparently unrelated characteristics, which is why concentrating on selecting
single features can lead to unwanted changes in other behavioural and physi-
ological features. Selecting for temperament as a single behavioural character-
istic in foxes, for example, led to unexpected changes in fur colour, oestrus
cycles, hormone profiles and subsequent changes to physical characteristics.
(GRANDIN and DEESING, 2014a, b).
For temperament, the most valued characteristic in terms of genetics (SCHUTZ
and PAJOR, 2001), a heritability of h² = 0.21–0.61 was determined (GAULY et al.,
2001). The temperament associated with weaning of calves was estimated by
LITTLEJOHN et al. (2016) as h² = 0.28–0.35. LUCENA et al. (2015) determined the
heritability of temperament as being h² = 0.21–0.26 (p < 0.05), with the correla-
tion between weaning weight and temperament being r = -0.33 to -0.34
(p < 0.05). HÖRNING (2008) summarised several studies by various authors on
the heritability of temperament in which figures range from h² = 0.08 to 0.6.
MENDOZA (1986) calculated heritability of the drinking behaviour of calves of
h² = 0.43 ± 0.21 for the drinking duration (time in s to drink 1 L liquid) and
h² = 0.52 ± 0.1 for the volume of liquid drunk.
This corresponds to studies by DAS et al. (2001) that identified breed-related
differences for several behavioural aspects, such as suckling behaviour (dura-
tion and bunting) and maternal behaviour, in pure-bred zebus and their cross-
breeds compared to an endemic African breed.
The effect of the described genotype–environment interactions is supported by
LE NEINDRE et al. (1979). The authors compared the intake of colostrum by
Salers calves in mother-bonded beef cattle breeding and Friesian calves that
are normally separated early from their mothers. Friesian calves accepted
Literature overview 51
colostrum administered by bottle significantly more frequently than Salers
calves (82% vs. 39%, p < 0.005). Suckled Salers calves ingested a significantly
higher quantity of colostrum than calves fed with a bottle (8.3% vs. 6.3% live
weight, p < 0.025). In the first few hours of life, separated Friesian calves were
considerably more active, stood up earlier and slept for shorter periods than
separated Salers. However, the Salers were more active with their mothers
than the Friesians. Therefore, Salers calves appear to be less well adapted to
the conditions of dairy husbandry than Friesian calves. For the Salers, the pres-
ence of the mother is essential while the Friesian calves did not appear to miss
her. The conditions of dairy husbandry have possibly led to selection in these
breeds that has directly affected their behavioural characteristics.
From studies of aggressive behaviour of dairy cows, GIBBONS (2009) concluded
that the correlating effects of selective breeding on behavioural characteristics
should be kept in mind. SCHUTZ and PAJOR (2001) assume that it should be
possible to consider temperament in the breeding objective. In Sweden and
Denmark it has long been part of the estimation of the breeding value (BRADE,
2003).
The variance in behaviour between individuals also enables targeted selection
(HÖRNING, 2008). Studies by HOPPE et al. (2010) indicate that genetic selection
of temperament characteristics does not affect production characteristics such
as the average daily weight gain of calves.
The use of genetic markers to determine the reproductive performance in ani-
mals with different behavioural patterns was of major importance for
HUNTINGFORD (1984) in behavioural research. The quantitative trait loci (QTL,
genetic location of a quantitative characteristic (GELDERMANN, 1976;
ANDERSSON, 2001; GOMERINGER, 2007)) have long been identified for various
behaviours; the QTL for temperament, for example, has been mapped to chro-
mosome 29 (SCHMUTZ et al., 2001; BALL et al., 2002; HIENDLEDER et al., 2003;
GUTIERREZ-GIL et al., 2008).
2.8 Animal welfare, animal friendliness and well-being
The discussion in Germany about improving animal husbandry started with the
requirement to consider the welfare of agricultural livestock. Consequently, the
German Animal Welfare Act came into effect on 29 July 1972.
Article 2 of the Animal Welfare Act states that any person who keeps, cares for
or is required to care for an animal must provide the animal with food, care and
housing appropriate to its species and needs; may not restrict the animal’s
Literature overview 52
opportunities for species-specific freedom of movement to such an extent as to
cause the animal pain or avoidable suffering or harm; and must possess the
knowledge and skills necessary for providing the animal with adequate food,
care and housing in accordance with its behavioural requirements.
According to CARENZI and VERGA (2009), animal welfare must adopt a scientific
position to be able to play a specific role in animal science. The methodology
based on applied ethology, which enables a comprehensive view of the entire
organism and its adaptability, needs to be validated.
The BRAMBELL COMMITTEE commissioned by the British government in the
1960s emphasised the need to consider the standards of ‘welfare’ of domesti-
cated animals, which includes both the physical and mental well-being of the
animals. Any attempt to evaluate well-being must consider current scientific
understanding of the feelings of animals that can be derived from their structure
and functions as well as from their behaviour. (BRAMBELL, 1965)
Modern animal welfare laws, including that of the Federal Republic of Germany,
are based on fundamental ethical principles. The aim of the German law is to
protect the life and well-being of animals; no-one may cause them unjustified
pain, suffering or harm. The analogous law in Switzerland also protects against
frightening animals. This law obliges all animal keepers to treat animals in such
a way that their needs are met as much as possible, which includes food, care
and housing. To implement and monitor these laws, and to enforce them if nec-
essary, requires information that enables the situation for an individual animal
to be evaluated. One option is to compare the situation with animals that are
kept in a natural environment. (TSCHANZ, 1987)
MCGLONE (1993) believed that animals are only in poor condition if the physio-
logical systems are so greatly impaired that survival or reproduction are nega-
tively impacted. This would mean, however, that if animals continue to produce
or grow, they would also have well-being. Intensive breeding for high outputs
has led, however, to animals continuing to produce even when their physical
fitness is compromised (THOMPSON, 2016).
Good animal welfare starts with physical health, but good well-being means
much more, specifically animals need to also experience positive emotions
such as pleasure and satisfaction but not negative emotions such as fear, frus-
tration or suffering (DAWKINS, 2006). Subtle awareness of the behaviour of the
animals and close observation of any deviations provide an opportunity to iden-
tify suffering (LOEFFLER, 1990). To measure well-being, VAN PUTTEN (1982)
called for observation of animals and recording the number of stereotypical
movements, among other features. According to MILITZER (1990), specific
Literature overview 53
activities such as fighting, relaxed lying and playing as well as behavioural dis-
orders and stereotypies are suitable to use as assessment criteria for well-be-
ing, provided that methodological and biological limitations are considered be-
cause as an expression of animal condition they can be interpreted more mean-
ingfully than biochemical data or clinical parameters measured by veterinari-
ans.
DUNCAN (1993) recommended determining what animals feel under the given
husbandry conditions. Negative feelings must be reduced while also attempting
to maximise positive feelings.
Husbandry systems and management practices that are associated with suf-
fering or pain are counterproductive for the effectiveness of production
(UNDERWOOD, 2002). Higher welfare rearing, in which calves feel comfortable
and are not exposed to avoidable stress, is definitely in the interests of agricul-
turalists, which has been recognised and also implemented in many instances
in recent years.
TSCHANZ (1984) differentiates between properties of the environment that relate
to the terms ‘species appropriate’ – food and care – ‘species specific’ – behav-
iour, life phases, needs – and ‘behaviourally appropriate’ – behavioural pat-
terns. ‘Species appropriate’ and ‘behaviourally appropriate’ refer to conditions
that can be derived from the characteristics of the animals. The author strongly
urges that animal-friendly husbandry must not only incorporate housing design
that is behaviourally appropriate and feed that is species appropriate but should
also provide animals with species-specific stimuli.
A husbandry system can be considered species appropriate and animal
friendly, according to STUBER (1980), when the totality of the associated facili-
ties, regarding shape, dimensions and physical properties of the materials
used, satisfies the needs of the animals for unimpeded progression of species-
specific life processes (welfare). SAMBRAUS (1985) has a similar view, charac-
terising housing as animal friendly if the animal can express essential behav-
ioural needs across every functional area. He states that although ethological
knowledge and technical options are sufficient to provide animals with appro-
priate housing conditions, in many cases this is not done. Economics often
plays a role here.
For BAUMGARTNER (1997) a husbandry system is animal friendly if the animal
receives what it needs to achieve self-sustenance and self-maintenance and it
can meet its needs and avoid harms by having the opportunity to express ap-
propriate behaviour. For SUNDRUM et al. (2004), the reactions of animals to their
Literature overview 54
current husbandry environment are an essential criterion for evaluating the an-
imal welfare of husbandry conditions.
To ensure well-being, all needs must be considered. The degree to which
needs are satisfied and the level of frustration can be evaluated by the intensity,
duration and frequency of output criteria and parameters of behaviour and/or
(patho)physiology. (BRACKE and HOPSTER, 2006)
The concepts and definitions of well-being make it clear that the ability to feel
pain and pleasure, cognitive processes or consciousness are central points in
the question of the value of the animal compared to humans and the resulting
moral obligations of humans towards animals (KRIJNEN, 1999; ÖZMEN and NIDA-
RÜMELIN, 1999; WEBER and VALLE ZÁRATE, 2005).
The animal friendliness of husbandry methods cannot be directly evaluated and
indicators are needed to confirm the level of welfare. REINHOLD et al. (2017) call
for a uniform definition of the term ‘animal welfare’ as well as specifications for
its measurement and evaluation.
SMIDT (1990) differentiates between indicators relating to the animal, technol-
ogy and the livestock manager. These can be measured or verified and thus
easily monitored or evaluated. BRINKMANN et al. (2016) developed a practical
guideline based on such parameters that can be used as a working basis for
animal keepers of dairy cows, breeding calves or beef cattle to verify that ani-
mal welfare standards are being met.
Husbandry-related indicators include parameters such as the space provided,
the ratio of animals to feeding places, the type of lying surfaces, and the pres-
ence and type of runs. Animal-based indicators are understood to be, for ex-
ample, time spent lying and eating, injuries, movement scores (lameness) but
also agonistic behaviour and behavioural disorders. These indicators must be
valid, able to be reliably recorded and practically implementable on an opera-
tional level. (WILLEN, 2004; LIEBHART, 2009; ZIPP, 2015; BRINKMANN et al., 2016)
SUNDRUM (1995) counts changes in the intensity and frequency of the occur-
rence of behaviours and behavioural disorders among the ethological indica-
tors for animal well-being and animal friendliness.
Indicators relating to the livestock manager include caring for and nurturing the
animals but also their management. In surveys of livestock managers, some
responses indicated that inadequate measures were applied when caring for
calves (e.g. various substances administered to calves with sucking problems,
additives in the milk, milk diluted with water), which suggested that acquiring
Literature overview 55
technical knowledge did not teach adequate conceptual implementation.
(LÜDTKE, 2004)
In Germany the Animal Needs Index (ANI) 200 was developed as a user-
friendly instrument to evaluate housing systems. It aims to identify weaknesses
and deficiencies so that animal husbandry can be improved. The evaluation
criteria are limited, however, to measurable parameters and do not consider
the behaviour of the animals. (SCHNEIDER, 1996)
SCHRADER et al. (2006) presented the legally non-binding ‘National Evaluation
Framework for Livestock Management’ for agricultural animals regarding their
welfare, which was developed on behalf of the German Federal Ministry for
Food, Agriculture and Consumer Protection and the Federal Ministry for the
Environment. For animal behaviour, the extent to which the structural and tech-
nical features of the husbandry system limit the behaviour of the animals was
evaluated, using indicators and based on a catalogue of criteria.
TSCHANZ (1995) evaluated the animal welfare of a husbandry system based on
the type and norm for the individual. The design features and the behaviour as
well as the degree to which individual characteristics are expressed must cor-
respond to the breed type. The norm is derived from the variability of the ex-
pression of characteristics and corresponds to the average of the population.
Well-being includes both health and an unimpeded, species-appropriate and
behaviourally appropriate progression of life processes (WENNRICH, 1978). Ac-
cording to LORZ and METZGER (2019), well-being is a state of physical and men-
tal harmony within an animal itself and with its environment. Regular signs of
well-being are health and behaviour that is normal in every regard.
Good welfare goes beyond purely physical health. In simple terms, ‘good wel-
fare means that animals are healthy and have what they want’. This intuitive
definition has the advantage of capturing what most people understand by ‘im-
proving welfare’ and avoids a potentially dangerous distinction between scien-
tific and non-scientific definitions of welfare. (DAWKINS, 2008, 2016)
According to FRASER (2008), the concept of animal welfare includes various
components that can be summarised under three primary complexes: health
and functionality, affective states, and a natural life (Figure 5). These com-
plexes overlap to a degree, meaning that satisfying one criterion does not nec-
essarily ensure a high degree of welfare when evaluated from the perspective
of the other criteria. Thus, free-range husbandry provides the opportunity to
experience natural behaviour but simultaneously carries a risk of infection by
Literature overview 56
parasites and exposure to harsh weather. (LUND and ALGERS, 2003; FRASER,
2008)
Figure 5: Three concepts for animal welfare
adapted from FRASER et al. (1997); APPLEBY (1999); LUND and ALGERS (2003);
FRASER (2008)
According to SANDØE et al. (2003), evaluations of animal welfare at an opera-
tional level must define which parameters have to be measured. If these are
added to form a composite parameter, it raises the question of whether to be
guided by the average, whether the worst value must be used as the criterion
for evaluation, or whether there is a baseline or a threshold value between still
and no longer acceptable.
A particularly good indicator for impaired well-being is also reduced or modified
immunocompetence, which directly correlates with animal health. To evaluate
the animal welfare of husbandry systems, ethological and physiological indica-
tors are suitable because changes in these indicators often occur well before
pathological clinical symptoms and diseases manifest. (STEFANSKI, 2016)
There are no husbandry systems in which the harmony of the animal with its
environment is perfect. An animal feels comfortable in its environment when it
lives in appropriate harmony. This is always the case even if stimuli in the en-
vironment are perceived by the animal as unpleasant but are still acceptable
and do not lead to disharmonious behaviour (VAN PUTTEN and ELSHOF, 1982).
However, the best ‘alternative’ production system is always a compromise be-
tween the requirements for efficient production and animal welfare. This re-
quires a discussion not only of animal welfare but also about a feasible com-
promise. Simply measuring well-being does not achieve anything and leads
only to endless debate. (JENSEN and SANDØE, 1997)
Literature overview 57
Good welfare is achieved if the expectations of an individual accord with per-
ceptions of the internal and external environments, and the individual has cop-
ing strategies or mechanisms that appropriately eliminate any difference or that
enable adequate adaptation to the specific environmental situation. Under hu-
man control, it would be possible to protect animals from all challenges that
possibly cause stress. A good life is, however, not a life without challenges,
provided that such challenges do not overwhelm the capabilities of an individ-
ual. Living a ‘natural’ life includes sustained states of stress for most animals.
(SACHSER, 2001; STEFANSKI, 2016)
DUNCAN (1993, 2005) explained that neither health nor a lack of stress or fitness
is necessary and/or sufficient to conclude that an animal is experiencing good
well-being. Well-being depends on what animals feel. Preference tests and mo-
tivation tests can investigate suffering as an expression of strong negative feel-
ings and pleasure as an expression of positive feelings in animal husbandry.
Using livestock ethology, husbandry conditions and management measures
can be reviewed for their animal welfare (VON BORELL, 2009). Recognising and
considering behaviour is a prerequisite for animal-friendly design of housing
and feeding (SCHRADER and MAYER, 2005).
Animal-friendly husbandry systems should contain elements that increase at-
tentiveness and exploratory behaviour. Routine workflows, such as providing
care or mucking out barns, should be done regularly and consistently, and an-
imals must be able to react appropriately to spontaneous changes in their en-
vironment through their behaviour. (STAUFFACHER, 1994)
Animal welfare costs money but does increase the efficiency of animal produc-
tion and improves working conditions and occupational health and safety. It
often also guarantees bonus effects for greater cost effectiveness. (HARMS,
2021)
SAMBRAUS (1985) derived the following suggestions for rearing calves from be-
havioural studies:
1. Calves must live in direct contact with other calves because only group
housing satisfies the need for social contacts.
2. Satisfying calves’ strong need for movement and play requires large areas
in group pens.
3. The stock density must enable sufficient space for lying down and standing
up.
4. The drive to suck must be satisfied by milk intake. Roughage must be of-
fered from a few weeks of age on.
Literature overview 58
2.9 Animal welfare and ethics
Since about the 1960s there has been a change of thinking regarding animal
husbandry that has flowed into the process of creating animal welfare legisla-
tion (TEUTSCH, 1982).
Emotional well-being of animals has only been intensively researched for about
the last two decades, research that is a basic requirement for assessing the
quality of life of animals. For a long time, animal welfare research focused on
potentially negative situations for animals, such as the effect of procedures (de-
horning of calves) or husbandry conditions, or behavioural disorders such as
stereotypies (cross-sucking in calves). There was less consideration given to
behaviours that indicate that an animal is not only not suffering but that it also
feels good. Play behaviour, particularly by young animals, is just such an ex-
ample and much more attention has been devoted to this area in recent years.
The new ‘quality of life concept’ expands the perspective of animal welfare re-
search and asks questions such as: is there a ‘degree of suffering’ that must
never be exceeded and that can never be compensated for by positive experi-
ences? How much in the way of positive experiences does an animal need in
life? Should animals be provided with a ‘life worth living’ or does a ‘good life’
suffice? Are intensive ‘highs’ desirable or are less positive but sustained con-
ditions sufficient? Is a switch between positive and (slightly) negative experi-
ences required? (WINCKLER and HINTZE, 2019)
Finally, reference must be made to an issue that SANDØE et al. (2003) ad-
dressed in their article on the relationship between ethical questions and the
evaluation of animal welfare. The following four key ethical questions are men-
tioned:
Is there a basic standard for morally acceptable animal welfare?
How is a good animal life designed?
Which agricultural production objectives are justified?
Which compromises are acceptable in animal husbandry?
What is considered acceptable animal welfare includes not just evaluating how
well animals fare in a particular production system but also evaluating the pur-
pose of the production, which requires that a compromise between animal wel-
fare and other social interests be considered (SANDØE et al., 2003).
Bearing responsibility means defending one’s actions and facing the conse-
quences. Animals may only be killed for sensible reasons, and they have no
ability to understand their origin or future. The needs of livestock must be
guided by the requirements of the livestock and must be animal friendly. It is
Literature overview 59
not correct, however, to anthropomorphise animals and to adapt them to hu-
man needs. (BREM, 2019)
Over the past decade, animal protection and improving animal welfare have
become an increasingly important component of animal husbandry systems
and animal food supply chains in many economically well-developed countries
around the world. If, according to the recommendation by the United Nations,
improved animal welfare as a goal consists of becoming a significant and rec-
ognised part of the global efforts towards more sustainable agricultural devel-
opment, the following aspects must be implemented:
1. Animal welfare must be an integral part of sustainability.
2. Animal welfare science and policy must articulate emerging concerns re-
garding the relationship between health and well-being of humans and an-
imals.
3. The animal welfare community needs appropriate representation within in-
ternational governmental structures, which are linked to sustainable agri-
cultural development.
4. At an international level, robust and comparable standards for the welfare
of livestock are required across a range of production systems.
5. Animal welfare science must respond to new scientific developments and
political challenges. (BULLER et al., 2018).
Animal welfare and cost effectiveness must not be in opposition (BADERTSCHER
FAWAZ, 2003).
Beyond simply complying with laws, ensuring the well-being of livestock in
many industrial countries has become a quality criterion for animal products.
Considering these new global agendas, the task for a broader animal welfare
community consists of providing healthy food produced in a sustainable man-
ner, which is equally important for both humans and animals. (BULLER et al.,
2018)
Research into animal welfare can be considered an instrument to improve our
knowledge of animals regarding their physical and mental aspects. Ethics can
help to develop a principle for how animals must be treated and used. It should
be combined with the science because scientific knowledge affects the per-
spective of ethicists, and scientists can be influenced by ethical values.
(CARENZI and VERGA, 2009)
Materials and methods 60
3 Materials and methods
The behaviour of female German Holstein calves in intensive off-cow rearing
was investigated in collaboration with an agricultural enterprise in Mecklenburg-
Vorpommern.
3.1 Study farm, housing and feeding of the calves
All calves were kept in hutches with a run after initial care up to the end of
week 4. The move to the subsequent group housing was continuous from week
3 to a pen with a litter-covered lying area and concrete running area (Table 2,
Figure
6 and Figure
7). The maximum age difference within a group was
4 weeks.
The maximum daily calf milk replacer feed-allowance (CMRA) of 12 L calf milk
replacer feed (CMR-feed) was offered up to day 49 (Table 2, rounds 1 to 5) and
on day 106 the calves were weaned (rounds 1 and 2). Over the entire feeding
period, a powdered calf milk replacer (CMR) containing 50% skim milk fraction
was used.
Table 2: Husbandry and feeding plan
Single keeping Maximum until 4th week of life
Form of husbandry Igloo with running area
Calf feeder Mixed colostrum/Transition milk 2 x daily
as needed
Group housing From the 3rd week of life
Form of husbandry Lying surface with,
running surface without bedding
Maximum number of calves per group 15, age difference up to 4 weeks
Maximum CMR-feed-allowance 12 l CMR-feed up to the 49th day of life
Weaning 50th – 105th day of life of life
Weaned 106th day of life
CMR-feed-concentration, g CMR per l 160 / l water
CMR = calf milk replacer
Table 3: Period of feeding and quality of solid feed
Hay From week 1 1st cut, 8.7 MJ ME, 124 g crude protein
/
kg T
Dry-TMR
TMR
From week 1
From the 6th week
10.7 MJ ME, 160 g crude protein
/
kg FM
37
% T; 10.8 MJ ME, 155 g crude protein
/
kg T
week = week of life, MJ ME = Megajoule Metabolizable Energy, FM = fresh matter,
T = dry matter, TMR = total-mixed-ration; g = gram, kg = kilogram
Materials and methods 61
Figure 6: Calf group pen with lying and running area, hay rack, dispensing sta-
tion, feeding trough, toy (brush, chairs and ball)
Figure 7: Schematic of the calf group pen with lying and running area, hay rack,
dispensing station, feeding trough, toys (brush, chains and ball) and dummy
teats as well as location and field of view of the cameras
Materials and methods 62
As supplementary feed, from week 1 hay and dry total mixed ration (TMR) was
available fresh daily ad libitum in bowls in individual housing and in hay racks
and troughs in the group housing. From week 6, a TMR gradually replaced the
dry TMR (Table 3, Figures 8 (a) – (d)). Water was also available ad libitum in
bowls in the individual housing and an automatic drinker in the group housing.
(a) (b) (c) (d)
Figures 8: (a) – (d): Drinking and feeding facilities in the calf pen: (a) CMR-feed
dispensing station, (b) automatic drinker, (c) trough and (d) rack
3.2 Data acquisition and processing
The behaviour of female suckling calves in group housing was continuously
recorded 24 hours a day from November 2017 to December 2018 using cam-
eras in the VisoTech surveillance system from the day the first calves were
placed in the pens until the day the last calves left (Figure 7).
The group pen was fitted with various elements to investigate which activity
options were preferred by the animals. A tyre freely hanging on a rope with a
ball (ball), a cattle brush for calves (brush) and board with four chains (chains)
were installed in the pen as toys (Figure 7, Figures 9 (a) – (c)).
In rounds 1 to 3, a calf feeder with ten teats was hung on the grate to the right
of the automatic drinker and was used to provide the animals with electrolytes
(Figure 7, Figure 10 (a)). The feeder stayed empty in the pen during the day
and thus was available as a dummy teat. In rounds 4 and 5, it was investigated
whether the calves use the dummy teats which did not provide them with any-
thing (boards with four blank plugs made of plastic, Figure 10 (b)). These were
fixed to the wall to the left of the dispensing station and on the gate to the open
area (Figure 7) so that they could also be used at night.
Materials and methods 63
(a) (b) (c)
Figures 9: (a) – (c): Toys: (a) ball, (b) chains, (c) brush
(a) (b)
Figures 10: (a), (b): (a) ‘Milk Bar’ calf feeder, (b) dummy teats
For the statistical analysis, the focal sampling method (HOY, 2009b) was used
to code the behaviour of selected calves recorded in the videos (n = 13) in the
Interact program from Mangold (Figure 11, Table 4).
A total of five rounds were evaluated for 13 calves over 212 days with 4,569.5
hours of video recording (Table 4, Table A
1). For nine calves their behaviour
was observed from week 3 to 7 with a maximum allowance of 12 L MR per
animal and day, and three of these calves and two others were observed during
weaning from week 8 to 15.
‘Resting’ was coded when the calves stayed on the ground for at least 1 minute
after lying down before standing up again. Resting, sleeping and ruminating
were not differentiated here (Table 5).
When coding the blind visits, it was not possible to differentiate between re-
warded and unrewarded visits.
Materials and methods 64
Table 4: Number of evaluated calves, days and hours per week
Age in Video evaluations, numbe
r
of
Week Days Calves Days Hours
3 15 bis 21 1 6 127.3
4 22 bis 28 4 14 270.9
5 29 bis 35 8 32 765.8
6 36 bis 42 824 685.0
7 43 bis 49 929 742.7
8 50 bis 56 417 239.0
9 57 bis 63 5 20 278.8
10 64 bis 70 4 15 262.6
11 71 bis 77 3 9 209.1
12 78 bis 84 311 256.9
13 85 bis 91 211 249.3
14 92 bis 98 4 16 327.4
15 99 bis 105 2 8 154.8
Total 212 4,569.5
Number 13
Table 5: Behaviour categories and their coding
Behaviour Coding of behaviour
Rest, sleep, lie down
Without and/or with rumination
Rest
Ingestion CMR-feed-intake
Solid-feed-intake (hay, dry-total-
mixed-ration, total-mixed-ration)
Water-intake
Visit at dispensing station without
CMR-feed-intake (blind visit)
Social behaviour or
Contacts between calves
Calf-calf-contact with licking and
smelling
Gaming behaviour Game with toys: ball, brush, chain,
Head butting
Exploratory behaviour Lick and smell objects
Sucking activities Cross-sucking, suckle on another calf,
suck on dummies
CMR = calf milk replacer
Materials and methods 65
Figure 11: Scheme of coded behavioural categories
(CMR = calf milk replacer)
3.3 Statistical analysis and presentation of the results
The recorded and processed data were statistically prepared using Interact
(Mangold) and Excel 2019 MSO from Microsoft (version 2207, means, minima,
maxima, standard deviation, and percentages of the duration and number per
time unit) and presented in diagrams and tables and analysed by week of life
and calves using the situation comparison defined by HOY (2009b) for the pur-
pose of ecological ethology. Using Interact from Mangold, contingency anal-
yses were performed to identify activities before and after the appearance of
one of the selected and recorded behaviour parameters.
Significances are indicated with p < 0.05, p < 0.01 and p < 0.001.
As a rule, the calves are placed in group housing in weeks 3 and 4 on the farm.
After changing housing from individual housing to group housing, calves need
about 5 days to re-establish their circadian rhythm (SCHULDT and DINSE, 2018a,
b; BALTROCK, 2019; SCHULDT and DINSE, 2021a), which is why the first week
after stabling in group housing is considered an adjustment period.
The feeding behaviour of the calves changes, regardless of the milk replacer
allowance and the start of weaning, after week 7 (FRIETEN, 2018; SCHULDT and
DINSE, 2018b, 2019a, 2021a). The resting behaviour develops in terms of the
duration of the lying and standing periods up to week 7 (SCHEURMANN, 1971).
For this reason, the behaviour is differentiated between the age groups from
the adjustment period to day 49, that is, from week 3 to 7, and day 50 until
weaning.
Materials and methods 66
The analysis up to day 49 (n = 9 calves) was carried out in the functional areas
resting, food and water-intake, social behaviour (calf–calf contacts), exploratory
(smelling and licking of objects) and play behaviour, suckling activities, and
‘other activities’. ‘Other activities’ include all activities without food and water-
intake including visits to the dispensing station without MR intake. The param-
eter locomotion behaviour (standing without activity, walking and running,
jumping, galloping without and with elimination) was calculated from the differ-
ence in the number and duration of all coded behaviour in an active phase.
Play behaviour (headbutting, playing with toys), visits without MR intake
(dummy visits) and suckling activities, including cross-sucking and sucking on
dummy teats, was assessed separately. A dummy visit was coded if a calf en-
tered the dispensing station and remained there, without ingesting MR, but in-
dicated a need to drink by playing with the teat. Entering the dispensing station
with no interest in the teat was not evaluated as a dummy visit and was included
in the assessments for locomotion.
The behaviour when weaning was recorded for two rounds with a duration of
55 days (day 50 to 105, weaning on day 106) and was observed and analysed
for five calves.
Because activities between the main phases of a 24-hour day differ signifi-
cantly, the time from 12 midnight to 06:00 a.m. was defined as ‘night’ and the
time from 06:00 a.m. to 12 midnight was defined as ‘during the day’ to simplify
the presentation. To illustrate behaviour over the course of the day, an assess-
ment was carried out in a 2-hourly rhythm, calculating the weekly average of
the duration and frequency of a behavioural category using the average for a
day.
Presentation of the results 67
4 Presentation of the results
4.1 Circadian rhythm
From the start, calves aged three to four weeks show a pronounced circadian
rhythm in group housing (Figure 12). Longer resting phases are recorded over-
night, that is, from 12 midnight to 6:00 a.m. During the day the calves alternated
between shorter resting and activity phases.
The calves are at their most active in the mornings from 6:00 a.m. to about
10:00 a.m. This is followed by resting times that are somewhat shorter between
2:00 and 4:00 p.m., 6:00 and 8:00 p.m. and between 10:00 p.m. and 12 mid-
night compared to between 10:00 a.m. and 2:00 p.m., 4:00 and 6:00 p.m. and
between 8:00 and 10:00 p.m.
Figure 12: Mean duration of activities and rest time during the day from the
week 3 to 7, n = 9 calves
To check if the season or the time of sunrise acted as a zeitgeber for the start
of activities, the mean number of CMR-feed-intakes was compared for 4:00 to
6:00 a.m. (night) and for 6:00 to 8:00 a.m. (start of the day) (Figure 13). In all
seasons CMR-feed was more frequently ingested from 6:00 a.m. onwards than
overnight, and the differences between the means across all rounds for the
start of the day compared to the night hours are significant.
Workflows of the farm personnel affect the activities of the calves. Resting times
during the day are interrupted by cleaning the runs and placing fresh straw
Presentation of the results 68
bedding on the lying area. After this, the animals move more intensely on the
fresh straw, running and jumping about, playing with each other and in the
straw. The resting periods of all calves are further interrupted when individual
animals are treated or trained how to ingest CMR-feed at the dispensing sta-
tion. In the first few days after moving to the group housing, the calves who
have not ingested enough CMR-feed are collected and taken to the dispensing
station, which leads to restlessness in the entire group.
Figure 13: Mean number of CMR-feed-intakes per animal and day from 4:00 to
6:00 a.m. as well as from 6:00 to 8:00 a.m. by seasons from week 3 to 7,
n = 9 calves, significance: 4:00 to 6:00 a.m. vs. 6:00 to 8:00 a.m., p = 0.01
During the weaning phase the circadian rhythm became even more apparent.
At night the animals rested from 12 midnight to about 6:00 a.m., often for more
than one to two hours (h) in a resting period (Figure 14). In the early morning
all calves are active and only rest briefly. From 2:00 to 4:00 p.m. and 10:00 p.m.
to 12 midnight additional periods of intense activity were observed.
Presentation of the results 69
Figure 14: Mean duration of activities and rest time in minutes during weaning
from day 50 to 105, n = 5 calves
4.2 Resting behaviour
4.2.1 Resting behaviour to day 49
All calves sought out lying surfaces with straw bedding to rest. After brief
phases of activity, the calves often lay down in the same place where they had
stood up. This was observed quite frequently overnight. Before lying down, the
animals sniffed the potential lying surface and any calves that were already
lying in this area were then also often intensively licked. The resting place was
often changed for shorter resting phases during the day. No abnormal pro-
cesses were observed when lying down, resting and standing up.
Short periods of lying down (< 1 minute lying duration) were not included in the
calculation of the resting duration, which is why 1.0 minute is often stated as
the minimum for the younger calves (Table A 2).
In group housing, the calves rested for 14.6 to 16.5 h each day from the start.
(Figure 15; Table A 2). On average, there were 17.3 to 23.2 resting periods per
calf with a mean duration of 41.9 ± 41.0 to 49.8 ± 46.4 minutes with a large
spread of up to 6.1 h at night and 4.8 h during the day. (Figure 16; Table A 2).
The mean duration of the resting phases increased from week 5 and the num-
ber of resting periods decreased.
Presentation of the results 70
Overnight, the weekly average for the length of the resting phase ranged from
65.4 to 109.7 minutes or 1.1 to 1.8 h (Table A 2). During the day, mean resting
phases of 33.3 to 42.2 minutes were observed. Large variations were noted in
the length of the resting phases both over the course of the day and between
the calves.
Figure 15: Mean duration of total rest time in hours per day from week 3 to 7,
n = 9 calves
Presentation of the results 71
Figure 16: Mean duration in minutes and number of rest sessions per day from
week 3 to 7, n = 9 calves
Figure 17: Mean duration of rest sessions in minutes over the course of the day
from week 3 to 7, n = 9 calves
Presentation of the results 72
Within a 2-hour interval, the calves rested overnight on average for 57.6 to
89.6 minutes (Figure 17). During the day, shorter resting periods of 26.2 to
44.0 minutes on average were observed.
Figure 18: Percentage of activities during 5 minutes before and after rest from
week 3 to 7, n = 9 calves
CMR = calf milk replacer-feed-intake, BV = visits without CMR-feed-intake
(blind visits), Feed = solid-feed-intake, Water = water-intake, CCC = calf-calf-
contacts, Suck = cross-sucking, suck on calves
In the period 5 minutes before and 5 minutes after the resting periods, 30% and
22% respectively of the calves’ activities up to week 7 involved eating supple-
mentary feed (Figure 18). Resting phases were noted before resting in 34%
and repeat resting phases were observed after resting in 35%. In the 5 minutes
before the resting phases, the calves visited the dispensing station with CMR-
feed-intake in 8% of cases and without CMR-feed-intake in 7%. After the rest-
ing phases, 17% of the activities involved CMR-feed-intake by the calves and
5% were unrewarded visits to the dispensing station.
Cross-sucking processes followed 0.1% of the resting phases, while two calves
sucked once after resting on three days and one calf once on one day.
Presentation of the results 73
4.2.2 Resting behaviour in the weaning phase
During weaning, the calves rested for 14.3 to 16.3 h per day until they were
weaned, except for week 12 which had a mean resting time of 17.6 h. (Figure
19; Table A 3).
Figure 19: Mean duration of total rest time in hours per day during weaning
from day 50 to 105, n = 5 calves
The daily resting period lasted on average 48.3 to 67.4 minutes (Figure 20). On
average, 14.0 to 17.9 periods were noted. The average figure is significantly
lower than that for the age group up to day 49, for which the duration is signifi-
cantly longer.
At night, the calves rested on average for 66.0 to 128.1 minutes. The maximum
continuous resting time observed was from midnight to 7:30 a.m. (451 minutes
or 7.5 h) and the minimum observed was a period of 1.1 minutes. (Table A3).
During the day, the resting time was on average 41.0 to 52.2 minutes with these
values being slightly higher than those for up to week 7.
Presentation of the results 74
Figure 20: Mean duration of rest sessions in minutes and number of rest ses-
sions per day during weaning from day 50 to 105, n = 5 calves
In the period 5 minutes before and after resting, eating supplementary feed
dominated, making up 43% and 34% of all activities respectively (Figure 21).
This was followed by another resting phase (33% and 31% respectively). CMR-
feed-intake and water-intake, blind visits and contacts were all at about the
same level of 2% to 10% before resting and 5% to 10% after resting.
Before resting, three calves cross-sucked, while one calf was observed cross-
sucking after resting on day 60 but in each case this occurred only once on one
day.
Presentation of the results 75
Figure 21: Percentage of activities during 5 minutes before and after rest during
weaning from day 50 to 105, n = 5 calves
CMR = calf milk replacer-feed-intake, BV = visits without CMR-feed-intake
(blind visits), Feed = solid-feed-intake, Water = water-intake, CCC = calf-calf-
contacts, Suck = cross-sucking, suck on calves
4.3 Food and water-intake behaviour
4.3.1 Food and water-intake behaviour up to day 49
4.3.1.1 Drinking behaviour
An CMR-feed meal lasts on average 4.1 to 4.9 minutes with a rising trend (Fig-
ure 22; Table A 4). The number of meals per day increases between week 3
and 4 from 4.3 to 5.3 and thereafter remains steady at about six meals per
animal per day.
The most intensive drink intake occurs from week 3 to 7 between 6:00 and 8:00
a.m. and from 2:00 to 8:00 p.m. with 44.7% of CMR-feed-intakes occurring in
this time (Figure 23). From 12 midnight to 6:00 a.m., 14.1% of the CMR-feed-
intakes were recorded.
Presentation of the results 76
Figure 22: Mean duration in minutes and mean number of CMR-feed-intakes
per animal and day from week 3 to 7, n = 9 calves
Figure 23: Percentage of CMR-feed-intakes over the course of the day from
week 3 to 7, n = 9 calves
CMR = calf milk replacer
Presentation of the results 77
A comparison of the duration of a CMR-feed meal overnight (12 midnight to
6:00 a.m.) vs. during the day (6:00 a.m. to 12 midnight) showed differences
only in the maximum values (Table A 4). The minimum values overnight in
weeks 3 and 4 must be disregarded because of the small number of calves and
datasets. In week 5, the longest duration of an CMR-feed-intake of
14.6 minutes is reported. Overall, the spread is high between the calves and
over the course of 24 hours (SD = 1.8 to 2.6 minutes).
The calves rested before 53% of CMR-feed-intakes and 21% of the supple-
mentary feed intakes (Figure 24). Water-intakes, playing and contacts with
other calves occurred at rates of 9%, 10% and 3%. Only 3% and 1% of the
calves respectively visited the dispensing station with or without CMR-feed-in-
take. CMR-feed-intake was followed within 5 minutes by eating supplementary
feed or resting each with 28% of all activities. Contact between the calves, play-
ing or water-intake was observed in 11%, 14% and 10% respectively. A total of
6% of the CMR-feed-intakes were followed by another, albeit unrewarded, visit
to the dispensing station. One calf cross-sucked after the CMR-feed-intake
once in week 7.
Presentation of the results 78
Figure 24: Percentage of activities during 5 minutes before and after CMR-
feed-intake from week 3 to 7, n = 9 calves
CMR = calf milk replacer-feed-intake, BV = visits without CMR-feed-intake
(blind visits), Feed = solid-feed-intake, Water = water-intake, CCC = calf-calf-
contacts, Suck = cross-sucking, suck on calves
4.3.1.2 Behaviour associated with supplementary feed intake
Up to week 6 the hay rack and trough were only briefly visited. Up to the end of
week 7, the calves ate supplementary feed on average 10.7 to 18.8 times in 24
h over 1.5 to 3.1 minutes per meal with a slight increase as the calves age
(Figures 25 (a), (b); Table A 5). The mean duration of meals for the hay and
dry TMR intakes is approximately the same but the number of dry TMR intakes
is higher.
The supplementary feed intake shows a clear difference in the number of meals
over 24 h, making up 8.5% of all meals in the 6 h overnight (12 midnight to 6:00
a.m., Figure 26) compared to 24.0% of meals from 6:00 to 10:00 a.m. and
43.0% from 2:00 to 10:00 p.m.
Presentation of the results 79
(a)
(b)
Figures 25: (a), (b): Mean duration in minutes and number of feed-intakes per
animal and day (a) total solid feed (b) hay and dry TMR (total mixed ration) from
week 3 to 7, n = 9 calves
Presentation of the results 80
Figure 26: Percentage of solid-feed-intakes over the course of the day from
week 3 to 7, n = 9 calves
Figure 27: Percentage of solid-feed-intakes per animal by duration of intake
from week 3 to 7, n = 9 calves
Presentation of the results 81
The increase in the duration of a supplementary feed-intake is seen particularly
for the maximum values for a week (from week 3 to 7: 6.4 to 26.2 minutes) and
the increase of the percentage of meals lasting more than 3 minutes, while at
the same time the spread of the data increased (Figure 27; Table A 5). How-
ever, the percentage of longer mealtimes > 10 minutes in week 7 was only 3%
with 83% of all supplementary feed-intakes lasting less than 5 minutes.
Resting periods were most frequently observed in the 5 minutes before visiting
the trough or rack (27%) and the rate after eating supplementary feed was even
higher, making up 35% of all activities (Figure 28). Before eating supplementary
feed, the calves played about as often as they ate supplementary feed again.
Visits to the dispensing station as well as drinking water and contacts between
the calves occur a little less often before eating. Cross-sucking was not ob-
served in connection with intake of supplementary feed up to week 7.
Figure 28: Percentage of activities during 5 minutes before and after solid-feed-
intake from week 3 to 7, n = 9 calves
CMR = calf milk replacer-feed-intake, BV = visits without CMR-feed-intake
(blind visits), Feed = solid-feed-intake, Water = water-intake, CCC = calf-calf-
contacts, Suck = cross-sucking, suck on calves
Presentation of the results 82
4.3.1.3 Behaviour associated with water-intake
Young calves use the automatic drinker for water-intake from the start of the
group housing. In week 3, more frequent water drinking was noted for the calf
observed here than for all 13 calves in the following weeks (Figure 29; Table
A 6). From week 4 to the end of week 7, the calves drank water on average 2.4
to 5.6 times over about 1 minute with the mean number and duration increasing
with age.
Figure 29: Mean duration in minutes and number of water-intakes per animal
and day from week 3 to 7, n = 9 calves
At night the calves only rarely drank water with only 8.4% of all water-intakes
observed between 12 midnight and 6:00 a.m. (Figure 30). The mean duration
of a water-intake lasted 0.8 to 1.2 minutes. Maximum values of 8.6 and 8.4
minutes were noted for two calves in weeks 5 and 7 between 12 midnight and
1:00 a.m. (Table A 6).
Presentation of the results 83
Figure 30: Percentage of water-intakes during the day from week 3 to 7, n = 9
calves
4.3.1.4 Visits to the dispensing station without CMR-feed-intake (blind visits)
A visit without CMR-feed-intake (blind visit) was coded if a calf stayed in the
dispensing station without ingesting CMR-feed.
In week 3, for one calf studied during the adjustment period, each day 3.5 visits
without CMR-feed-intake of an average of 0.9 ± 0.9 minutes were noted (Figure
31; Table A 7). From week 4 to 7 the daily average of visits to the dispensing
station without CMR-feed-intake was 2.1 to 3.2 and a blind visit lasted on aver-
age 0.7 to 1.0 minutes. As a maximum, one blind visit in week 7 lasted more
than 6.5 minutes.
Presentation of the results 84
Figure 31: Mean duration in minutes and number of blind visits per animal and
day from week 3 to 7, n = 9 calves
Figure 32: Percentage of blind visits by mean duration in minutes from week
3 to 7, n = 9 calves
Presentation of the results 85
On average, from week 3 to 7 the calves stayed for less than 1 minute in the
dispensing station without CMR-feed-intake in 70% of cases (Figure 32). About
one-quarter of the blind visits lasted 1 to 2 minutes and only 8% lasted more
than 2 minutes.
From 12 midnight to 6:00 a.m., 9.4% of all blind visits were recorded (Figure
33). The highest number of blind visits of 15.1% in 2 hours is seen between
4:00 and 6:00 p.m.
In the 5 minutes before and after a blind visit, none of the assessed calves
sucked up to week 7 (Figure 34). Before 25% of the unrewarded visits to the
dispensing station, the calves rested, before 12% to 17% they ate supplemen-
tary feed or drank water or played, and the same percentage made repeat blind
visits within 5 minutes. CMR-feed-intakes and social contacts occurred a little
less frequently (8% and 5% respectively). After 26% and 24% of the blind visits,
the calves ingested supplementary feed or water respectively, and after 15%
and 13% they rested or played respectively. CMR-feed-intakes and contacts
between the calves do not play a role after unrewarded visits to the dispensing
station.
Figure 33: Percentage of blind visits over the course of the day from week 3 to
7, n = 9 calves
Presentation of the results 86
Figure 34: Percentage of activities during 5 minutes before and after blind visits
from week 3 to 7, n = 9 calves
CMR = calf milk replacer-feed-intake, BV = visits without CMR-feed-intake
(blind visits), Feed = solid-feed-intake, Water = water-intake, CCC = calf-calf-
contacts, Suck = cross-sucking, suck on calves
4.3.2 Food and water-intake behaviour during the weaning phase
4.3.2.1 Drinking behaviour
One CMR-feed-intake lasted on average 6.3 minutes in weeks 9 and 10 and
4.6 to 4.8 minutes in weeks 8, 11 and 15 (Figure 35; Table A 8). Between week
12 and 14 the mean duration of an CMR-feed-intake decreased to 3.5 minutes
and then increased again in week 15. The number of daily meals in the weaning
phase depends on the number of allowances and decreases in week 14 to two
per animal and day. A mean CMR-feed-intake lasts 4.9 minutes over the entire
weaning phase and is thus significantly longer than with the maximum allow-
ance up to day 49 (4.6 minutes on average, p < 0.05).
Presentation of the results 87
Figure 35: Mean duration in minutes and number of CMR-feed-intakes per an-
imal and day during weaning from day 50 to 105, n = 5 calves
The circadian rhythm described in Section 4.1 is apparent in the weaning phase
in the frequency of CMR-feed meals (Figure 36). From 12 midnight to 6:00 a.m.,
11.7% of all meals were recorded. The restrictions in the daily milk replacer
allowances are apparent in the percentage of CMR-feed meals that were rec-
orded with a total of 50.3% in the mornings from 6:00 to 8:00 a.m. and in the
evenings from 6:00 to 8:00 p.m.
Presentation of the results 88
Figure 36: Percentage of CMR-feed-intake over the course of the day during
weaning from day 50 to 105, n = 5 calves
Before a CMR-feed-intake, the calves most frequently rested or they ate sup-
plementary feed (38% and 28% respectively, Figure 37). After drinking, visits
to the trough or rack dominated (49%), and resting periods followed only in 8%
of cases. For all other activities, there are only negligible differences in the per-
centages before and after drinking.
Two calves cross-sucked once (week 8 to 14) before a CMR-feed-intake and
on one or four days once each after that.
Presentation of the results 89
Figure 37: Percentage of activities during 5 minutes before and after CMR-
feed-intake during weaning from day 50 to 105, n = 5 calves
CMR = calf milk replacer-feed-intake, BV = visits without CMR-feed-intake
(blind visits), Feed = solid-feed-intake, Water = water-intake, CCC = calf-calf-
contacts, Suck = cross-sucking, suck on calves
4.3.2.2 Behaviour associated with supplementary feed-intake
With the start of weaning, the dry TMR is gradually replaced by a TMR in the
supplementary feeding. It is placed in front of the front opening of the pen en-
closure. The supplementary feed thus includes hay and TMR with a decreasing
proportion of dry TMR.
The mean daily duration and number of supplementary feeds increases during
the weaning phase from 1.1 h and 16 meals to 2.7 h and > 20 meals per animal
and day (Figure 38; Table A 9). In week 15 a reduced duration and number of
the supplementary feeds was recorded compared to the average for weeks 8
and 14.
Presentation of the results 90
Figure 38: Mean duration of total solid-feed-intake in hours per day and number
of solid-feed-intakes per animal and day during weaning from day 50 to 105,
n = 5 calves
Figure 39: Percentage of solid-feed-intake over the course of the day during
weaning from day 50 to 105, n = 5 calves
Presentation of the results 91
The circadian rhythm of the calves is also apparent in the supplementary feed-
intakes (Figure 39). Overnight, only a few meals (7.4% of the supplementary
feeds) were recorded. During the day, the meals followed the rhythm with on
average approx. 10% in 2 h. A mean supplementary feed-intake lasted signifi-
cantly longer in the weaning phase than in the weeks with the maximum allow-
ance (2.5 minutes vs. 5.0 minutes, p < 0.001, Tables A 5, A 8).
Figure 40: Percentage of solid-feed-intake by duration of intake during weaning
from day 50 to 105, n = 5 calves
With increasing age, the proportion of shorter meals of less than three minutes
fell in favour of longer supplementary feed-intakes (Figure 40, Table A 10). The
percentage of meals of 10 minutes and more eating time in week 10 was still
7% and it was only in week 12 that the percentage increases to 11%, achieving
the highest values of 23% and 21% in weeks 13 and 14 respectively. In week
15 the percentage is 4%. The high percentage of short supplementary feed-
intakes of > 80% corresponds to a decrease in the total duration in the last
week.
From week 8 to 14 the calves consistently eat a hay meal on average over 4.4
to 6.1 minutes with a maximum of 39.7 minutes (Figure 41 (a)). In week 15 an
average of 3.3 minutes was noted. With the uptake of TMR, an increase in the
duration of a meal was identified during weaning (Figure 41
(b)). The calves
increased their mean TMR intake from 3.4 minutes per meal in week 8 to 8.2
Presentation of the results 92
and 7.0 minutes in weeks 13 and 14 respectively with a maximum of 59.7
minutes. In week 15 the observed TMR intake decreased to 2.6 minutes on
average.
In the 5 minutes before and after a supplementary feed, all activities occurred
with about the same frequency (Figure 42). Blind visits were recorded before
10% of the supplementary feeds while the rate recorded after visiting the rack
or trough was 9%. The calves often played before and after eating supplemen-
tary feed. Several sequential supplementary feed-intakes took place in 26% of
cases, with the calves often switching between rack and trough, but this is not
shown separately here. Of all the activities, rest made up 21% of all activities
before and 25% after a supplementary feed-intake.
Cross-sucking was observed twice before and once after a visit to the rack or
trough in each case by one calf.
(a)
Presentation of the results 93
(b)
Figures 41 (a), (b): Mean and maximum duration of intake of (a) hay and (b)
TMR (total mixed ration) during weaning from day 50 to 105, n = 5 calves
Figure 42: Percentage of activities during 5 minutes before solid-feed-intake
during weaning from day 50 to 105, n = 5 calves
CMR = calf milk replacer-feed-intake, BV = visits without CMR-feed-intake
(blind visits), Feed = solid-feed-intake, Water = water-intake, CCC = calf-calf-
contacts, Suck = cross-sucking, suck on calves
Presentation of the results 94
4.3.2.3 Behaviour associated with water-intake
The mean number of water-intakes at the automatic drinker increased during
the weaning phase from three to eight to ten times per day (Figure 43). The
mean duration of a water-intake remained relatively steady at 0.7 to 1.5
minutes. The total duration over 24 h thus increased with increasing age from
2.4 to 11.0 minutes per animal (Table A 11). There is a clear circadian rhythm
apparent in the water-intake of the calves (Figure 44). Most visits to the auto-
matic drinker occurred during the main times for food intake of 6:00 to 8:00
a.m., 2:00 to 4:00 p.m., and 6:00 to 8:00 p.m. The highest percentage in 2 h of
19.1% was recorded from 6:00 to 8:00 p.m.
Figure 43: Mean duration in minutes and number of water-intakes per animal
and day during weaning from day 50 to 105, n = 5 calves
Presentation of the results 95
Figure 44: Percentage of water-intake over the course of the day during wean-
ing from day 50 to 105, n = 5 calves
4.3.2.4 Visits to the dispensing station without CMR-feed-intake (blind visits)
With the decrease in the milk replacer allowance, the number of visits to the
dispensing station without an allowance increased during the weaning phase
but the duration of a blind visit remained about one minute on average (Figure
45; Table A 12). The number of unsuccessful attempts to ingest CMR-feed in-
creased gradually from an average of 2.5 blind visits per animal and day in
week 8 to 11.8 in week 13. The dummy visits then declined to an average of
5.3 during the day in week 15.
More than half were short visits of less than one minute in duration (54%). Only
14% of the blind visits lasted two minutes or more (Figure 46).
Like the food intake, the blind visits correspond to the circadian rhythm of the
calves (Figure 47). Frequent unsuccessful visits to the dispensing station were
recorded between 6:00 and 8:00 a.m., from 2:00 to 6:00 p.m. (17.0% and
11.0%–11.8% of the mean daily dummy visits) and between 6:00 and 8:00 p.m.
(27.2%), while overnight and over the rest of the day it was considerably less
(1.3%–6.7% in 2 h).
One-third of the blind visits took place before and after eating supplementary
feed up to day 105 (Figure 48). Before blind visits, water-intake (19%) and
Presentation of the results 96
resting phases (16%) were commonly recorded. There were also several blind
visits one after the other within 5 minutes (24%). CMR-feed-intakes, social con-
tacts and cross-sucking were each observed before a maximum of 2% of the
dummy visits.
In terms of frequency, the activities before and after blind visits are similar, ex-
cept for resting periods, which make up 16% before and 5% after visiting with-
out CMR-feed-intake (Figure 48). Supplementary feed and water-intake as well
repeat blind visits dominate before and after blind visits.
Two calves cross-sucked in the 5 minutes before a blind visit (week 8, 14) and
two calves after a blind visit (week 10, 14). Another calf from the 1st round
cross-sucked twice on one day within 5 minutes and twice visited the dispens-
ing station with no intake in this period. On the same day it cross-sucked again
after a blind visit.
Figure 45: Mean duration in minutes and number of blind visits per animal and
day during weaning from day 50 to 105, n = 5 calves
Presentation of the results 97
Figure 46: Percentage of blind visits by mean duration in minutes during wean-
ing from day 50 to 105, n = 5 calves
Figure 47: Percentage of blind visits over the course of the day during weaning
from day 50 to 105, n = 5 calves
Presentation of the results 98
Figure 48: Percentage of activities during 5 minutes before and after blind visits
during weaning from day 50 to 105, n = 5 calves
CMR = calf milk replacer-feed-intake, BV = visits without CMR-feed-intake
(blind visits), Feed = solid-feed-intake, Water = water-intake, CCC = calf-calf-
contacts, Suck = cross-sucking, suck on calves
4.4 Social behaviour
4.4.1 Social behaviour up to day 49
Social relationships between the calves are described using the parameter
‘calf–calf contacts’ and include contacts in which the calves sniff and lick one
another but not cross-suck. The daily average duration of each of these con-
tacts is 1.3 to 2.0 minutes (Figure 49; Table A 13). From week 3 onwards, there
are more frequent contacts and from week 4 to the end of week 7 the daily
average is 4.6 to 5.6.
Overnight (12 midnight to 6:00 a.m.) there are only few social contacts and only
7.0% of all contact occur in these six hours (Figure 50). In the mornings, the
calves lick one another most frequently from 6:00 a.m. to 12 midday (39.3% of
all contacts). Maximum values for the contacts were recorded from week 3 to
7 with 16.4 and 15.7 minutes (Table A 13).
Presentation of the results 99
Figure 49: Mean duration in minutes and number of calf–calf contacts from
week 3 to 7, n = 9 calves
Figure 50: Percentage of calf–calf contacts over the course of the day from
week 3 to 7, n = 9 calves
Presentation of the results 100
4.4.2 Social behaviour in the weaning phase
Contacts between the calves last on average 1.1 to 2.1 minutes per event (Fig-
ure 51; Table A 14). Until week 10, frequent contacts between the animals were
observed but the number decreased with increasing age.
Figure 51: Mean duration in minutes and number of calf–calf contacts per ani-
mal and day during weaning from day 50 to 105, n = 5 calves
4.5 Other activities and play behaviour
4.5.1 Other activities
Until the end of week 6, the calves spend 6.8 to 8.4 h a day on other activities
that are not related to eating food (activities without CMR-feed-intakes, supple-
mentary feed and water, blind visits), which corresponds to 85% to 91% of the
active time (Figure 52). In week 7, the percentage decreases to 80% (6.0 h).
Other activities were observed predominantly during the day (Figure 53). In the
second half of the night, when the calves rest deeply for long periods, the calves
only ingest water and supplementary feed briefly during the awake phases. The
calves frequently urinate and defecate and run briefly around the pen without
any apparent purpose, with only 11.2% of the other activities in a complete 24-
h day taking place at night. Shortly before lying down, the calves seek contact
with the other animals. Peaks in the other activities were observed between
6:00 a.m. and 12 midday and between 4:00 and 10:00 p.m.
Presentation of the results 101
Figure 52: Mean duration of active time and other activities (locomotion, explo-
ration, play) in hours per animal and day and other activities as a percentage
of active time from week 3 to 7, n = 9 calves
Figure 53: Percentage of other activities over the course of the day from week
3 to 7, n = 9 calves
Presentation of the results 102
The locomotion behaviour, which is included in the other activities, decreases
from week 3 to 7 from an average of 8.0 to 5.7 h per day (Figure 54; Table A
15).
Figure 54: Mean duration of active time and locomotion behaviour per animal
and day from week 3 to 7, n = 9 calves
Licking objects, one element of exploratory behaviour, was observed for 6.5 to
15.5 minutes on average per calf and day (Figure 55; Table A 15) with a large
range in the duration of the activity recorded from 1.6 to 39.1 minutes. The
smaller number of up to five calves per week and a maximum seven datasets
up to week 6 that can be assessed must be noted here.
Presentation of the results 103
Figure 55: Licking objects: mean duration in minutes per day, maximum num-
ber per animal and day and mean duration in minutes from week 3 to 7, n = 9
calves
When assessing behaviour during the weaning phase, locomotion and explor-
atory behaviour were not considered separately because there were insufficient
data available. Other activities include the active time without food and water-
intake.
From week 8 to 10 the percentage of other activities relative to the overall active
time stayed at 79% to 83% (Figure 56). Starting at week 11, the percentage of
other activities decreased to 71% and remained between 57% and 70% from
week 12 to 15. The total duration of all activities from week 9 to 15 is between
7.9 and 9.7 h, except for week 12 when the assessed calves were active on
average for only 6.4 h per day.
At night only few activities took place (Figure 57). The calves are briefly awake,
drink CMR-feed and/or supplementary feed or urinate and defecate. The per-
centage overnight makes up only 11.9% of all other activities during the day.
During the day, the peak times are recorded from 6:00 a.m. to 12 midday, 2:00
to 4:00 p.m. and 6:00 to 8:00 p.m.
Presentation of the results 104
Figure 56: Mean duration of active time and other activities (locomotion, explo-
ration, play) in hours per animal and day and other activities as a percentage
of active time during weaning from day 50 to 105, n = 5 calves
Figure 57: Percentage of other activities over the course of the day during
weaning from day 50 to 105, n = 5 calves
Presentation of the results 105
4.5.2 Play behaviour
Playing with toys – ball, brush and chains – was described as play behaviour
along with headbutting. Headbutting was observed in three calves as early as
week 4. In most cases only two animals butted their heads together with alter-
nating pairs observed less often during a game up to week 7. The activities are
instigated by all calves with even smaller animals encouraging larger members
of the group.
In the daily average, the calves play for an average of 7.6 to 12.6 minutes,
ranging from 0.2 to 36.1 minutes per calf and day (Figure 58; Table A 16). A
game lasted on average 1.1 to 1.4 minutes with one game recorded that lasted
9.1 minutes, the maximum.
Figure 58: Mean total duration in minutes and number of games per animal and
day and mean duration of games per animal and day from week 3 to 7,
n = 9 calves
To illustrate play over the course of the day, in Figure 59 the mean percentage
of games in 2 hours is plotted against the number of games in a day. Calves
play at all times of the day. However, the most common play activities occur
between 4:00 to 10:00 p.m. (42.8% of all play events). In this time the calves
run, jump and gallop most intensively around the pen, play fight or use the toys.
At night between 12 midnight and 6:00 a.m. the calves hardly play.
Presentation of the results 106
Figure 59: Percentage of games over the course of the day from week 3 to 7,
n = 9 calves
The play behaviour of the calves was studied in terms of the number and dura-
tion of the activities (Table A 17). The maximum value of 9.1 minutes cited
above was recorded for playing with the chains (week 6) while playing with the
ball for 7.9 minutes was observed (week 7).
From week 4 to 7 the calves preferred to play with a ball hanging from a chain
(40.0% of the play time, Figure 61; 4.0–5.2 times a day over 5.2–7.2 minutes
Figures 60 (a), (b); 52.9% of the frequency per calf, Table A 18). Headbutting
and playing with the brush made up 20.7% and 18.5% respectively of the total
play time. The calves played with the chains for 20.8% of the total play period
over 24 h.
The animals used the brush and chains about equally for 18.5% and 18.3% of
the play events. Headbutting makes up 10.3% of the play activities of the indi-
vidual calves (Table A 18).
Presentation of the results 107
(a)
(b)
Figures 60 (a), (b): Mean (a) duration and (b) number of games with ball, brush
and chains as well as headbuttings per animal and day from week 3 to 7,
n = 9 calves
Presentation of the results 108
Figure 61: Percentage of playtime with ball, brush or chains as well as
headbutting in the total playtime per animal and day from week 3 to 7,
n = 9 calves
During the weaning phase, calves play on average for up to 20.2 minutes a day
with a toy or headbutting each other (Figure 62). A game lasts on average 1.3
to 2.0 minutes with a maximum of 18.5 minutes (Table A 19). The mean number
per animal and day is 5.1 to 10.0.
Over the course of the day, the calves play according to the circadian rhythm
(Figure 63). They were observed playing particularly in the mornings, between
2:00 and 4:00 p.m., and from 6:00 to 8:00 p.m. Playing was barely recorded
overnight.
The calves played most intensively with the ball for an average of 3.9 to 15.1
minutes per day (Figure 64). Mean daily play periods of 1.0 to 7.2 minutes were
observed with the other play equipment.
In the weaning phase playing with the ball dominated, making up 50.2% of all
games (Figure 65). There was barely any difference in the calves playing with
the brush, chains (18.5% and 18.2% respectively) or headbutting (13.0%).
Presentation of the results 109
Figure 62: Mean total duration in minutes per animal and day, mean duration
in minutes and number of games per animal and day during weaning from day
50 to 105, n = 5 calves
Figure 63: Percentage games over the course of the day during weaning from
day 50 to 105, n = 5 calves
Presentation of the results 110
Figure 64: Mean duration of games with ball, brush and chains as well as
headbutting per animal and day during weaning from day 50 to 105,
n = 5 calves
Figure 65: Percentage of playtime with ball, brush or chains as well as
headbutting in the total playtime per day during weaning from day 50 to 105,
n = 5 calves
Presentation of the results 111
4.6 Sucking activities without visiting the dispensing station
4.6.1 Cross-sucking on another calf
Up to the end of week 7, cross-sucking was only observed for one calf (Figure
66; Tables A 20, A 21). In week 6 this calf cross-sucked once for 9.7 minutes
and in week 7 once on each of three days for an average of 3.1 minutes. Over-
night there were no sucking activities among the observed calves in the five
rounds up to day 49. One calf attempted after its CMR-feed meal to cross-suck
the calf that used the dispensing station after it. It was fended off forcefully twice
and made no further attempts.
Of the five calves that were observed during weaning, three cross-sucked on
another calf (Figure 67; Tables A 20, A 21). One calf started to cross-suck in
week 14, with eight events over 0.6 to 4.5 minutes observed for this calf in week
15, in each case one cross-sucking event per week at night. Another animal
cross-sucked in week 11 once over 1.4 minutes and in week 13 once a day
over 0.9, 1.4 and 1.6 minutes.
Presentation of the results 112
Figure 66: Mean duration in minutes, maximum number of sucking sessions
per day and number of days with sucking activities from week 3 to 7,
n = 9 calves, 1 sucking calf
Figure 67: Mean duration of a sucking session in minutes, maximum number
of sucking sessions per day and number of days with sucking activities during
weaning from day 50 to 105, n = 5 calves
Presentation of the results 113
The activities before and after cross-sucking are assessed over the entire drink-
ing period. Before cross-sucking, 62% of activities were related to the dispens-
ing station, half of which were without a milk replacer intake, while after cross-
sucking only 41% of activities were related to the dispensing station (Figure
68). Cross-sucking was followed by drinking water in 25% of cases while the
calves did not drink water before cross-sucking. No animal cross-sucked sev-
eral times consecutively. Eating supplementary feed, playing and contacts oc-
curred before cross-sucking in 8% to 15%, afterwards no playing was observed
while visits to the trough or rack occurred in 17% of cases and contacts be-
tween the calves in 8%. Within one minute of cross-sucking, no calf rested,
while after cross-sucking activity resting made up 8% of the subsequent ob-
served activities.
Figure 68: Percentage of activities during 1 minute before and after sucking
activities, from week 6 to 15, n = 5 calves
CMR = calf milk replacer-feed-intake, BV = visits without CMR-feed-intake
(blind visits), Feed = solid-feed-intake, Water = water-intake, CCC = calf-calf-
contacts, Suck = cross-sucking, suck on calves
Presentation of the results 114
4.6.2 Sucking on dummy teats
An empty calf feeder with ten teats and two boards each with four blank plugs
were installed in the calves’ pen for use as dummy teats. The calf feeder is only
in the pen during the day but the dummy boards are kept there over 24 h. One
board was placed next to the dispensing station and the second was placed
next to the gate to the open area. Because of the small number of datasets, the
use of the dummy teats is assessed across the entire group housing.
The ten animals for which there are assessable data sucked on average for 0.6
to 1.5 minutes on a dummy teat for a total of 183 times on 56 days (Figure 69,
Table 6). On one day a maximum of 12 events on the calf feeder was regis-
tered. The high mean of 5.8 minutes in week 12 is due to two sucking events
by one animal over 3.5 and 8.1 minutes.
On average a sucking event lasted 1.3 minutes with a maximum of
12.4 minutes at the calf feeder, 2.4 minutes at the dummy board next to the
dispensing station and 2.9 minutes at the board near the gate (Tables A 22 to
A 24). One animal from the 1st round, two animals from the 2nd round and
three from the 3rd round sucked on the teats of the empty calf feeder a total of
154 times on 35 days for an average of 1.3 ± 1.4 minutes.
One calf in the 4th round and two in the 5th round sucked on the dummy teat
next to the dispensing station and two calves from the 4th and one from the 5th
round sucked on the dummy board that was mounted next to the gate to the
lying area. In total, only 29 sucking events were observed, meaning that the
data can only be assessed to a limited degree. The calves sucked on the
dummy teats next to the dispensing station 16 times on 12 days for 0.7 ±
0.5 minutes on average, of which 8.3% was overnight (Table A 23). A total of
13 sucking events on the dummy teats next to the outer gate were observed
on 11 days with an average duration of 1.1 ± 0.9 minutes, of which 19.0% was
overnight (Table A 24).
Presentation of the results 115
Figure 69: Mean duration of sucking activities at the calf feeder, at the dummy
teats next to the dispensing station (du sta) and at the gate (du gate), week 4
to 14, n = 10 calves
Presentation of the results 116
Table 6: Statistical parameters of sucking on dummy teats, proportion of sucking events at night and proportion of visits
to the dispensing station with and without CMR-feed-intake in all activities up to 5 minutes before sucking events from
week 6 to 15
C / R / du Sucking sessions
Number During
n d max
/
d
Mean duration perc.
at night
perc.
CMR
/
BV
before sucking
Average min max s
3 / 1 / cf 40 10 10 1.4 0.04 8.1 1.5 - 9 / 21 %
4 / 2 / cf 31 5 12 1.3 0.1 6.6 1.3 - 20 / 8 %
5 / 2 / cf 31 7 8 1.4 0.2 3.5 0.8 - 35 / 4 %
6 / 3 / cf 11 3 8 1.3 0.6 2.9 0.8 - 0 / 11 %
8 / 3 / cf 20 5 10 1.5 0.03 12.4 2.7 - 28 / 28 %
9 / 3 / cf 21 5 10 1.0 0.3 3.7 0.7 - 18 / 18 %
10 / 4 / sd 2 1 2 0.6 0.6 0.7 0.1 46.0 % 0 / 0 %
11 / 4 / sd 6 4 2 1.0 0.4 2.6 0.8 10.7 % 22 / 11 %
12 / 5 / sd 5 4 4 0.9 0.3 2.4 0.9 0 % 0 / 25 %
13 / 5 / sd 16 12 4 1.1 0.2 2.9 0.8 37.4 % 13 / 6 %
Total 183 56 12 1.3 0.03 12.4 1.4 27.7
% 17 / 15
%
C = calf, R = round, du = dummy, cf = calf feeder, sd = suction dummy, n = sucking sessions, d = days, max = maximum, s = standard deviation, perc. = percentage,
at night = 00:00 to 06:00 a.m., CMR / BV = CMR-feed-intake/ blind visits, before sucking = up to 5 minutes before sucking
Presentation of the results 117
Table 7: Number of sucking sessions and sucking activities on the dummy teats by calves, n = 13 calves
Calf / R
Number
Sucking sessions Sucking activities on the dummies
Calf feeder sd next station sd gate
1 / 1 9 - - -
2 / 1 0 - - -
3 / 1 0 40 - -
4 / 2 0 31 - -
5 / 2 4 31 - -
6 / 3 0 11 - -
7 / 3 0 - - -
8 / 3 0 20 - -
9 / 3 4 21 - -
10 / 4 0 - 0 2
11 / 4 0 - 3 3
12 / 5 0 - 5 0
13 / 5 0 - 8 8
R = round, sd = suction dummy, station = dispensing station, gate = at the gate, - = no dummy in the group box on observation days
Presentation of the results 118
Four calves sucked on the calf feeder and four other calves sucked on the dummy
teats but never cross-sucked (Table 7). Cross-sucking was recorded by two
calves 4 times each and these calves also sucked on the empty calf feeder. For
three cross-sucking calves (sucking in weeks 14 and 15) and two calves that did
not cross-suck, the behaviour in the weeks when there was a calf feeder in the
group pen was not assessed.
In 36% of the assessed activities, up to 5 minutes before sucking on a dummy
teat the calves visited the dispensing station, almost half of which attempts were
unrewarded (Figure 70). 30% of the activities involved play, 23% involved eating
supplementary feed with contacts between the calves occurring before 5% of the
sucking activities. After the calves sucked on a dummy teat, they moved to eat at
the trough or rack (38%), rested, visited the dispensing station or drank water
(10% to 16%). Contacts between the calves were recorded after 7% and play
after only 2% of the sucking events. The dummy teats were not used before or
after cross-sucking.
Figure 70: Percentage of activities in the 5 minutes before and after sucking ac-
tivities on a dummy teat, from week 4 to 15, n = 10 calves
CMR = calf milk replacer-feed-intake, BV = visits without CMR-feed-intake (blind
visits), Feed = solid-feed-intake, Water = water-intake, CCC = calf-calf-contacts,
Suck = cross-sucking, suck on calves
Discussion 119
5 Discussion
5.1 Methodology used to observe and evaluate the behaviour
The most beneficial tool for improving animal welfare is animal observation
(HARMS, 2021). Behavioural observations are, however, elaborate, which is why
the behaviour of calves is observed only on selected days or over a few hours
during the day because of the high amount of time required (WEBSTER and
SAVILLE, 1982; KEIL et al., 2002; BRUMMER, 2004; EGLE, 2005; FISCHER, 2006; UDE
and GEORG, 2006; KÜRN, 2017; UGWU, 2020).
ALIMIRZAEI et al. (2020) compared the behaviour of intensively (IF) and conven-
tionally (CF) fed calves during weaning. One week before to two weeks after
weaning, the CF calves spent more time eating and drinking and rested longer in
the 60 minutes after drinking the morning milk or supplementary feed than did the
IF calves, but the lying behaviour of both groups of weaning calves was the same.
The authors conclude that intensive feeding before weaning would be detrimental
to well-being during weaning. We must disagree with this, however, because the
calves’ behaviour was only observed twice a week for one hour, which would not
permit any reliable insights into the behaviour of the calves over 24 hours (h). The
maximum allowance for the IF calves of 1.02 kg dry matter per calf and day also
does
not
correspond
to ‘intensive’
feeding,
even
if
weaning was only
started
on
day
51.
Random observations are not likely to allow meaningful conclusions to be drawn
about behaviour across the entire rearing period or in the day–night rhythm. Sev-
eral animals must be observed because extrapolating from individuals to the an-
imals as a whole is only possible to a limited degree. A few hours’ observation
will also not be indicative of the behaviour of the animals over the entire day and
observations during the day do not permit conclusions to be drawn about activi-
ties at night. Identifying qualitative and quantitative changes over the course of a
day requires instead uninterrupted 24-hour observation. (PORZIG, 1964; WEBSTER
and SAVILLE, 1982)
For our studies on the behaviour of calves in off-cow, intensive rearing, we there-
fore analysed data from up to nine calves per week with daily milk replacer allow-
ances of 12 L CMR-feed up to day 49 over 24 hours on two to seven days in each
case throughout the entire group housing phase, that is, from week 3 to the end
of week 15. Even though the behaviour of one animal only permits limited con-
clusions to be drawn about the herd behaviour (PORZIG, 1964), these studies must
remain limited to observations of 13 calves on selected days over 5 rounds as
Discussion 120
focus animals as defined by HOY (2009a) because of the enormous amount of
time required.
This procedure is also used by UGWU (2020) who observed and investigated the
behaviour of four calves, each in different husbandry systems, over 24 h directly
and using video recordings.
KILGOUR (1978) and TSCHANZ (1984) urged that ethograms be developed for ag-
ricultural livestock to determine if the design of housing systems is appropriate
for a species and to develop optimal processing technologies with reference to
the welfare of the animals. Along with innate typical behaviours, the ethograms
must also include learnt behaviours to provide an overview of the full spectrum of
behavioural modifications.
This requirement is satisfied in this paper with a presentation of the behaviour of
calves in intensive, off-cow rearing. We investigated which behaviours can be
considered normal behaviour for calves reared in off-cow husbandry systems.
Another essential objective of this study is to identify behaviour that deviates from
the norm and to evaluate the effects of such deviations on the welfare of the
calves.
Of the various methodologies used in livestock ethology, we selected descriptive
behavioural recording in which the inventory of behaviours is identified in space
and time and described using means and variations to illustrate the behaviour of
calves in intensive rearing (HOY, 2009a).
RUSHEN et al. (2012) discussed issues with the use of automated methods to
measure behaviour in animals as part of evaluating animal welfare. Automatic
calf feeders can help to identify sick animals in group housing by making a com-
parison with average values. Devices attached to animals (e.g. accelerometers
or GPS trackers) measure the level of activity of animals and thus their well-being
with a high degree of accuracy. Automated image analysis offers significant po-
tential to evaluate movement within animal groups. Tracking individual animals
is, however, difficult to realise and the number of behaviours is still limited.
The authors of the current paper opted for recording behaviours using video cam-
eras but viewing of the videos and observing the behaviour of individual animals
was not automated. To process the data for statistical analysis, the program In-
teract from Mangold was used as it appeared best suited for this task.
The basis for displaying normal behaviour in cattle is the opportunity to move
freely (FLINT et al., 2016). This is ensured in the study farm.
Discussion 121
In the publication of the University of Applied Sciences in Neubrandenburg ‘Rear-
ing of female calves and young cattle in agricultural enterprises, Part 2: Influence
of CMR-feed supply in calf rearing on health, outputs and well-being’, the authors
describe the prerequisites related to feeding that must be satisfied to enable off-
cow rearing of calves in accordance with their needs and wants and thus to avoid
negative consequences associated with off-cow rearing (SCHULDT and DINSE,
2021a).
The farm where the behavioural studies presented here were carried out satisfies
the requirements of Germany’s Animal Protection – Livestock Ordinance regard-
ing the husbandry system used for calves. All animals are provided with a gener-
ous amount of space with straw bedding in the lying area and a concrete running
area. The animal to eating place ratio enables all calves to eat at the trough at
the same time. Until day 49, the animals are provided with a milk replacer allow-
ance of 12 L CMR-feed. Until the end of week 7, the calves demanded the full
allowance of 12 L CMR-feed on only 3% of the drinking days and no calf drank
12 L CMR-feed per day on average. It can thus be assumed that the calves were
largely able to cover their drinking needs. Weaning was carried out at a moderate
rate up to day 105 with the animals weaned on day 106. (SCHULDT and DINSE,
2021a)
To be able to evaluate the welfare of calves in off-cow rearing, a comparison of
the behaviour of the observed animals with an allowance of 12 L CMR-feed up to
day 49 and weaning from day 50 to day 105 (reduction of the CMR-feed allow-
ance by 0.22 L CMR-feed per day) with the normal behaviour of calves that vari-
ous authors have described for wild cattle or breeds kept in a manner close to
nature is discussed below (SCHLOETH, 1961; THORPE, 1965; KOCH, 1968; ZEEB
and MACK, 1970; SCHLICHTING and SMIDT, 1986; SCHEUNERT and TRAUTMANN,
1987; BROOM, 1991; TSCHANZ, 1995).
We review whether an ethogram based on the collected data on the behaviour of
calves in intensive rearing – at least 12 L CMR-feed until day 49, weaned by day
106 – is suitable to be described as exemplary of ‘normal behaviour of calves in
off-cow rearing’.
5.2 Behaviour of calves in off-cow intensive rearing
5.2.1 Circadian rhythm
Cattle, like all animals, have a 24-hour rhythm. Active and rest phases are dis-
tributed in a particular way over the whole day. (BARTUSSEK, 2008)
Discussion 122
A circadian rhythm for almost all behaviours can be observed in the calves from
as young as three to four weeks. In the second half of the night, from 12 midnight
to about 6:00 a.m., the calves rest for long periods and, with increasing age, con-
tinuously for several hours. Resting phases were included in the analysis if the
animals spent at least 1 minute lying down.
In the night during short wake periods the calves urinate and defecate, visit the
dispensing station and/or hay rack or trough or run seemingly aimlessly around
the pen for a while. Before calves lie down again, they seek out contact with the
other animals, smelling and licking them.
From 6:00 a.m. the calves start being active, in isolated cases from as early as
4:00 a.m. The younger calves first visit the dispensing station while older animals
increasingly start the day by eating supplementary feed and drinking water. Up
to 12 midnight, phases of intense activity alternate with resting phases but these
are considerably shorter than in the hours after midnight.
Because of these significant differences in the resting and active phases in the
circadian rhythm, we differentiated between night (12 midnight to 6:00 a.m.) and
day (6:00 a.m. to 12 midnight) when presenting the behaviour of the calves over
24 h.
The circadian rhythm develops independently of the milk replacer allowance and
of the start of work in the stables. The intensity and duration of weaning also does
not have an effect. (SCHULDT and DINSE, 2021a)
The circadian rhythm is described in many studies on the behaviour of calves.
On pasture calves suckle at any time of the day in principle but a circadian rhythm
can nevertheless be identified that is determined by the activities of the herd. All
calves drink at daybreak. Towards midday and at dusk suckling events are ob-
served more frequently. Around midnight there is a slight increase, otherwise ac-
tivities during the night are insignificant. (SAMBRAUS et al., 1978)
WAGNON (1963) observed that calves are suckled by the dam at all hours of day
and night but most commonly between 5:00 and 6:00 a.m. and often around
noon, towards evening and around midnight. With a stable dam–calf relationship,
RITTER and WALSER (1965) observed fixed individual suckling times among the
calves that occurred primarily between 9:00 and 11:00 a.m. and between 2:00
and 4:00 p.m. If a calf did not go to its mother of its own accord, she called for it
loudly.
In the first two months after birth, VITALE et al. (1986) observed the main suckling
phases between 9:00 and 11:00 a.m. and between 2:00 and 4:00 p.m. in semi-
wild free-ranging Maremma calves (Bos primigenius taurus).
Discussion 123
In a herd of Polled Hereford and Simmental cows, ODDE et al. (1985) observed
5.0 ± 0.1 suckling events in 24 h. Peaks in sucking activity occurred from 5:00 to
7:00 a.m., 10:00 a.m. to 1:00 p.m. and 5:00 to 9:00 p.m. The most suckling events
in an hour occurred between 5:00 and 6:00 a.m. and the fewest between 10:00
and 11:00 p.m. Similar peak suckling phases are also described by RIESE et al.
(1977).
In a herd in New Zealand containing different genotypes and in a herd of East
African Zebu cattle, the main suckling times were recorded at daybreak, midday,
in the evenings and between 10:30 p.m. and 1:00 a.m. After 1:00 a.m. almost all
the animals rested for several hours. Between 1:30 and 3:00 a.m., only a few
suckling events by very young calves were observed. (WALKER, 1962; REINHARDT
and REINHARDT, 1981)
On pasture, the duration of all activities by the calves differed significantly be-
tween night and day (SCHAKE and RIGGS, 1970).
Calves at automatic feeder had a specific, although certainly not rigid, cycle over
24 h in studies by ZEEB and MACK (1970). As described in the studies presented
in this publication, more intense activities started with a deep drink in the morn-
ings at about 6:00 a.m. Around 9:00 a.m. and from 8:00 to 9:00 p.m. resting times
were observed with additional suckling interactions around 11:00 a.m. and 4:00
p.m. After 9:00 p.m. the calves were active until about 1:00 a.m.
Sunrise and sunset, which are described as zeitgebers on pasture grazing
(WALKER, 1962; PORZIG et al., 1969; SÜSS and ANDREAE, 1974; FRASER et al.,
1978; SAMBRAUS et al., 1978; SCHEUNERT and TRAUTMANN, 1987), are of no im-
portance in the current analyses.
BREER and BÜSCHER (2006) confirmed in their studies on trial farms that there is
also a biorhythm apparent for activities in calves in off-cow rearing.
Feeding times for calves kept in barns are the primary zeitgebers with another
peak in activity observed by SAMBRAUS and STEINEL (1978) around midnight, in-
dependent of the administration of feed. This is also described by FRÖHNER
(2011), who identified a strong effect of feeding times on the circadian rhythm
while sunrise on the other hand did not trigger any locomotor responses.
Studies by UGWU (2020) suggest that the behaviour of dairy calves is affected by
day and sex but not by the husbandry system. The author prepared ethograms
using direct and video observations over two consecutive days of four male and
12 female calves that were kept in different stables. The influence of the time of
day was determined by comparing the ethograms with the behaviour of five
calves that was recorded on two days every hour between 10:00 a.m. and 5:00
Discussion 124
p.m. using a combination of direct and video observations. Although these results
need to be confirmed, they indicate that the behaviour of dairy calves can also be
influenced by the time of day.
Because the calves were consistently offered the high allowance of 12 L CMR-
feed over 24 h in the current studies, these observations cannot be confirmed.
The start of the workday of the farm personnel was also not identified as a zeit-
geber for the first feed-intake after the night-time rest. The activities of the calves
were most influenced by the farm personnel when the running area in the calves’
pen was cleaned and fresh bedding was spread on the lying area. Lying periods
were interrupted and shortened as a result. The calves were particularly active in
fresh straw, jumping about vigorously and playing together. Lying times were also
briefly interrupted by treatments or when individual calves were shown how to
freely access the CMR-feed in the dispensing station.
In the analysis of the resting data, FRÖHNER (2011) considered operational work-
flows on the farm, such as spreading fresh straw bedding, and observed intense
locomotor activities on freshly spread straw. KOCH (1968) also identified that feed-
ing times and the presence of humans had a greater effect on the activities of the
calves than dusk.
The rounds analysed in this publication were carried out in winter 2017, spring,
summer and autumn 2018. Across all seasons, CMR-feed-intake was signifi-
cantly more often from 6:00 a.m. onwards than between 4:00 and 6:00 a.m. Be-
cause younger calves mostly start their activities by eating feed, this comparison
suggests that in off-cow rearing in the barn, there are different timers operating
than on pasture. The number of early feeds from 4:00 a.m. in spring of 0.8 per
calf is slightly and not significantly above the mean for the other seasons (0.4 to
0.6). From 4:00 to 6:00 a.m. in summer a mean number of 0.4 CMR-feed-intakes
and from 6:00 to 8:00 a.m. of 1.2 CMR-feed-intakes per calf and day were rec-
orded. In winter the number was on average 0.6 from 4:00 to 6:00 a.m. and 1.0
from 6:00 to 8:00 a.m. When the sun was rising later, the calves thus started their
activities earlier.
5.2.2 Resting behaviour
Because the group size in the group pen of the study farm did not exceed 15 in
any of the rounds and there was sufficient space available, all the calves could
rest undisturbed. The calves lay down and stood up in the normal manner as
described by SAMBRAUS (1971), SCHEURMANN (1971) and SAMBRAUS (1985)
among others. All calves sought out lying surfaces with straw bedding to rest and
no calves lay down on the running area with no straw. Deviations from the normal
Discussion 125
lying down and standing up processes and short rest times indicate inadequacies
in the housing facility, such as insufficient available space or a substrate that is
too hard (GRAF et al., 1976; SAMBRAUS, 1985; KETELAAR-DE LAUWERE, 1989;
FERRANTE et al., 1998), which was not the case in the calf pen that was investi-
gated.
Features of the functional area ‘resting behaviour’ are established as animal-
based and husbandry-relevant indicators for the animal friendliness of husbandry
systems because a large part of the day is spent resting, meaning it is of consid-
erable importance for the animal. This behaviour can also be relatively clearly
defined and observed. Lying duration and frequency are used as analysable pa-
rameters. (SCHEURMANN, 1971; SCHLICHTING and SMIDT, 1986)
Until they were weaned in the current studies, the calves rested for 14.6 to 16.5 h
a day with a resting period before weaning lasting on average 41.9 ± 41.0 to 49.8
± 46.4 minutes with a large range of up to 6.1 h at night and 4.8 h during the day.
During weaning, the mean duration of the resting periods increased significantly
to up to 67.4 ± 61.4 minutes in the daily average with a maximum of 6.1 h. The
number of resting periods from week 3 to 7 is on average between 17.3 and 23.2
in 24 h. During weaning, the calves rested on average for 14 to 18 times a day
for 48.3 to 67.4 minutes. The differences in the means are significant between
the age groups (up to day 49 vs. day 50 to 105) in the number and duration of
the resting periods.
The calves rested at night on average for 65.4 to 128.1 minutes continuously
throughout the entire group housing phase of the drinking period with a rising
trend, while during the day they rested on average for 33.3 to 52.2 minutes. As
indicated by the age-dependent change in the resting times, the total of the daily
lying period decreased from week 8 to week 15 in the calves investigated here.
On pasture calves spend 80% of the day lying down in the first days of life while
they rarely stand for longer than 15 minutes (SAMBRAUS et al., 1978).
The duration of the lying and standing periods changes up until day 43. The pro-
portion of nocturnal standing time decreases in young calves in favour of standing
time during the day. At the age of 40 days, calves lie down for 14 h per day while
later they only lie down for 12 h. (SCHEURMANN, 1971; PORZIG, 1987)
Calves of fighting cattle raised in semi-wild conditions in the French Camargue
form groups a few days after birth, lie down in groups and rest for most of the 24-
hour day (SCHLOETH, 1961; KOCH, 1968; ZEEB and MACK, 1970). In the first days
of life, individual lying phases last initially for 2.5 h and shorten until day 5 to 1.5 h
(LANGBEIN et al., 1998).
Discussion 126
FRÖHNER (2011) established that calves kept in stable groups and fed using au-
tomatic calf feeders had an average total daily lying time of 18.4 h. SUTHERLAND
et al. (2018) counted an average of 17.4 ± 1.16 lying periods a day in healthy
calves.
Lying times in the current studies were often interrupted by the calves when, for
example, dry cows or pregnant heifers ran on an adjacent paddock or vehicles
drove past the stable. All the animals usually jumped up suddenly and ran to the
edge of the pen to watch the event. This curiosity-driven behaviour is also de-
scribed by SAMBRAUS (1971) in calves on pasture where young cattle spend their
resting times, rushing over to something that catches their attention.
In the current studies, the calves often lie down at night in the same place where
they had previously stood. In contrast, during the day the lying areas were
changed frequently. Before lying down, the animals intensively sniffed the ground
and any calves that were already lying down, which were also very often licked.
This behaviour was recorded particularly at night when the animals were only
briefly active. KILEY-WORTHINGTON and PLAIN (1983) also observed this behaviour
in suckler cow husbandry.
Calves rest in off-cow rearing for just as long and distributed across day and night
in the same way as in mother-bonded calf rearing (WALKER, 1962; SAMBRAUS et
al., 1978; PORZIG and ENGELMANN, 1991). Calves prefer to lie down near the walls
(SAMBRAUS, 1971; SAMBRAUS and STEINEL, 1978). These observations, regarding
both the lying duration and the locations, were confirmed. Lying positions against
the walls were also preferentially sought out in our studies.
Calves spend about 70% of their time lying down until week 8 (CHUA et al., 2002).
This corresponds to the resting time of the calves with the 12 L CMR-feed-allow-
ance (CMRA) until day 49. Up to five weeks of age, the animals rested for 14.6
to 16.5 h during the day or 61% to 69% of 24 h. The calves also spend most of
their early lives sleeping in suckler cow husbandry (KILEY-WORTHINGTON and
PLAIN, 1983).
In the present studies, calves often lay down again in the 5 minutes before and
after resting, which occurred with 34% and 35% of the resting periods respec-
tively. Before 30% and after 22% of the resting phases, the calves ate supple-
mentary feed, often hay from the rack on the rear wall of the lying area. If the
animals only briefly interrupted a resting period, they usually defecated and/or
urinated, which was not recorded separately here, however. Before resting
phases, the calves often sought out contact with other animals, which made up
12% of the activities 5 minutes before lying down. The third most common activity
after resting is CMR-feed-intake followed by blind visits and water-intake with 5%
Discussion 127
and 6% respectively. A higher proportion of CMR-feed-intakes than dummy visits
after a resting phase confirmed the feeding rhythm that had developed in the
calves.
In the period between week 3 and 7, a calf cross-sucked on another calf for about
4 minutes after resting once it had nibbled on some hay in the rack.
5.2.3 Food and water-intake behaviour
5.2.3.1 Drinking behaviour
To be able to evaluate the drinking behaviour of off-cow reared calves, their drink-
ing behaviour is compared to natural behaviour in mother-bonded rearing. This
method is not consistently defined, however (KUNZ, 2017). In this publication com-
paring the drinking behaviour with natural suckling behaviour refers to rearing of
calves with their mothers as is practised in suckler cow husbandry, for example.
This method mostly relates to beef-producing breeds, which must be taken into
account when considering the MR volumes.
Another method is mother-bonded and fostered calf rearing in dairy husbandry,
which is recommended by some associations in organic farming (BARTH et al.,
2009; SPENGLER NEFF et al., 2017). However, because the calves are not assured
of free access for milk intake in this method, it cannot be equated to natural
mother-bonded rearing.
In the current studies, an CMR-feed meal lasted on average 4.1 to 4.9 minutes
up to day 49 with a slight increase in the duration as the animals get older. The
number increased from week 3 to 4 from 4.3 to 5.3 meals per animal and day and
thereafter remained constant at five to six meals. The duration and number of
daily CMR-feed-intakes is therefore in the same range as the number of times
calves suckle from their mother. The mean duration of an CMR-feed-intake in-
creases with slow weaning from 4.6 minutes in week 8 to 6.3 minutes in weeks 9
and 10. From week 11 to 15, an CMR-feed meal lasted on average 3.5 to 4.8
minutes. Over the entire long weaning phase, the average duration of an CMR-
feed-intake was significantly above the mean for a full milk replacer allowance
(4.9 vs. 4.6 minutes). The number of CMR-feed meals decreases because of the
declining CMR-feed-allowance , which also reduces the frequency of supply, from
week 7 to 10 to a daily average of three and up to week 14 to two per day. The
four CMR-feed meals in week 15 is due to the higher frequency of milk replacer
intake of one of the two calves that were assessed.
On average, calves suckle for five to ten minutes at their mother, with the suckling
time varying greatly between calves from one to 30 minutes, with a total daily time
Discussion 128
of between 15 and 115 minutes or a daily average of 45–60 minutes (WALKER,
1950; WAGNON, 1963; SCHEURMANN, 1974b; RIESE et al., 1977; SAMBRAUS and
STEINEL, 1978; KILEY-WORTHINGTON and PLAIN, 1983; ODDE et al., 1985; FRASER
and BROOM, 2002). AHMED (1987) reported a total 22 minutes during the day with
154 seconds per suckling event on average for different calves at their mothers.
In mother-bonded rearing, calves only drink a small volume at a meal, in some
cases less than a litre (KASKE, 2018b). Older calves suckle less frequently at their
mother than younger calves, with the duration of a suckling event consequently
increasing. On average, younger calves suckle up to eight times a day and older
calves up to six times. (W
ALKER, 1962; HAFEZ and LINEWEAVER, 1968;
SCHEURMANN, 1974b; NICOL and SHARAFELDIN, 1975; ARAVE and ALBRIGHT, 1981;
PORZIG and ENGELMANN, 1991)
With increasing age, the duration of suckling as well as the daily number de-
crease but not the duration of the individual suckling events (SCHRADER, 2007).
In the first few days of life, calves suckle up to eight times for less than 1 minute,
in the first month of life as a rule not more often than six times, and at 3 months
three to at most five times for 5 to 10 minutes each during the day (WALKER, 1962;
SCHEURMANN, 1974a; RIESE et al., 1977; SAMBRAUS et al., 1978; SOMERVILLE and
LOWMAN, 1979; KILEY-WORTHINGTON and PLAIN, 1983; SAMBRAUS, 1985; PORZIG,
1987). However, the duration of a suckling event does not provide any indication
of the volume of milk ingested with only a weakly positive correlation between
these two parameters (CAMERON, 1998).
A meal according to PORZIG (1987) lasts 9 to 15 minutes and shortens with in-
creasing age of the calves only from month 6 to 8, with 60 to 140 minutes suckling
duration observed in 24 h. SOMERVILLE and LOWMAN (1979) noted approximately
the same number of suckling events at night as during the day in Charolais
crosses. At night, however, the individual meals are shorter (SAMBRAUS et al.,
1978).
On the other hand, the age, breed and sex of a calf did not influence the suckling
frequency or duration according to ODDE et al. (1985)
Calves from one month of age suckled at their dam for 8 to 10 minutes, in isolated
cases up to 30 minutes, suckling at each teat for 5 to 30 seconds and equally
from the right and left side, for a total of 60 to 70 minutes distributed over the day
(PORZIG, 1964; SCHEURMANN, 1974b; PORZIG and ENGELMANN, 1991; SAMBRAUS,
1992, 1997a). The hunger of a calf is sated within 10 to 15 minutes of a suckling
event at the mother (WALKER, 1950; HAFEZ and BOUISSOU, 1975). If this takes too
long for the cow, she becomes restless, kicks out towards the calf with her rear
legs or moves away (SAMBRAUS et al., 1978).
Discussion 129
SPENGLER NEFF et al. (2017) consider four to five suckling periods per day each
over about 10 minutes in weeks 2 to 8 to be natural behaviour, from which they
deduce that calves in mother-bonded and fostered calf rearing should be able to
suckle several times a day for a total of 50 minutes.
In a comparison of calves with and without dam contact aged from 1 to 3 months
(Schwarzbuntes Milchrind × Salers crosses), SCHLEYER (1998) measured 25, 40
and 59 minutes’ suckling time daily at the mother over three to four suckling
events vs. 32, 26 and 23 minutes’ drinking time at automatic calf feeders over
about six sucking events. The sucking duration of calves with no dam contact
reduced in month 3 to 2 to 3 minutes per sucking bout on average. The calves
spent about the same time at the dispensing station and played with the teat. The
suckling bouts of the calves with dam contact at this age lasted 15 to 20 minutes.
In both groups the total sucking activity made up about the same time and in-
creased with age. With the automatic feeder calves, the number of unrewarded
and dummy sucking bouts increased at the expense of milk intake.
With free access to the dispensing station, a similar sucking duration was ob-
served for calves in off-cow rearing as for mother-bonded calves (HAFEZ and
LINEWEAVER, 1968; WEBSTER and SAVILLE, 1982).
With ad libitum supply, male Holstein Friesian calves ingested CMR-feed from
the automatic feeder for a total of 8.1 meals over 57.0 minutes in 24 h with a meal
lasting on average 7.1 minutes (WEBSTER and SAVILLE (1982), Table 8).
Table 8: Ethogram of feed-intake behaviour of male HF calves in group housing
with automatic feeders at the age of 8 weeks
Paramete
r
In 24 hours
Numbe
r
of CMR-feed-intakes 8.1 (± 2.2)
Mean duration of CMR-feed-intakes in minutes 7.1 (± 1.6)
Total CMR-feed-intake in minutes 57.0 (± 13.0)
Source: quoted at WEBSTER and SAVILLE (1982)
WAGNON (1963) concluded that calves suckle for longer if they get too little milk
from their mother. He noted 44 minutes suckling time on pasture with supplemen-
tary feeding and 55 minutes without. Calves aged up to 3 months suckled 3 to 11
times a day at the dams with an average of 4.7 suckling periods a day noted by
WAGNON (1963) About 83% of the time the calves sought the dams out, otherwise
the cows called their young to suckle. A meal was usually ended by the calves
(81%) but in 13% of cases the cows ended the suckling.
In studies by BORDERAS et al. (2009) with an ad libitum supply, calves drank milk
7 to 12 times a day from the automatic feeders. Healthy calves visited the dis-
pensing station 6.7 times ± 0.93 a day in studies by SUTHERLAND et al. (2018).
Discussion 130
RIESE et al. (1977) observed that the first eating time for calves reared in group
housing was after the morning drink.
In the present studies, 53% of the activities in the 5 minutes before an CMR-feed-
intake were resting, while 21% of the activities before an CMR-feed-intake were
eating supplementary feed. After a successful visit to the dispensing station, the
calves rested in 28% of the observed activities with eating supplementary feed
recorded equally as often. Playing and drinking water occurred with almost the
same frequency (9% to 14%) before and after an CMR-feed-intake. No calf cross-
sucked on another calf in the first weeks of life immediately before or after drink-
ing. A repeat visit to the dispensing station within ±5 minutes with or without CMR-
feed-intake was made only in 1% to 6% of cases.
5.2.3.2 Behaviour associated with supplementary feed-intake
The hay rack is on the rear wall of the group pen of the study farm and thus at
the back of the lying area. The trough for the dry TMR is on the front left wall of
the running area. The youngest calves made short, frequent visits to the trough
and rack up to week 6 and amused themselves almost playfully with the supple-
mentary feed, initially only nibbling the hay. The intake of hay and dry TMR ap-
pears at first to be less motivated by a feeling of hunger rather than seen as an
opportunity for entertainment. That calves ingest hay before drinking rather as a
distraction was also observed by PORZIG and ENGELMANN (1991).
Up to the end of week 7, supplementary feed-intake (hay and dry TMR) in the
present studies lasted on average 1.5 to 3.1 minutes and was observed in the
daily average 10.7 to 18.8 times per animal and day. No difference was identified
here between the duration of the intake of hay and dry TMR. In terms of the num-
ber of meals, there is a clear rising trend in the number of visits to the trough from
week 4 to 7 (11.6 to 19.4 per animal and day). The number of hay intakes in-
creased in the same period only slightly from 6.0 to 9.3 per animal and day. In
week 7 the number of longer eating periods lasting more than ten minutes in-
creased to 3% with the eating time remaining under five minutes in 83% of cases.
With the start of weaning, on the study farm for the supplementary feeding dry
TMR was gradually replaced by a TMR that is offered in young cattle rearing after
weaning. This mixture is placed in front of the front opening of the pen enclosure.
The supplementary feed in the weaning phase thus includes hay and TMR with
a decreasing proportion of dry TMR.
For the observed calves, the duration of the supplementary feed-intake increased
significantly from week 8 during weaning. This is also apparent in the percentage
of eating periods lasting more than 10 minutes, which is 7% in week 10, rises to
Discussion 131
11% in week 12, and reaches the highest values of 23% and 21% in weeks 13
and 14 respectively. However, only the trough feed-intake increased to about 7
minutes per intake while the hay intake remained at an average of 4–6 minutes
per meal. A maximum of 59.7 minutes was recorded. The supplementary feed-
intake follows the circadian rhythm in terms of its frequency. At night in 6 h only
8.5% of the intakes were solid food before weaning and 7.4% during weaning.
Before and after about one-third of the supplementary feed-intakes, the calves
rested, with several eating periods often shortly following each other, particularly
in the weaning phase. Older calves often eat together, particularly when fresh
supplementary feed is provided.
Eating activity has a rhythmical structure over 24 h. Over 16 h of observation,
RIESE et al. (1977) recorded eating times of 4.2 h in calves kept in suckler cow
husbandry. In indoor housing, calves started the first meal, which lasted 30
minutes to one hour, after the morning drink. Additional meals were observed at
around midday and after the evening drink each of about one hour. In the running
pen, the calves spent considerably less time eating than with full-day pasture
access. The number of eating periods was accordingly lower. The authors sus-
pect that the grazing takes longer overall than ingesting hay and concentrate.
From day 7 to 42, calves ate hay for an average of 12.7 minutes a day with an
allowance of 12 L whole milk in studies by ROSENBERGER et al. (2016). Until day
43 the intake of concentrate by calves fed ad libitum was also low in studies by
BORDERAS et al. (2009).
Feeding silage should not be started before week 5 according to VAN ACKEREN
(2013). From the perspective of the behaviour associated with trough feed, we
must agree with this.
In the current study until week 7 resting periods were most commonly observed
within ±5 minutes before and after a visit to the trough or rack (27% and 35%
respectively). The calves also often played or switched between the trough and
rack. Visits to the dispensing station as well as drinking water and contacts be-
tween the calves occur a little less often before eating. Until week 7 no cross-
sucking activities on other calves were observed in connection with supplemen-
tary feed-intake.
At night meals lasted on average only 15 minutes in a number of studies, while
during the day the calves ate for a total of about three hours and thus significantly
longer (SCHAKE and RIGGS, 1970; PORZIG and ENGELMANN, 1991). Humans act as
a generator of rhythm in that the animals are always fed and watered at specific
times (PORZIG and ENGELMANN, 1991).
Discussion 132
Calves aged 2 to 4 months also eat at night, but this was no longer observed from
5 to 6 months (LIEBENBERG, 1965).
Studies by MILLER-CUSHON and DEVRIES (2015) showed that behavioural patterns
such as feed-intake behaviour that develop at an early age persist once they have
been learned and can have long-term effects on health and well-being. For ex-
ample, providing ad libitum access to a feeder not only leads to more growth but
also to meal patterns that closely resemble the natural behaviour of a calf that
suckles at its mother. However, the physical form and presentation of the feed
can also affect the feeding behaviour of calves, particularly feed sorting, which
can affect both the direct intake of nutrients and the development and persistence
of this behaviour.
Before supplementary feed-intake, PORZIG and ENGELMANN (1991) observed wa-
ter drinking, which was not the case in the studies presented here. In our own
studies, the calves rested more often or ate feed from the trough or rack. Play
was also recorded before eating hay or trough feed.
5.2.3.3 Behaviour associated with water-intake
It is frequently mentioned in the literature that calves should be offered fresh wa-
ter ad libitum right from the start (SCHRAG et al., 1987; BEEDE, 1993; ULBRICH et
al., 2004; SPRENG, 2011; HERRMANN, 2014; VAN ACKEREN, 2014; HORN, 2019).
On the study farm, calves have access to fresh water ad libitum from automatic
drinkers. The calves quickly adapt to the automatic drinkers and use them without
having to be taught. After the adjustment to the group housing, the animals drank
water on average 2.4 to 5.6 times daily from week 4 to the end of week 7 with the
mean number of drinks rising with age and increasing supplementary feed-intake.
Until weaning the number of drinks was up to ten times a day. Along with the
frequency, the mean duration of the water-intake increased slightly, but only to
1.2 minutes on average.
SPRENG (2011) also noticed increased water-intake with age in terms of frequency
and volume.
The calves observed in the current studies rarely interrupted the night rest up to
week 7 to drink water and only 8.3% of all visits to the automatic drinker were
recorded between 12 midnight and 6:00 a.m. During the weaning period, the per-
centage of nocturnal water-intake over the total intake in 24 h decreased to 7.0%
because of the longer rest periods. From 6:00 a.m. the water-intake follows the
circadian rhythm. During the day the calves drink most frequently when they are
particularly active, that is, in the morning, early afternoon and evening.
Discussion 133
According to OLSON and WILLIAMS (1960) cited in PORZIG and ENGELMANN (1991),
calves drink water up to seven times a day.
Young calves only get sufficient fluids through CMR-feed in the first few days of
life (SCHRAG et al., 1987; PIATKOWSKI et al., 1990). The minimum requirement for
the calves in the first 3 weeks is 1 L water per animal and day (ULBRICH et al.,
2004). Because a calf needs about 10% of its body weight in water daily, the
requirement in the first 3 months of life is a total of 8 to 12 L fluids (milk meals +
water) per day (SCHRAG et al., 1987; MEYER et al., 2005; HORN, 2019).
There is no justifiable reason to withhold drinking water from young calves. Feed-
intake and growth rate are increased by providing access to freely available drink-
ing water. Under heat-stress conditions, the water requirement is 1.2 to 2 times
higher. (BEEDE, 1993)
Up to day 30, the water-intake in studies by NELLE et al. (2005) remained at a
consistently low level. With the increase in concentrate intake, the water-intake
also increased from day 31. However, it must be noted that these calves were
only supplied with a low allowance of a maximum 8 L CMR-feed until day 23.
5.2.3.4 Visits to the dispensing station without CMR-feed-intake (blind visits)
Visits to the dispensing station with no CMR-feed-intake are triggered by the mo-
tivation to suck and thus to ingest liquids (SCHULDT and DINSE, 2020a), which is
why this behaviour is classified as food intake behaviour in these studies. A blind
visit was coded if a calf entered the dispensing station and remained there, with-
out ingesting CMR-feed, but showing a clear need to suck by playing with the
teat. Entering the dispensing station with no interest in the teat was not evaluated
as a blind visit and was included in the assessments for locomotion.
From week 4 to 7 the daily average for visits to the dispensing station without
CMR-feed-intake was 2.1 to 3.2 visits and a blind visit lasted on average 0.7 to
1.0 minutes. Only 8% of the blind visits lasted more than two minutes with a max-
imum of 6.5 minutes recorded. In 70% of cases the calves that needed to suck
spent less than one minute in the dispensing station if unrewarded.
In the publication of the University of Applied Sciences in Neubrandenburg on
intensive animal-friendly calf rearing, the authors describe the adjustment of the
calves to group housing. The young calves learn very quickly if they have an
allowance in the dispensing station. Within eight days the number of dummy visits
falls from 15 in week 1 to eight in week 2. With an CMR-feed-allowance of 12 L,
the management program of the dispensing station recorded a daily average of
2.1 to 3.5 visits with no allowance after the adjustment period. Most animals need
less than one week to develop a circadian rhythm in their drink intake. Until day
Discussion 134
49, 53% to 60% of the calves only entered the station if they wanted to drink and
thus had no unrewarded attempts. (SCHULDT and DINSE, 2021a)
In studies by NIELSEN (2008) large portions combined with a low flow rate
(300 ml/min) extended the duration of the visits with an allowance but with no
drink intake. This may have been only play behaviour and thus should also not
have been evaluated as a visit with sucking motivation.
PIRKELMANN (1981) reported that most of the calves had adjusted to the automatic
feeders in 1 to 2 days. An adjustment period of 3 to 5 days only had to be calcu-
lated for a few calves.
BORDERAS et al. (2009) also observed that a high milk volume reduced the num-
ber of unrewarded visits to the dispensing station. The authors report that calves
adjust readily to the rhythm of milk availability and can thus reduce the number
of blind visits.
In studies by ROSENBERGER et al. (2016) with an allowance of 12 L CMR-feed per
animal, up to day 42 an average of 0.4 ± 0.8 visits with no drink intake per day
were recorded.
SUTHERLAND et al. (2018) recorded a daily average of two to four visits to the
dispensing station with no milk allowance for healthy calves that had an allow-
ance of 2 L whole milk supplied three times a day over a total of at least 360
minutes.
In the weaning phase the number of unrewarded visits to the dispensing station
increased because of the decreasing allowance but the duration did not change.
If the calves were rapidly weaned over 20 days, the number of blind visits rose to
a daily average of 16.9 in week 10 (SCHULDT and DINSE, 2021a). With slow wean-
ing up to day 105, this high number was not recorded with the maximum being
11.8 visits per animal and day in week 13. The mean number then fell to a daily
average of 5.3 in the last week before weaning was complete. Longer blind visits
lasting ≥2.0 minutes only made up 6% of visits during weaning up to day 70
(SCHULDT and DINSE, 2021a) but with slower weaning this figure was 14%. This
difference corresponds to the number of unrewarded attempts and is an indica-
tion that calves in this age still have a pronounced need to suck and more rest to
act this out when they are weaned at a moderate rate.
In this study blind visits frequently occurred after resting when a new allowance
was still not available. The older calves also frequently sought out the dispensing
station without success after ingesting supplementary feed. In the entire drinking
period, several consecutive unrewarded visits to the dispensing station were rec-
orded in 15% to 16% of the blind visits. Playing and contacts in the 5 minutes
Discussion 135
before and after dummy visits did not play a role with older calves, while younger
calves visited the automatic drinker, rack or trough before and after blind visits or
played with equal frequency. Up to day 49 the calves had ingested CMR-feed
before 8% of the blind visits, indicating that the time to eat a meal obviously was
not sufficient to satisfy the need to suck. The blind visits corresponded to the
circadian rhythm in their duration and frequency. At night before weaning 9.5%
of all unrewarded attempts were recorded with a mean duration of 0.7 to 1.2
minutes, while during the day the duration of a blind visit increased to 6.5 minutes
maximum with an average of 0.7 to 1.1 minutes. During weaning the frequency
of blind visits at night fell to 7.8%.
High milk volumes and gradual weaning reduce the number of visits to the dis-
pensing station with no allowance compared to abrupt weaning (LIDFORS and
ISBERG, 2003; DE PASSILLÉ et al., 2004). During the weaning phase, the number
of visits with no allowance by animals that previously had ad libitum access in-
creased to a daily average of 6.9 (PATT et al., 2017), which corresponds to our
findings. For ROSENBERGER et al. (2016) a high number of visits with no allowance
indicates that the calves are constantly hungry.
5.2.4 Social behaviour
About more than half of the respondents in a survey conducted by BUSCH et al.
(2017) believe that calves should be separated later from the cow. Particularly for
reasons of animal welfare, this practice is disputed, however (JOHNSEN et al.,
2016).
Practitioners explain the reasons behind the early separation to defuse the emo-
tionally charged debates. The separation distress for the calves with earlier sep-
aration is lower and pens are also not currently set up for shared cow–calf rearing.
Additional arguments for early separation are the lower microbial pressure and
controlled colostrum intake on the first day as well as an increase in the occur-
rence of diarrhoea due to excessive milk intake with restricted contacts with the
mother. (BARTH et al., 2009; EVERS, 2021; LAHMANN, 2021)
In off-cow rearing of calves, the mother–child relationship is missing in the func-
tional area of social behaviour. This can only be experienced by cow and calf in
a suckler herd as is conventional in beef husbandry. Even with foster mothering,
in which cows suckle two to four calves, possibly including their own, the relation-
ship between mother and calf is disrupted or even completely severed. Usually,
the foreign suckling calves are only endured but not adopted. In mother-bonded
rearing, in which calves are only allowed to go to their mother to suckle, cow and
Discussion 136
calf cannot freely choose the contacts, which means that a true bond cannot form.
(ROTH, 1978; VOIGT, 1996; EHRLICH, 2003)
The social behaviour of off-cow reared calves is limited to contacts between sim-
ilarly aged calves in a group and is described in the present studies with the eval-
uation ‘calf–calf contacts’ that is expressed in mutual sniffing and licking. The
exception to these contacts is cross-sucking, although it does result in some
cases from the contacts between the calves (SCHULDT and DINSE, 2020a). The
calves lick each other particularly intensely before lying down. In week 3 the most
frequent contacts between the calves were observed with 8.0 per animal and day,
which is arguably related to combining the animals into a group. Calves smelled
and licked each other from week 4 to 7 on average 4.6 to 5.6 times a day for 1.3
to 2.0 minutes each time and with a rising trend. At night there were fewer con-
tacts between the calves compared to the day because of the longer resting
phases (7.0% of all contacts in 6 h). At the start of weaning the number of con-
tacts increases and then declines with the duration remaining on average about
1 to 2 minutes per contact. Slow weaning manifested in the present studies as a
moderate decline in the number of daily social contacts.
Calves love to be stroked on the neck by humans or to be licked by other calves
(LÜRZEL, 2019). They scratch and lick themselves or brush or rub themselves
against objects, behaviour that increases significantly with age (KILEY-
WORTHINGTON and PLAIN, 1983).
Overall, calves spend on average 25 minutes a day licking themselves or other
calves (GROTH, 1978). SCHLOETH (1961) observed that cattle lick one another for
up to 10 minutes, also with brief interruptions and changing of roles.
Over about 80 hours of observation, SAMBRAUS and STEINEL (1978) recorded 7.7
± 5.7 licking activities per calf in off-cow rearing, with some animals preferably
licking specific members of the herd. Off-cow reared calves had fewer contacts
among each other but mutually licked more often than suckling calves (SCHLEYER,
1998).
Spatial proximity between dam and calf is an important component of the bond
(BRENNINKMEYER et al., 2005). Rearing with contact with the mother in the first 12
weeks can, even if such contact is very limited, have consequences for later be-
haviour and possibly lead to improved social competence in dairy cows (WAGNER
et al., 2012; WAGNER et al., 2015).
Studies on the effects of three different rearing systems (free, half-day [6:45 a.m.
– 6:00 p.m.] and no maternal contact in the first nine weeks of life) on lying be-
haviour after incorporation of heifers into the dairy cow herd showed that the dam-
Discussion 137
reared heifers exhibited less stress caused by entering the herd. Other effects of
the rearing conditions, such as on the productivity, were not identified. (ZIPP and
KNIERIM, 2015)
Abrupt separation of the calves that were reared with their mothers induces stress
responses that have a similar effect on behaviour as the pain after dehorning
(DAROS et al., 2014).
In a review of studies MEAGHER et al. (2019) conclude that early separation (within
24 h of birth) reduces the acute stress responses of cows and calves. Although
longer contact amplifies the acute stress responses between cow and calf on
separation and reduces the quantity of saleable milk while the calves are being
suckled, it can have positive effects in the longer term on the behaviour of calves,
encouraging more normal social behaviour while reducing abnormal behaviour
and sometimes the response to stressors. However, the authors remark in their
summary of the studies that only a few studies described indicators for long-term
well-being effects other than abnormal and social behaviour of the calves.
Many other authors also point out that contact with the mother has a positive
effect on the development of the young (BARTH et al., 2009; WAGNER et al., 2012;
WAGNER et al., 2015; BEAVER et al., 2019).
In the behaviour of the young calves in the group phase, the authors of the pre-
sent study observed no signs that the calves are stressed by the absence of the
mother–child relationship.
5.2.5 Other activities and play behaviour
In the present study activities were predominantly observed during the day, while
at night, if the animals had rested deeply for long periods, the calves only briefly
drank and ate supplementary feed in their wake phases. In the wake phases,
liquids and supplementary feed were ingested and the calves engaged in other
activities such as play, locomotion or social contacts between the calves. This
was observed particularly from 6:00 to 12 midday and 4:00 to 10:00 p.m.
When calves are reared to day 49 with an allowance of 12 L CMR-feed per animal
and day, they spend 8.4 to 6.9 h a day engaging in locomotion, exploratory and
play behaviour up to week 6, which corresponds to 91% to 85% of the active time
with a decreasing trend. In week 7, the percentage is 80% (6.0 h).
The authors described the influence of the milk replacer allowance, that is, the
intensity of the calf rearing, on the locomotion, exploratory and play behaviour in
a number of publications. It is apparent that ‘other activities’ reach a level of 80%
and more only with intensities of at least 12 L CMR-feed per animal and day
Discussion 138
(SCHULDT and DINSE, 2018a, 2019b, c, 2020c, 2021a). As a result of the increas-
ing supplementary feed-intake, the proportion of other activities decreases during
weaning. In week 13 such activities make up 59% of the active time while in the
other weeks it ranges from 66% to 83%.
A drastic decline associated with intensive or rapid weaning is closely related to
the rapid increase in the dry feed-intake and should be considered another sign
of a higher stress load on the calves. As a parameter for evaluating animal wel-
fare in calf rearing, the proportion of other activities should be given particular
attention. Until weaning, these activities should make up more than 80% of the
active time and the level should not fall below 60% during weaning. (SCHULDT and
DINSE, 2021a)
5.2.5.1 Locomotion and exploratory behaviour
In the present study, locomotion behaviour includes standing without ingesting
food or water, play or social behaviour, slow and fast running, jumping and gal-
loping. The duration is calculated from the difference between the data for all
activities and the food intake.
In the present studies, the calves moved individually through the pen but often
encouraged one another to romp about, often and intensively in the evening
hours from 4:00 to 10:00 p.m. in particular. Up to week 7, the daily locomotion
was observed for a daily average of 8.0 to 5.7 hours per animal with a decreasing
trend. This decline can be attributed to the gradually increasing duration of sup-
plementary feed-intake.
Licking and sniffing objects, such as pipes or walls, is a behaviour belonging to
the functional area of exploratory behaviour and was performed while standing
or slowly walking and was recorded in six calves up to week 7. After increasing
from 11.7 to 15.5 minutes per animal and day from week 3 to 4, from week 5 to
weaning it remained at a daily average of 11.4 to 6.5 minutes per animal. The
somewhat higher value at the start of group housing confirms the curious nature
of the calves and the associated intensive exploration of the pen after being in-
troduced to the group housing. The duration of the licking remains constant at 1.3
to 3.3 minutes per activity.
The intensity of the exploratory activities changes with the age of the cattle in the
form of an inverted U. Initially, the intensity increases among the calves. Of all
age groups, weaned but not yet sexually mature young cattle are the most inquis-
itive and engage most intensely in exploratory activities. With the onset of sexual
maturity, the intensity declines again. (MURPHEY; DUARTE; et al., 1981)
Discussion 139
In studies by RIESE et al. (1977), the locomotion activities of calves aged 2 to 12
weeks started to decline right from the start. Over the course of the day, a brief
active phase was followed in the morning by a longer rest time, and towards
evening the calves were again very active and covered longer distances.
BREER and BÜSCHER (2006) investigated the activity of off-cow reared calves us-
ing pedometers. In the early morning hours and the evenings, the calves were
particularly active while at night and in the afternoons the animals rested. The
level of activity in the afternoon was considerably higher than overnight. With in-
creasing age, the number of activities increased but with considerable differences
between individual animals. These studies, however, did not provide any infor-
mation about the type of activity during movement.
5.2.5.2 Play behaviour
Play is described as a purely instinctive action (BROWNLEE, 1954; TEMBROCK,
1964). TEMBROCK (1958) defines play as behavioural sequences the function of
which cannot be directly derived from the actions or their effects and that do not
lead to a fundamental change in the environment of the individual concerned.
Play trains muscles, joints, tendons and ligaments that are otherwise only used
for fighting, fleeing or reproduction but also trains the circulatory and respiratory
systems. When playing, animals also gain confidence in executing the behav-
iours that are necessary for developing a stable hierarchy and sexual behaviour.
(WEINREICH, 1968; FRÖHNER, 2011)
Play behaviour encourages group dynamics through active invitations to a play
mate and helps to train cognitive strategies and develop social relationships.
Playing together is thus more effective than playing alone. (FAGEN, 1981)
The calf play investigated in the current study (with toys or headbutting) was ob-
served across the entire drinking period in the circadian rhythm, that is, less often
and only briefly at night, more often after the first intake of CMR-feed and/or sup-
plementary feed, and quite pronounced in the afternoon and evening from about
4:00 to 10:00 p.m. Until week 7, this play lasted only a little more than a minute
ranging from 0.04 to 9.1 minutes, with a total of 7.6 to 12.6 minutes a day with a
maximum of 36.1 minutes in which two calves played with one another or a toy
in weeks 6 and 7.
During the weaning phase, calves played for up to 20.2 minutes a day with a toy
or each other. Play lasted on average 1.3 to 2.0 minutes and for each calf 5.1 to
10.0 play periods on average were counted during the day. For these slowly
weaned calves, the pronounced circadian rhythm is also seen in play behaviour.
At night the calves only played a little.
Discussion 140
This play duration in off-cow reared calves is confirmed by GRÖßBACHER et al.
(2020). With an allowance of up to 12 L milk per animal until week 8, each day
play activities were observed for a total of 15.2 minutes in 18 h (Table 9). In these
studies, the CMR-feed-allowance but not the age of the calves had a significant
effect on the play frequency and duration. With higher ambient temperatures, the
total and mean duration of play fights and games between several calves de-
creased, which may serve as an indicator of heat stress according to the authors.
Table 9: Mean number and duration of play activities of calves within 18 hours
with a CMR-feed-allowance of 9 – 12 L milk per animal and day up to week 8
Game Number and duration in 18 hours
Fighting games
Single games
Total
58.4 ±
3.3 in 15.3 ±
0.4 seconds
11.6 ±
0.7 minutes
15.2 ±
0.9 minutes
Source: GRÖßBACHER et al. (2020)
On pasture healthy, well nourished calves play predominantly in the late after-
noon, less around midday and often after suckling or feeding (BROWNLEE, 1954;
PHILLIPS, 1993). PORZIG (1964) also observed play behaviour only at specific
times of the day, particularly after feeding. SANTHA (1977) recorded the most in-
tense play activities of calves at dusk and in the morning, regardless of the milking
or feeding times of the herd, which confirms the circadian rhythm of play behav-
iour observed in the current studies.
VALNÍČKOVÁ et al. (2015) did not identify any significant differences in the play
behaviour of calves that remained with their mother for the first four days of life
and calves of the same age that were immediately separated from their mother.
If there is plenty of space available, calves enjoy running and jumping and prefer
long spaces over square spaces (JENSEN and KYHN, 2000; MINTLINE et al., 2012).
A social hierarchy does not play a role in suckling calves and forms at the earliest
between three to six months or at sexual maturity. Even among younger calves
in group housing, quarrels were observed, which cannot be viewed as an expres-
sion of the superiority of one calf over another, however. (SCHLOETH, 1961;
LIEBENBERG, 1965; SAMBRAUS et al., 1978; SAMBRAUS, 1985; FRÖHNER, 2011;
SCHULDT and DINSE, 2021a, b)
Headbutting (Figure 71) is therefore evaluated in the current study as play be-
haviour in accordance with studies conducted by SAMBRAUS and STEINEL (1978).
Calves often start to headbutt when moving to the group housing when they ac-
tively move through the pen, run quickly, jump or gallop, which is assessed as
‘locomotion’ in the current studies. Usually only two calves headbutt each other
Discussion 141
and several calves alternating headbutting was rarely observed. The activity is
initiated by all calves and it is not uncommon for smaller animals to incite larger
members of the group to play.
Figure 71: Headbutting calves
Playful headbutting was observed by REINHARDT (1980) from week 2. HÜNERMUND
(1969) only observed hierarchical disputes after weaning and therefore assumes
that there is no hierarchy for calves up to the age of six months. In contrast,
SCHLEYER (1998) reported hierarchical disputes from 3 months of age but only
after weaning.
A comparison of the play behaviour between calves with mother contact and
calves without contact with their mother in studies by WAIBLINGER et al. (2020)
indicated greater well-being among the contact calves. This was manifested by
frequent, individual locomotor play activities. Calves without mother contact initi-
ated agonistic games more often, which could contribute to the development of
higher social competence.
The group pen used for the studies presented in this publication contained ele-
ments for play to test options for the calves to occupy themselves. The calves
play constantly with the ball hanging on a chain in a tyre (Figure 72 (a)), often
with several playing together at the same time. The animals suck on the chain
and hit their snouts against the ball so that it swings, which appears to be of
particular interest for the calves. The central position in the running area of the
pen is advantageous, because the animals must always move past the toy when
moving from the front running area to the lying area or the dispensing station. The
game is often triggered when a calf randomly butts against the ball and starts it
swinging.
Discussion 142
The brush is located on the right side of the front wall of the pen (Figure 72 (b)).
The animals play more frequently with the brush as they get older, with the calves
brushing their snout or forehead against the brush fixed to a steel spring or they
sniff or lick it. The actual function of the cattle brush of enabling scratching of
head and neck is rather the exception before week 7.
(a) (b)
(c)
Figures 72: (a) – (c): Toys for calves (a) ball, (b) brush, (c) chains
GEORG and UDE (2007) investigated the acceptance of an automatic cleaning
machine by 144 calves. 98% of the calves used the brushes at any time of day
and night and more heavily in the evening hours from 8:00 to 10:00 p.m. The
animals preferred to clean their heads with the horizontal brush but their neck
and rump considerably less and only as they got older. The automatic operation
of the brush was triggered when a calf gently raised the brush using its head,
which the calves frequently did as they became more familiar with the brush,
about 30 to 40 days after entering the group housing saturation was reached and
no additional increase was recorded. Playing with the brush was gradually re-
placed by its active use for skin care.
The chains were installed to the front wall of the pen as part of the ongoing studies
in this investigation in the second week of the first round (Figure 72 (c)) so that
the calf, which moved into the group housing in week 3, was not able to
Discussion 143
immediately play with them. The types of play were therefore only assessed from
week 4 on. The chains are sucked by the calves and, because they also move,
are made to swing. As described for the ball, the animals often unwittingly bump
into the chains, which starts them moving and the calves then play briefly with
them.
The ball was preferred by the calves, making up 52.9% of the frequency and
40.0% of the duration of all play activities up to week 7. On average the calves
played four to five times a day with the ball for a few seconds to 7.9 minutes.
18.3% of the play was spent playing around with the chains, even though they
are not located directly in the running area. One calf played continuously with the
chains for 9.1 minutes. The brush, which made up 18.5% of the play activities,
was played with for just as often and long (18.5% of the play period) as the chains
(20.8% of the play period). During the weaning phase about half the time is spent
playing with the ball while the other three toys are used for about the same length
of time.
In studies by MORROW-TESCH (1997), only two of 23 calves played with freely
hanging chains. A ball lying in the pen also did not stimulate the calves to play.
The author nevertheless views enrichment in calf pens as a useful instrument to
improve the housing environment.
Calves often play after feeding, in new but not strange environments and with
familiar moving objects such as swinging items (THORPE, 1965).
Studies with fattening calves were intended to show how different toys affect the
well-being of the animals and how often they are used during the day. However,
only the daytime activities of the calves were assessed and then only on three
consecutive days at pre-, mid- and end-fattening. The calves preferred a moving,
hanging ‘toy hedgehog’ that they could suck and a scratching wall, showing much
weaker preference for a chain. The scratching wall was only used for body care
rather than as a toy, however. If only one play element was available, several
calves also played with it at the same time. As the take-home message, the au-
thors recommend installing play material in farms where animals are displaying
behavioural disorders. (ZIRON and DICKS, 2019)
We cannot agree with this conclusion without comment, however. Behavioural
disorders are an indication of inadequate housing and feeding conditions. There-
fore, the causes of the abnormal behaviour must first be clarified before employ-
ing a symptomatic treatment. A toy is an opportunity for calves to play but is no
substitute for an insufficient milk supply, for example, which does not satisfy the
calves sucking motivation and may encourage cross-sucking and other behav-
iours.
Discussion 144
A comparison of the behaviour of calves and contamination of their skin with soil-
ing with no environmental enrichment (control group) vs. a group with a cattle
brush vs. with a Jolly Ball™ showed that calves in the control group lay down for
significantly longer than the calves with environmental enrichment and also ran
and jumped significantly less often. The social behaviour of calves with environ-
mental enrichment was more pronounced and the calves with the brushes
groomed themselves less than the animals in the control group. There were no
differences in the play behaviour between calves with the brush or Jolly Ball™.
Regarding skin contamination, there were no differences between any of the
groups. (BULENS et al., 2014)
PEMPEK et al. (2017) see advantages in providing toys. The calves used all ob-
jects – artificial teats, a stationary brush, a calf lolliball and rubber chain links –
but depending on the time of day, they spent the most time playing with the brush.
Calves that were housed in hutches with a toy spent almost 50% more time with
locomotion games, but also the same amount of time sucking on objects, as
calves that were housed in standard hutches. There were no effects observed on
growth, feed-intake or behavioural responses or in response to social or environ-
mental novelties after weaning.
The motivation for calf play behaviour increases for some but not all elements
over time in studies by BERTELSEN and JENSEN (2019). Therefore, differentiating
between play elements can be important when using calf play behaviour to eval-
uate animal welfare. The link between the health of a calf and headbutting indi-
cates that a healthier calf initiates more social play.
Healthy animals that are fed according to their needs and housed appropriately
play more often than sick, hungry animals (BROWNLEE, 1954; PORZIG, 1964;
THORPE, 1965; AHLOY-DALLAIRE et al., 2018). When they experience negative
events (e.g. pain), play behaviour may also be reduced, which is why it should be
observed as an indicator of the well-being of young animals (JENSEN et al., 1997;
JENSEN et al., 1998; MANTEUFFEL, 2006; DUVE et al., 2012; MINTLINE et al., 2013;
RUSHEN et al., 2016; GRÖßBACHER et al., 2020). JENSEN et al. (2015) concluded
that more intensive feeding leads to longer periods spent playing, which suggests
an advantage for animal welfare. This can also be concluded from our own stud-
ies.
Discussion 145
5.2.6 Sucking activities without visiting the dispensing station
5.2.6.1 Cross-sucking on another calf
Cross-sucking among cattle has long been a known behaviour. In the older liter-
ature it was described as ‘self-sucking’ by cows (ZÜRN and MÜLLER, 1885), while
from the middle of the last century it was referred to as ‘milk sucking’, ‘sham
sucking’, ‘inter-sucking’ or ‘mutual sucking’ of calves, young cattle and cows
(SCHOTT, 1956; HEIDRICH and RENK, 1963; PORZIG, 1964; KITTNER and KURZ,
1966; MOTSCH et al., 1975; REINHECKEL, 1975; SCHLÜTER et al., 1975; KELZ, 1977;
SÜSS and SEBESTIK, 1982; KAPHENGST, 1984).
With an allowance of 12 L CMR-feed per animal in the current studies, cross-
sucking was only observed by one animal up to day 49. In week 6 this calf cross-
sucked once for 9.7 minutes and in week 7 three times on three days each time
for an average of 3.1 minutes. Overnight there were no cross-sucking activities
among the observed calves in the five rounds up to day 49. In the weaning phase
three of the five observed calves cross-sucked. One calf must be considered to
be a ‘cross-sucker’. It began in week 14 and a total of three to five events were
recorded on four days. The other calves only cross-sucked 1 to 3 times on one
to three days. Usually the calves only cross-sucked briefly with a mean duration
of 2.1 minutes and only two events were recorded at night.
On 13 of 212 evaluated days, 19 cross-sucking activities by five calves were rec-
orded. Of these, 62
% occurred within 60 s of a visit to the dispensing station and
therefore must be classified as ‘nutritive’. In their publication ‘Investigations into
cross-sucking and possibilities of reducing this during calf rearing’ (SCHULDT and
DINSE, 2020a), the authors suggest not using milk intake as a criterion, that is,
differentiating between ‘nutritive’ and ‘non-nutritive’, that is, as defined by RUSHEN
and DE PASSILLÉ (1995) and KASKE (2018a), but based on the sucking motivation,
which is the actual cause of cross-sucking, and thus differentiating between
‘sucking activities’ and ‘non-sucking activities’. Sucking activities refer to those
sucking events that occur after CMR-feed-intake and/or unrewarded visits to the
dispensing station. All other cross-sucking events are ‘non-sucking activities’,
even those after eating supplementary feed, which would have to count among
the ‘nutritive’ sucking activities based on the content. Only by differentiation
based on the sucking motivation is it clear that this behaviour is not causally re-
lated to a feeling of hunger but instead with the latent, always present sucking
need of young calves.
Individual calves cross-suck and even maximum CMR-feed allowances do not
prevent this. Cross-sucking cannot be eliminated but can only be limited by
Discussion 146
satisfying the sucking motivation as much as possible by the drink intake, which
requires an allowance of at least 12 L CMR-feed up to day 49. (SCHULDT and
DINSE, 2020a)
NIELSEN (2008) did not record any effect of the portion sizes on the proportion of
cross-sucking calves but there was a reduction in the percentage of animals that
cross-sucked for 60 minutes or more after drink intake.
To be able to evaluate a behaviour, we should ask not only ‘why?’, that is, what
is the biological function or purpose of a behaviour, but also ‘how?’, that is, what
is the causal control. The functional consideration asks about the adaptation and
function of a behaviour, about the causes that have led to its evolution. The
causal control asks about the immediate causes, including triggering stimuli.
(WECHSLER, 1993)
Cross-sucking is genetically determined and is considerably more pronounced in
breeds such as Brown Swiss, Simmental or Jersey than in German Holstein and
is part of the natural spectrum of behaviours of calves. It has developed to give
calves a chance of survival when their mother cannot provide enough milk. These
calves very quickly learn how they can suckle on foreign cows without being
fended off by them. Up to three suckling calves on one cow were observed by
SCHÄFFER et al. (1999) in a suckler herd. This is even wanted because this helps
with rearing calves whose mothers provide insufficient milk. Calves attempt to
suckle on different cows, which is most likely to succeed when the dam’s own calf
is being suckled, with some cows caring for three or four calves without being
able to differentiate between their own and foreign calves (ARAVE and ALBRIGHT,
1981). In studies by FRÖBERG and LIDFORS (2009), most calves that were kept
together with their mothers in a barn with an automatic milking system attempted
to suckle on foreign cows at least 1 or 2 times.
Calves usually suckle a foreign cow from behind between the legs, which is also
the most common variant of cross-sucking on other calves along with reversed
parallel, which is preferred when suckling from a calf’s own mother. Suckling from
behind was also observed by RITTER and WALSER (1965) as a normal position for
calves at their own mother.
If calves and young cattle also intensively cross-suckle after weaning, they should
be excluded from breeding. This is particularly recommended if the mother of
these animals is also known to be a cross-sucker. Along with providing high CMR-
feed allowances, this can also help to modify the cross-sucking problem by breed-
ing to minimise the issue in the herd. (SCHULDT and DINSE, 2020a, 2021a)
Discussion 147
Cross-sucking is not related to a hierarchy (SAMBRAUS and STEINEL, 1978). The
close bond observed between calves with this behaviour resembles the bond with
its own mother. SAMBRAUS, H.H. (1993) classified cross-sucking in the functional
area of ‘eating behaviour’, however.
It can be assumed that calves also cross-suck in mother-bonded rearing, but it is
only rarely observed and described.
SCHLOETH (1961) observed young calves playing ‘mother and child’ in Camargue
cattle kept in semi-wild conditions. One calf approached another in the posture it
would use to approach its mother and started, with bunting movements, to play-
fully suck in a relaxed manner between stomach and hindquarters. In older young
animals, the suckling posture occurred at the end of games with sexual over-
tones. SAMBRAUS and STEINEL (1978) recorded 0.6 (SD = 1.0) cross-sucking ac-
tivities in a suckler herd over 80 hours of observation.
Of 14 calves that were suckled by their mother, ROTH et al. (2008) observed one
calf cross-sucking another calf. At 10 weeks of age, the likelihood that an animal
cross-sucked another or not peaked. The frequency of cow–calf contacts,
whether twice a day or unlimited, had no effect on cross-sucking, which suggests
that permanent contact with the dam has minimal effect.
HULSEN (2015) published a photo (Figure 73) in which one calf on pasture cross-
sucks a calf standing in front of it from behind through the legs and recommends
separating the cross-sucker from the other animals because it will presumably
display this behaviour throughout its life.
Photo: HULSEN (2015)
Figure 73: A calf suckles another calf on the pasture
5.2.6.2 Sucking on dummy teats
A ‘Milk Bar’ calf feeder with ten teats (Figures 10 (a) (b)), which was used to
provide the calves with electrolytes, was installed in the group pen in the current
Discussion 148
study farm in rounds 1 to 3 as a sucking dummy in addition to the dispensing
station. The feeder stayed empty in the pen during the day and was used by the
animals as a sucking dummy. In rounds 4 and 5 it was investigated whether
calves accept sucking dummies that do not provide any liquids. Two boards each
with four blank plugs made of plastic (dummy boards) were attached to the wall
next to the dispensing station and to the outer gate at the height of the calves’
heads.
There are assessable data available from ten calves on the use of the dummy
teats. On 56 days a total of 183 sucking events were recorded of which 27.7%
were overnight. On average, the calves sucked on the teats of the empty calf
feeder for 1.0 to 1.5 minutes with extreme values of 8.1 and 12.4 minutes as a
maximum and on the dummy boards for 0.6 to 1.1 minutes, which corresponds
to the duration of the dummy visits to the dispensing station.
The calves sucked more often on the teats of the calf feeder than on the blank
plugs of the dummy boards. Whether this is related to the nature of the teats,
which resemble those of the dispensing station, or whether the calves received
electrolytes through the dummy teats could not be investigated. The blank plugs
of the dummy boards are rigid and not malleable, which is possibly why the calves
did not suck on them as often or for as long.
Sucking on a dummy teat was followed predominantly by visits to the dispensing
station, trough or rack with 24% and 38%.
Three of six calves sucked exclusively on the dummy teats (dummy board) at the
gate, which was not specifically sought out for sucking, however. Usually the
calves sucked on the blank plugs if they had initially only sniffed and licked them.
Obviously, playing with the plugs triggered the sucking motivation.
No clear relationship between cross-sucking of calves and use of the dummy
teats could be confirmed. It can therefore be assumed that the dummy teats can-
not be used to limit cross-sucking or even to prevent it.
Which elements the calves prefer for sucking was examined by MORROW-TESCH
(1997) in a group pen. The author compared the use of empty small and large
Kongs (a rubber toy for dogs that resembles a snowman in shape) with a feeding
bottle and a calf lolly, which was fixed to a pipe filled with treacle flakes. When
the calves sucked on the lolly, the treacle flakes fell out through small holes in
the pipe. At the start of the studies, the small Kong was most frequently sucked
on, while in week 2 it was the feeding bottle. The duration of sucking on the bottle
increased with the age of the animals while the other elements were used with
the same frequency. The small Kong toy was better accepted than the large.
Discussion 149
5.3 Normal behaviour of calves in intensive off-cow rearing
Artificial selection by humans has affected the adaptability of the animals to
changing environmental conditions. Selecting particular characteristics in domes-
ticated animals could also affect the behavioural flexibility of the individuals along
with their general learning performance because resources are diverted to
productivity parameters. Study results from NAWROTH et al. (2021) indicate that
selecting for milk production may have affected behavioural flexibility in goats.
Thus, breed-specific differences in the adaptation to changing environmental
stimuli can affect the parameters for species-appropriate husbandry.
Compared to life in the wild, animals in every husbandry system are subject to
restrictions, the extent of which are determined by the structural and engineering
features of the husbandry system and by their management (FLINT et al., 2016).
Husbandry systems always represent a compromise between different require-
ments such as investment costs, labour economics and animal welfare (BRADE,
2005).
Deviations from normal behaviour are signs that the environment is not capable
of satisfying the needs of the animals. We therefore discuss below which behav-
iours indicate inadequate feeding and housing conditions and thus can be used
as indicators of calf rearing that is not animal friendly.
Some clearly natural behaviours are, however, detrimental to animal welfare. This
includes emergency behaviours, such as flight responses that shift the animal to
a stress state, or even harmful behaviours, such as hierarchical or illness-related
aggression in which other animals are injured or deprived of resources. Never-
theless, considering these caveats, natural behaviour can be used as a guide for
improving existing husbandry systems. (ŠPINKA, 2006)
For VAN PUTTEN (1982) a behaviour can be considered ‘normal’ if the appetitive
behaviour leads an animal to find a specific stimulus by which it can completely
satisfy its need.
Behaviours occur in a particular context or are executed against an appropriate
object. Only as an exception can one and the same behaviour occur in different
functional areas and belong in each case to normal behaviour. A behaviour that
is defined as ‘normal’ must not necessarily satisfy a function as is, for example,
regularly observed with play. Although the motivation for play is high, particularly
in young animals, it rarely leads to consummatory responses. Nevertheless, play
is included in normal behaviour. (SAMBRAUS, 1997b)
From about 1960, the behaviour of agricultural livestock became a topic of inter-
est, initially only in the comparison with free-range animals. In older specialist
Discussion 150
literature, behaviours were only mentioned if they deviated from the norm.
(SAMBRAUS, 2002)
Animal checks should be conducted twice a day in calf rearing, which according
to SCHÄFFER et al. (2007) includes physiological signs (breathing, lameness and
posture) and
behaviour
in
addition
to
the
requirements
of the
German
ordinance
on
the protection of animals kept for farming purposes (monitoring,
feeding
and
care).
A frequently used method is comparative studies in which the same features can
be collected on pasture as a reference method for the behaviour of cattle
(SCHRADER, 2020).
Features of behaviour show a normal distribution with deviations within a 95%
limit considered ‘normal’ if they correspond to a particular type (TSCHANZ, 1985).
Certain behavioural forms are environmentally stable and have barely changed
as a result of domestication and breeding. Differences between wild and domes-
ticated animals involve the degree to which behaviours manifest and the fre-
quency of occurrence of behaviours with increases and decreases possible. New
behaviours that are not simply a variation of a basic form have not been observed
to date. (TSCHANZ, 1984)
SAMBRAUS (2002) differentiates between categories of behaviours that deviate
from the norm using sequences of movements that do not belong to the behav-
ioural repertoire of the animal species; behaviour directed toward an object that
does not serve to satisfy a need; vacuum activities; behaviour that causes harm
to the acting animal or the recipient; changes in the frequency or duration of the
behaviour as well as apathy. Drink intake that is shortened at each meal because
of a sucking speed that is too fast can be assigned to these categories. Social
contacts that are limited to contacts between calves of the same age and are
elevated compared to calves in a suckler herd can also be allocated using this
category.
According to GRAUVOGL (1983), frustration that develops because a behaviour
cannot be expressed is only relevant for animal welfare if the animal does not
behave ‘correctly’ as a result.
For the intensively reared calves observed in the current studies, this is obviously
not the case. Cross-sucking occurred only sporadically but the frequency during
weaning was significantly higher than when calves are reared with their own
mother. No other behaviours that deviate from the norm were observed in the
calves that were studied.
Discussion 151
BRENNINKMEYER et al. (2005) reported on practical experiences whereby the in-
tensity of the separation response increased with the duration of the time the cow
and calf spent together.
This was confirmed in studies by FLOWER and WEARY (2003) that established that
the responses of the calf to separation increases when the calf has spent more
time with the cow. Nevertheless, longer contact has advantages, particularly re-
garding sociality, anxiety and future maternal behaviour. Health, weight gain and
productivity are also improved if the calf can spend time with the cow.
The absence of the mother had no effect on the behaviour of Friesian calves in
studies conducted by LE NEINDRE et al. (1979).
The study presented here on the behaviour of the calves with a high CMR-feed-
allowance up to day 49 revealed similarities and differences to the natural behav-
iour of calves in mother-bonded rearing. Correlations are seen in the circadian
rhythm, in the resting behaviour in terms of lying down and standing up, the rest-
ing positions and the duration of the resting hours. In the functional area of feed-
intake behaviour, we observed a similar age for when calves started to increase
their solid-feed-intake. Locomotion behaviours, inquisitiveness and the need for
play and social contacts with animals of the same age are further correlations.
An innate sucking motivation enables free drink intake from the sta tion in a rhyth m
that is also seen when suckling from the mother.
Differences in the behaviour of mother-bonded and off-cow reared calves are
listed in Table 10.
Table 10: Differences
in
the
behaviour
of
mother-bonded
and
off-cow
reared
calves
Categorie Calf rearing
Mother-bound Motherless
Daily rhythm
Zeitgeber Sunrise and
sunset
Possibly sucking motivation,
thirst, hunger
Food-intake behaviour: milk- or CMR-feed-intake
Mean duration of a session 5
–
10 minutes 4
–
6 minutes
Unsuccessful attempts at weaning
or weaning
Not described 2 to 3 in 24 hours,
each about 1 minute
Sucktion activities
Sucking, cross-sucking Rarely
described
Of individual animals, in-
creased in the weaning phase
Sucking on substitute objects
(dummies)
Not described Independent of feed-intake
Sources: WALKER (1950); WALKER (1962); WAGNON (1963); SCHEURMANN (1974b); RIESE et al. (1977);
SAMBRAUS et al. (1978); SAMBRAUS and STEINEL (1978); KILEY-WORTHINGTON and PLAIN (1983); ODDE et al.
(1985); AHMED (1987); SCHÄFFER et al. (1999); (FRASER and BROOM, 2002); FRÖBERG and LIDFORS (2009)
Discussion 152
Unrewarded attempts at the dispensing station can be compared to calves being
fended off by their own mother during weaning and can therefore be classified as
a normal behaviour. Because these attempts only occur briefly and only up to 3.2
times a day until weaning is complete, they represent only a slight stress load for
the calves. Several blind visits follow within 5 minutes for about one-fifth of the
subsequent activities only, with the calves frequently ingesting supplementary
feed or water. With smaller CMR-feed-allowances, up to 13 blind visits were ob-
served on average and the duration of each visit is also considerably higher with
means of 5 to 12 minutes (SCHULDT and DINSE, 2021a). BRUMMER (2004) reported
up to 150 visits with no allowance per day in Simmental calves that were reared
with 12 L CMR-feed-allowance up to day 20 and 10 L CMR-feed from day 30 to
42, that is, considerably less than in the study farm used in the present behav-
ioural observations.
The zeitgebers for the start of activities in the morning and the start of the night
rest cannot be used as indicators of animal welfare because their motivation in
off-cow rearing can only be surmised.
The duration of a drink intake should be adjusted to that of natural milk intake by
reducing the sucking speed. Because the calf feeders that are currently mostly in
use are usually set to a sucking speed that is too high (SCHULDT and DINSE,
2020b, 2021a), the duration of an average CMR-feed-intake should be used to
evaluate animal welfare of the feed supply.
The parameters ‘unrewarded attempts at milk or replacer intake’, which can be
evaluated in off-cow rearing as ‘visits to the dispensing station without CMR-feed-
intake’, and cross-sucking are suitable indicators of the well-being of calves. Both
parameters can be objectively and numerically collected and analysed. The
dummy visits can be recorded by the calf feeder software and the cross-sucking
by behavioural observations. Calves that intensively cross-suck are usually
known to farm personnel and such events should be documented.
Sucking on dummy teats is not a suitable indicator for animal welfare because it
is not related to abnormal behaviours.
Enriching the housing environment of animals with play elements encourages
cognitive capacities and has a positive impact on well-being (ELIZABETH BOLHUIS
et al., 2013; GRIMBERG-HENRICI et al., 2016; LECORPS et al., 2022). Even if the
calves in the current studies only briefly play with the play equipment such as the
ball or chains, it can be assumed that providing
such
elements benefits
well-being.
Discussion 153
5.4 Ethogram for calves in off-cow rearing
With CMR-feed-allowances of at least 12 L per animal up to day 49 and weaning
from day 105, the behaviour of the off-cow reared calves largely corresponds to
natural behaviour in mother-bonded rearing.
Circadian rhythm
Calves develop a distinct circadian rhythm that underlies almost all behaviours.
The behaviour at night, that is, from 12 midnight to about 6:00 a.m., with long
resting phases interrupted briefly by defecation and urination as well as ingesting
CMR-feed, supplementary feed and/or water, must be differentiated from activi-
ties during the day, that is, from 6:00 a.m. to 12 midnight. During the day, the
calves alternate activities and resting phases, which are considerably shorter
than overnight.
Resting behaviour
The calves seek out protected places, such as corners or near walls, to rest. After
a brief interruption, they often lay down in the same place where they had stood
up. The typical resting positions correspond to those familiar from behavioural
observations of herds on pasture. Young calves rest daily for 14 to 18 hours for
an average of 30 to 45 minutes during the day and for 60 to 180 minutes at night.
In the mornings and evenings the resting times are usually somewhat shorter
than over midday.
Feeding behaviour
The sucking behaviour at the dispensing station corresponds to the natural suck-
ing at a cow in terms of the posture, bunting and tail movements as well as the
number of four to five meals per day. The individual meals in the present studies
are each four to six minutes long with a rising trend and are therefore shorter than
when suckling on the cow. Increasing the suction resistance should increase the
sucking duration, so that a natural speed of about 0.2 L/min is achieved, which
was determined in mother-bonded rearing and is recommended as the optimum
for feeding with calf feeders (5–7 min/L milk, STEINHÖFEL and LIPPMANN (2000);
KASKE (2018b)).
Visits to the dispensing station without feed-intake occur briefly up to day 49 for
less than one minute on average. In the week when the calves are adjusting to
the free feed-intake, up to four of these dummy visits per animal and day are
tolerable, thereafter one to three up to weaning. In the weaning phase, slightly
longer blind visits in an increasing number can be considered normal if they do
not significantly exceed a daily average of 10 per animal.
Discussion 154
Up to the end of week 7, calves frequently ingest supplementary feed (hay, trough
feed) over the day with only a short eating duration. The number increases to
about 20 meals per day and the duration to about 3 minutes per meal. The calves
drink water from the start for about one minute and the frequency and duration
increase with the intake of supplementary feed.
Because sucking on animals other than the mother, e.g., on another cow, is part
of natural behaviour, cross-sucking of calves in a group cannot be prevented.
Individual animals have a distinct sucking need that cannot be satisfied through
the feed-intake even with the highest CMR-feed-allowances. If these are isolated
cases, the cross-sucking can be tolerated. Sucking calves from a maternal suck-
ing family should be excluded from breeding, however. Dummy teats are ac-
cepted
by
the
calves
but
cannot
prevent
cross-sucking
and are
used less
than a
moveable toy.
Social behaviour
Calves are herd animals and maintain intensive social contacts. They smell and
lick each other, play together and rest closely with one another. Hierarchical dis-
putes are not observed before weaning.
Locomotion and play behaviour
Young animals have a marked need to move. Calves run, jump and gallop
around, often together and encouraging each other. Intense activities are often
observed in the evening hours. Playful headbutting starts as early as week 2.
Calves prefer to use moveable objects that they can make swing as toys. These
toys are licked, sucked or sniffed, often by several calves at the same time. It is
beneficial to place a toy in an area in the pen where the calves primarily run and
where randomly knocking against the freely hanging objects stimulates the calves
to play.
Locomotion and play behaviour can be summarised as ‘other behaviours’ to eval-
uate the well-being of calves. The percentage of active time spent on these be-
haviours should be a daily average of at least 80% up to day 49. During weaning,
the percentage of other activities decreases because of the increasing intake of
supplementary feed but should not be less than 60% of the active time. During
weaning over at least 50 days, the animals rest for long periods, visit the dispens-
ing station not too often without an allowance, and reach a high intake of supple-
mentary feed upon weaning. Often the older calves stand together for a longer
period at the rack or trough and eat hay or trough feed.
Summary 155
6 Summary
The studies on the behaviour of female German Holstein calves in intensive off-
cow breeding were carried out in an agricultural enterprise in Mecklenburg-Vor-
pommern. The calves were kept in hutches with a run to the end of week 4 at the
latest. The calves were moved from week 3 to subsequent group housing in a
pen with a litter-covered lying area and concrete running area. The maximum age
difference within a group was 4 weeks. The calves were reared with a maximum
daily allowance of 12 L CMR-feed up to day 49 and weaned on day 106. Over
the entire feeding period, CMR powder (50% skim milk fraction) was used at a
concentration of 160 g/L water. Hay and dry total mixed ration (TMR) were used
as supplementary feed provided fresh daily ad libitum in racks and troughs with
TMR gradually replacing the dry TMR during weaning.
The calf pen was fitted with elements to investigate which activity options were
preferred by the animals. A tyre freely hanging on a rope with a ball (ball), a cattle
brush for calves (brush) and board with four chains (chains) were installed in the
pen. It was also investigated whether the calves accepted dummy teats.
The behaviour of the calves was recorded continuously over 24 hours with video
cameras from the day they moved into the barn to the day they moved out. In
total, data from 13 calves in five rounds over 212 days with 4,569.5 hours of video
recordings were coded with the Interact program from Mangold and statistically
processed and analysed by week of life (week 3–7 and week 8–15) or day of life
(day 50 to 105) with Interact and Excel 2019 MSO from Microsoft (Version 2207).
The analysis of behaviour with maximum allowance (12 L CMR-feed per animal
up to day 49, n = 9 calves) was carried out for the functional areas rest, food and
water-intake, social behaviour (calf–calf contacts), and ‘other activities’, which in-
clude exploratory, play, locomotion and elimination behaviours. Exploration (lick-
ing of objects) and play were coded, while locomotion (standing without activity,
slow and fast running, jumping, galloping) and elimination were calculated from
the difference in the duration of the activities and the behaviour coded in this
phase and summarised as locomotion behaviour. The use of toys, visits without
CMR-feed-intake and sucking activities (cross-sucking, sucking on dummy teats)
were assessed separately.
Summary 156
From the studies, an ethogram for calves in off-cow rearing can be derived:
Circadian rhythm
Calves develop a distinct circadian rhythm that underlies almost all behaviours.
The behaviour at night, that is, from 12 midnight to about 6:00 a.m., alternates
between long resting phases with brief wake phases for defecation and urination
as well as the intake of CMR-feed, supplementary feed and/or water. During the
day, from 6:00 a.m. to 12 midnight, the calves alternate between active and rest-
ing phases, which are considerably shorter than overnight.
Resting behaviour
The calves seek out protected places to rest. After a brief interruption, they often
lay down in the same place where they had stood up. Young calves rest daily for
14 to 18 hours for an average of 30 to 45 minutes during the day and for 60 to
180 minutes at night. In the mornings and evenings the resting times are some-
what shorter than over midday.
Feeding behaviour
The sucking behaviour at the dispensing station corresponds to the natural suck-
ing at a cow in terms of the posture, bunting and tail movements as well as the
mean number of four to five meals per day. The individual meals last on average
four to six minutes with a rising trend.
Visits to the dispensing station without milk intake occur briefly up to day 49 and
last less than one minute on average. In the week when the calves are adjusting
to the free feed-intake, up to four of these blind visits per animal and day are
tolerable, thereafter a daily average of one to three up to weaning. In the weaning
phase slightly longer blind visits in an increasing number can be considered nor-
mal if they do not significantly exceed a daily average of 10 per animal.
Up to the end of week 7, calves frequently ingest supplementary feed over the
course of the day with only a short eating duration. The number increases to
about 20 meals per day and the duration to about three minutes per meal. The
calves drink water from the start for about one minute and the frequency and
duration increase with the intake of supplementary feed.
Individual animals have a distinct sucking need that cannot be satisfied through
the feed-intake even with the highest CMR-feed-allowances. If these are isolated
cases, cross-sucking of another calf can be tolerated. Sucking calves from a ma-
ternal sucking family should be excluded from breeding, however. Dummy teats
are accepted by the calves but cannot prevent cross-sucking and are used less
than a moveable toy.
Summary 157
Social behaviour
Calves smell and lick each other, play together and rest closely with one another.
Hierarchical disputes are not observed before weaning. There were no signs in
the behaviour of the young calves that indicate that the calves are stressed by
the absence of the mother–child relationship.
Locomotion and play behaviour
Calves run, jump and gallop around, often together and encouraging each other.
Intense activities are often observed in the evening hours. Playful headbutting
starts as early as week 2.
Calves prefer to use moveable objects that they can make swing as toys. These
toys are licked, sucked or sniffed, often by several calves at the same time. Lo-
comotion and play behaviour can be summarised as ‘other behaviours’ to evalu-
ate the well-being of calves. The daily average of the percentage of the active
time spent on these behaviours should be at least 80% up to weaning. When
weaning, the percentage decreases because of the increasing intake of supple-
mentary feed but should not be less than 60% of the activities.
Weaning
Moderate weaning is recommended from the perspective of animal behaviour so
that the animals rest for long periods, only make few unrewarded visits to the
dispensing station and are ensured of having a high supplementary feed-intake
upon weaning. Because the maximum milk replacer allowance must be provided
up to day 49, this results in a recommended weaning age of at least 105 days.
Zusammenfassung 158
Zusammenfassung
Die Untersuchungen zum Verhalten von weiblichen Kälbern der Rasse Deutsche
Holsteins in der mutterlosen intensiven Aufzucht wurden in einem landwirtschaft-
lichen Unternehmen in Mecklenburg-Vorpommern durchgeführt. Die Kälber wur-
den bis maximal zum Ende der 04. Lebenswoche in Iglus mit Auslauf gehalten.
Die Umstallung in die anschließende Gruppenhaltung erfolgte ab der 03. Lebens-
woche in eine Box mit eingestreuter Liegefläche und betonierter Lauffläche. Die
maximale Altersdifferenz innerhalb einer Gruppe betrug 4 Wochen. Die Kälber
wurden mit einem maximalen täglichen Tränkeanrecht von 12 l Milchaustau-
schertränke (MAT) bis zum 49. Lebenstag (LT) aufgezogen und am 106. LT ab-
gesetzt. Über die gesamte Tränkeperiode wurde eine MAT mit Milchaustauscher-
pulver (50
% Magermilchanteil) in einer Konzentration von 160
g/l Wasser einge-
setzt. Als Beifutter wurden Heu und Trocken Totale Mischration (T-TMR) täglich
frisch ad libitum in Raufe und Trog vorgelegt, in der Abtränkphase ersetzte eine
TMR allmählich die T-TMR.
Die Kälberbucht wurde mit Elementen ausgestaltet, um zu untersuchen, welche
Beschäftigungsmöglichkeiten von den Tieren bevorzugt werden. Dafür wurden
ein frei an einer Kette hängender Reifen mit einem Ball (Ball), eine Viehbürste für
Kälber (Bürste) sowie ein Brett mit 4 Ketten (Ketten) als Spielzeug in der Bucht
installiert. Weiterhin wurde geprüft, ob die Kälber Saugattrappen annehmen.
Das Verhalten der Kälber wurde mit Videokameras durchgehend über 24 Stun-
den vom Tag der Einstallung bis zum Ausstallen aufgezeichnet. Insgesamt wur-
den von 13 Kälbern aus 5 Durchgängen über 212 Tage mit 4.569,5 Stunden Vi-
deoaufzeichnungen mit dem Programm Interact der Firma Mangold codiert, mit
diesem und dem Programm Excel 2019 MSO von Microsoft (Version 2207) sta-
tistisch aufbereitet sowie nach Lebenswochen (03. – 07. LW und 08. – 15. LW)
oder Lebenstagen (50. – 105. LT) ausgewertet.
Die Auswertung des Verhaltes bei maximalem Tränkeanrecht (12 l MAT je Tier
und Tag bis zum 49. LT, n = 9 Kälber) erfolgte in den Funktionskreisen Ruhe-,
Nahrungs-, Wasseraufnahme- und Sozialverhalten (Kalb-Kalb-Kontakte) sowie
nach „sonstigen Aktivitäten“, welche Erkundungs-, Spiel-, Lokomotions- und Eli-
minationsverhalten umfassen. Erkundung (Belecken von Gegenständen) und
Spiel wurden codiert, Lokomotion (Stehen ohne Aktivität, langsames und schnel-
les Laufen, Springen, Galoppieren) und Elimination aus der Differenz der Dauer
der Aktivitäten und des in dieser Phase codierten Verhaltens berechnet und als
Lokomotionsverhalten zusammengefasst. Die Nutzung von Spielzeug, Besuche
Zusammenfassung 159
ohne MAT-Aufnahme und Saugaktivitäten (gegenseitiges Besaugen, Saugen an
Attrappen) wurden gesondert ausgewertet.
Aus den Untersuchungen kann ein Ethogramm für Kälber in der intensiven mut-
terlosen Aufzucht abgeleitet werden:
Tagesrhythmik
Kälber entwickeln eine ausgeprägte Tagesrhythmik, der nahezu das gesamte
Verhalten unterliegt. In der Nacht, d.
h. von Mitternacht bis etwa 06:00 Uhr, wech-
seln lange Ruhephasen mit kurzzeitigen Wachphasen zum Koten und Harnen
sowie Tränke-, Beifutter- und/oder Wasseraufnahmen. Am Tage, von 06:00 bis
24:00 Uhr, wechseln Aktivitäts- und Ruhephasen, die deutlich kürzer ausfallen
als in der Nacht.
Ruheverhalten
Zum Ruhen suchen die Kälber geschützte Plätze auf. Nach kurzer Unterbre-
chung legen sie sich häufig auf derselben Stelle wieder ab. Junge Kälber ruhen
täglich 14 bis 18 Stunden über durchschnittlich 30 bis 45 Minuten am Tage und
60 bis 180 Minuten in der Nacht. Vormittags und abends sind die Ruhezeiten
etwas kürzer als über Mittag.
Nahrungsaufnahmeverhalten
Das Saugverhalten an der Tränkestation entspricht dem natürlichen Saugen an
einer Kuh hinsichtlich Körperhaltung, Kopfstößen und den Schwanzbewegungen
sowie der mittleren Anzahl von vier bis fünf Mahlzeiten pro Tag. Die einzelnen
Mahlzeiten dauern durchschnittlich vier bis sechs Minuten mit steigender Ten-
denz.
Besuche an der Tränkestation ohne Tränkeaufnahme finden bis zum 49. LT kurz-
zeitig statt, im Durchschnitt dauern sie weniger als eine Minute. In der Woche der
Eingewöhnung an die freie Tränkeaufnahme sind bis zu vier dieser Blindbesuche
pro Tier und Tag tolerabel, danach bis zum Abtränken ein bis drei im Tagesmittel.
In der Abtränkphase können geringfügig längere Blindbesuche in einer steigen-
den Anzahl als normal angesehen werden, wenn sie einen Tagesmittelwert von
10 pro Tier nicht deutlich übersteigt.
Bis zum Ende der 07. Lebenswoche nehmen Kälber häufig über den Tag verteilt
mit nur kurzer Fressdauer Beifutter auf. Die Anzahl steigt bis auf etwa 20 Mahl-
zeiten pro Tag, die Dauer auf etwa drei Minuten pro Mahlzeit. Wasser wird von
Beginn an über jeweils etwa eine Minute aufgenommen, Frequenz und Dauer
steigen mit der Beifutteraufnahme.
Zusammenfassung 160
Einzelne Tiere haben ein ausgeprägtes Saugbedürfnis, das auch mit höchsten
Tränkeanrechten nicht über die Tränkeaufnahme befriedigt werden kann. Wenn
es sich um Einzelfälle handelt, kann das Besaugen eines anderen Kalbes toleriert
werden. Saugende Kälber aus einer mütterlichen Saugerfamilie sollten jedoch
von der Zucht ausgeschlossen werden. Saugattrappen werden von den Kälbern
angenommen, können das Besaugen aber nicht verhindern und werden weniger
genutzt als bewegliches Spielzeug.
Sozialverhalten
Kälber beriechen und belecken sich gegenseitig, spielen zusammen und ruhen
dicht beieinander. Rangauseinandersetzungen werden vor dem Absetzen nicht
beobachtet. Im Verhalten der jungen Kälber wurden keine Anzeichen gefunden,
dass die Kälber durch die fehlende Mutter-Kind-Beziehung Stress erleiden.
Lokomotions- und Spielverhalten
Kälber laufen, springen und galoppieren, oft zusammen und sich gegenseitig ani-
mierend. Besonders in den Abendstunden sind intensive Aktivitäten zu beobach-
ten. Das spielerische Hornen beginnt bereits ab der 02. Lebenswoche.
Als Spielzeug bevorzugen Kälber bewegliche Gegenstände, die sie zum Schwin-
gen bringen können. Diese werden beleckt, belutscht oder beschnuppert, oft von
mehreren Kälbern gleichzeitig. Lokomotion und Spielverhalten können zum
„sonstigen Verhalten“ zusammengefasst werden, um das Wohlbefinden von Käl-
bern zu beurteilen. Dessen Anteil an der aktiven Zeit sollte im Tagesdurchschnitt
bis zum Abtränken mindestens 80
% ausmachen. Beim Abtränken verringert sich
der Anteil wegen der zunehmenden Beifutteraufnahme, sollte aber 60
% der Ak-
tivitäten nicht unterschreiten.
Abtränken
Ein moderates Abtränken ist aus der Sicht des Tierverhaltens zu empfehlen, da-
mit die Tiere lange ruhen, nur wenige Fehlversuche an der Tränkestation auftre-
ten und eine hohe Beifutteraufnahme beim Absetzen gesichert wird. Da das ma-
ximale Tränkeanrecht bis zum 49. Lebenstag angeboten werden muss, ergibt
sich ein zu empfehlendes Absetzalter von mindestens 105 Tagen.
Acknowledgements 161
Acknowledgements
The studies on the behaviour of female Holstein calves during rearing were car-
ried out in collaboration with an agricultural enterprise in Mecklenburg-Vor-
pommern.
We thank the management of the business for their support of the project and
providing data. The employees and students from the University always had ac-
cess to the calf village and were comprehensively supported on site with data
collection. During and even outside regular working hours, the stable gates were
open for us, and we were provided access to the calves and the technical equip-
ment. We give sincere thanks for the trust placed in us.
A special thank you goes to the herd manager, Sandra Dorn. Your interest and
extensive support made the studies and the preparation of this publication possi-
ble. The construction of the ball as a calf toy and its installation in the pen moti-
vated us to observe and analyse the play behaviour of the calves and to test the
chains as another play element.
Cameras recorded the behaviour of the calves over months. We thank the calf
rearers for agreeing to allow us film them while working.
Our student and research assistant Kajetan Baltrock reliably managed the data
recording on the study farm and thus made a fundamental contribution to the
success of this project. We therefore give him our special thanks.
Birgit Pfeiffenberger of the Animal Nutrition and Feed Science unit edited and
formatted this publication in her usual best manner. We are very grateful for her
knowledgeable and reliable work and the extensive critical and highly construc-
tive feedback.
The project was supported by the National and State Female Professors Pro-
gramme II and received financial support from additional sponsors. We are grate-
ful for the funding provided by:
Hauptgenossenschaft Nord AG, Kiel
Holm & Laue, Westerrönfeld
Professor Dr Anke Schuldt and Dr Regina Dinse
Neubrandenburg, June 2023
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Attachment of tables 193
Attachment of tables
Table A 1: Calf, date of birth and round as well as number of days and hours
evaluated ..................................................................................................... 195
Table A 2: Average, minimum, maximum and standard deviation of parameters
of resting behaviour up to the 49th day of life .............................................. 196
Table A 3: Average, minimum, maximum and standard deviation of parameters
of resting behaviour during weaning from the 50th to the 105th day of life .. 197
Table A 4: Average, minimum, maximum and standard deviation of parameters
of CMR-feed-intake up to the 49th day of life .............................................. 198
Table A 5: Average, minimum, maximum and standard deviation of parameters
of solid-feed-intakes up to the 49th day of life ............................................. 199
Table A 6: Average, minimum, maximum and standard deviation of parameters
of Water-intakes up to the 49th day of life ................................................... 200
Table A 7: Average, minimum, maximum and standard deviation of parameters
of visits at the dispensing station without CMR-feed-intake (blind visits) up to
the 49th day of life ....................................................................................... 201
Table A 8: Average, minimum, maximum and standard deviation of parameters
of ingestion during weaning from the 50th to the 105th day of life ............... 202
Table A 9: Average, minimum, maximum and standard deviation of parameters
of solid-feed-intakes during weaning from the 50th to the 105th day of life . 203
Table A 10: Percentage of solid-feed-intake by duration of intake during weaning
from the 50th to the 105th day of life, n = 5 calves ...................................... 203
Table A 11: Average, minimum, maximum and standard deviation of parameters
of water-intakes during weaning from the 50th to the 105th day of life ........ 204
Table A 12: Average, minimum, maximum and standard deviation of parameters
of visits at the dispensing station without CMR-feed-intake (blind visits) during
weaning from the 50th to the 105th day of life ............................................. 204
Table A 13: Average, minimum, maximum and standard deviation of parameters
of calf-calf-contacts up to the 49th day of life ............................................... 205
Table A 14: Average, minimum, maximum and standard deviation of parameters
of calf-calf-contacts during weaning from the 50th to the 105th day of life .. 205
Table A 15: Average, minimum, maximum and standard deviation of parameters
of locomotion and exploring behaviour (licking of objects) up to the 49th day of
life ................................................................................................................ 206
Attachment of tables 194
Table A 16: Average, minimum, maximum and standard deviation of parameters
of games up to the 49th day of life .............................................................. 206
Table A 17: Average, minimum, maximum and standard deviation of parameters
of games by type of toy as well head butts up to the 49th day of life .......... 207
Table A 18: Percentage of games with toys as well head butts by calves up to the
49th day of life ............................................................................................. 208
Table A 19: Average, minimum, maximum and standard deviation of parameters
of games during weaning from the 50th to the 105th day of life .................. 208
Table A 20: Average, minimum, maximum and standard deviation of parameters
of sucking sessions activities by weeks of life in the 06th to 15th week of life, n
= 13 examined calves ................................................................................. 209
Table A 21: Average, minimum, maximum and standard deviation of parameters
of sucking sessions by calves in the 5th and 15th week of life, n = 13 examined
calves .......................................................................................................... 209
Table A 22: Average, minimum, maximum and standard deviation of parameters
of sucking sessions at calf feeder, percentage at night as well visits at
dispensing station up to 5 minutes before sucking in the 5th and 15th week of
life ............................................................................................................... 210
Table A 23: Average, minimum, maximum and standard deviation of parameters
of sucking sessions at suction dummy next by dispensing station, percentage
at night as well visits at dispensing station up to 5 minutes before sucking in
the 5th and 6th week of life ......................................................................... 210
Table A 24: Average, minimum, maximum and standard deviation of parameters
of sucking sessions at suction dummy at gate, percentage at night as well visits
at dispensing station up to 5 minutes before sucking in the 5th and 6th week of
life ............................................................................................................... 211
Attachment of tables 195
Table A 1: Calf, date of birth and round as well as number of days and hours
evaluated
Calf Date of birth Round Number of evaluated
Days Hours
1 20/11/17 1 6 144.0
2 19/11/17 1 17 323.6
3 29/11/17 1 33 669.4
4 04/01/18 2 26 552.5
5 02/02/18 2 43 889.2
6 19/04/18 3 13 311.0
7 22/04/18 3 16 374.7
8 24/04/18 3 14 318.9
9 24/04/18 3 31 740.5
10 30/07/18 4 14 328.7
11 08/08/18 4 19 396.3
12 13/09/18 5 16 371.3
13 29/09/18 5 43 1,006.3
Total 5212 4,569.5
Attachment of tables 196
Table A 2: Average, minimum, maximum and standard deviation of parameters
of resting behaviour up to the 49th day of life
Week c n Average min max s
Resting behaviour, total during in hours per animal and day
3 1 5 14.6 12.6 16.0 1.3
4 4 11 15.1 8.9 18.4 3.4
5 9 33 16.2 6.7 19.0 8.3
6 9 31 15.9 8.8 21.6 8.3
7 9 31 16.5 7.2 19.2 8.4
Mean duration of rest periods in minutes
3 1 104 44.6 1.0 286.5 47.3
4 4 256 46.1 1.1 367.9 52.8
5 9 786 41.9 1.0 263.1 41.0
6 9 640 46.2 1.0 295.0 47.3
7 9 636 49.8 1.1 290.9 46.4
Total
Average
2,422
45.3
1.0 367.9
45.7
Mean duration of rest periods in minutes, 00:00 to 06:00 a.m.
3 1 26 70.0 1.7 249.4 65.9
4 4 27 109.7 2.0 367.9 117.8
5 9 160 65.4 1.2 234.5 55.7
6 9 119 79.8 1.1 295.0 69.4
7 9 131 76.0 1.8 290.9 61.0
Mean duration of rest periods in minutes, 06:00 to 24:00 a.m.
1 22 33.3 1.0 286.5 39.2
4 4 230 35.2 1.1 158.5 31.1
5 9 627 35.0 1.0 263.1 33.3
6 9 521 37.9 1.0 214.4 36.7
7 9 505 42.2 1.0 254.9 38.9
Week = week of life, c = number of calves, n = number of records, min = minimum, max
= maximum, s = standard deviation
Attachment of tables 197
Table A 3: Average, minimum, maximum and standard deviation of parameters
of resting behaviour during weaning from the 50th to the 105th day of life
Week c n Average min max s
Resting behaviou
r
. total during in hours per animal and day
8 3 10 16.3 10.1 18.2 2.4
9 4 11 14.5 4.8 18.3 4.0
10 4 7 14.3 11.1 16.5 2.5
11 3 9 16.1 8.9 19.1 2.8
12 3 9 17.6 16.0 19.6 1.1
13 2 11 16.1 13.9 17.7 1.2
14 4 10 14.9 9.0 17.5 2.5
15 2 7 14.6 14.0 16.8 1.1
Mean duration of rest sessions in minutes
8 3 173 56.6 1.2 310.4 52.6
9 5 271 61.6 1.7 369.4 57.7
10 5 253 52.7 1.4 289.9 48.6
11 4 184 50.2 1.1 297.4 48.6
12 3 197 58.0 1.6 451.0 56.1
13 2 158 67.4 3.9 324.0 61.4
14 4 229 57.1 1.1 307.2 48.6
15 2 134 48.3 1.9 194.6 36.5
Total
Average
1,599
56.7
1.1 451.0
52.3
Mean duration of rest sessions in minutes, 00:00:00 to 06:00:00 a.m.
8 3 42 77.9 3.9 203.2 61.3
9 5 63 92.6 2.4 369.4 85.3
10 5 66 84.2 3.1 289.9 65.3
11 4 39 84.3 1.1 297.4 71.3
12 3 43 93.9 2.0 451.0 87.7
13 2 29 128.1 16.1 324.0 91.3
14 4 52 90.0 3.3 307.2 69.2
15 2 34 66.0 1.9 136.2 36.8
Mean duration of a rest period in minutes, 06:00:00 a.m. to 12:00:00 p.m.
8 3 131 49.8 1.2 310.4 47.7
9 5 208 52.2 1.7 242.2 42.3
10 5 187 41.6 1.4 166.5 35.1
11 4 145 41.0 1.9 153.4 35.5
12 3 154 48.0 1.6 222.6 37.7
13 2 127 51.7 3.9 164.1 35.2
14 4 177 47.5 1.1 173.9 36.9
15 2 100 41.2 1.9 136.2 36.8
Week = week of life, c = number of calves, n = number of records, min = minimum.
max = maximum, s = standard deviation
significance: mean number and mean duration of rest periods up to the 49th day of life
(23.0 in 45.3 minutes) vs. from the 50th to the 105th day of life (17.2 in 56.7 minutes).
p < 0.001
Attachment of tables 198
Table A 4: Average, minimum, maximum and standard deviation of parameters
of CMR-feed-intake up to the 49th day of life
Week c n Average min max s
Mean duration of CMR-feed-intakes in minutes
3 1 26 4.3 0.6 9.7 2.7
4 4 68 4.1 0.3 10.9 1.8
5 9 198 4.4 0.5 14.6 2.2
6 9 174 4.7 0.6 11.8 2.3
7 9 192 4.9 0.4 12.8 2.4
Total
Average
658
4.6
0.3 14.6
2.3
Mean duration of CMR-feed-intakes in minutes, 00:00 to 06:00 a.m.
3 1 3 6.9 4.6 9.7 2.6
4 4 6 5.6 3.8 6.6 2.0
5 9 28 4.3 0.9 8.0 1.8
6 9 22 4.7 0.9 9.0 2.3
7 9 40 4.9 0.4 11.8 2.6
Mean duration of CMR-feed-intakes in minutes, 06:00 a.m. to 12:00 p.m.
3 1 23 3.9 0.6 8.5 2.5
4 4 62 4.0 0.3 10.9 1.8
5 9 170 4.3 0.5 14.6 2.2
6 9 152 4.7 0.6 11.8 2.2
7 9 152 4.9 0.6 12.8 2.4
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation, CMR = calf milk replace
r
Attachment of tables 199
Table A 5: Average, minimum, maximum and standard deviation of parameters
of solid-feed-intakes up to the 49th day of life
Week c n Average min max s
Mean duration of solid-feed-intakes in minutes
3 1 114 1.5 0.1 6.4 1,3
4 4 154 1.9 0.1 8.7 1,5
5 9 501 2.1 0.1 17.3 1,9
6 9 488 2.6 0.2 26.2 2,2
7 9 597 3.1 0.1 20.5 2,8
Total
Average
1,854
2,5
0.1 26.2
2.3
Mean duration of solid-feed-intakes in minutes, 00:00 to 06:00 a.m.
3 1 12 2,2 0,4 4,7 1,1
4 4 9 1,8 0,2 3,3 1,0
5 9 45 1,8 0,1 6,5 1,3
6 9 36 2,1 0,2 6,3 1,1
7 9 75 2,8 0,1 13,2 2,3
Mean duration of solid-feed-intakes in minutes, 06:00 a.m. to 12:00 p.m.
3 1 102 1,4 0,1 6,4 1,3
4 4 145 1,9 0,1 8,7 1,5
5 9 456 2,1 0,1 17,3 1,9
6 9 450 2,5 0,2 26,2 2,2
7 9 522 3,1 0,1 20,5 2,9
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation
Attachment of tables 200
Table A 6: Average, minimum, maximum and standard deviation of parameters
of Water-intakes up to the 49th day of life
Week c n Average min max s
Mean duration of water-intakes in minutes
3 1 46 1.2 0.1 3.9 1.0
4 4 31 0.9 0.2 2.2 0.5
5 9 128 0.8 0.1 8.6 0.9
6 9 143 1.0 0.1 3.9 0.8
7 9 164 1.2 0.03 8.4 1.1
Mean duration of water-intakes in minutes, 00:00 to 06:00 a.m.
3 1 1 0.4 0.4 0.4 -
4 4 2 1.0 0.7 1.2 0.4
5 9 9 1.8 0.2 8.6 2.7
6 9 14 0.8 0.1 2.5 0.7
7 9 20 1.6 0.2 8.4 1.9
Mean duration of water-intakes in minutes, 06:00 a.m. to 12:00 p.m.
3 1 45 1.2 0.1 3.9 1.0
4 4 9 0.9 0.2 2.2 0.5
5 9 119 0.8 0.1 2.8 0.6
6 9 129 1.0 0.1 3.9 0.8
7 9 144 1.1 0.03 5.7 0.9
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation
Attachment of tables 201
Table A 7: Average, minimum, maximum and standard deviation of parameters
of visits at the dispensing station without CMR-feed-intake (blind visits) up to the
49th day of life
Week c n Average min max s
Mean duration of blind visits in minutes
3 1 25 0.9 0.04 3.1 0.9
4 4 24 0.9 0.1 4.7 1.0
5 9 95 0.9 0.1 3.6 0.8
6 9 112 0.7 0.03 4.1 0.6
7 9 59 1.0 0.1 6.5 1.1
Mean duration of blind visits in minutes, 00:00 to 06:00 a.m.
3 1 0 - - - -
4 4 4 0.4 0.2 0.7 0.2
5 9 3 0.7 0.3 1.1 0.4
6 9 14 1.2 0.1 4.1 1.1
7 9 13 0.7 0.1 1.8 0.5
Mean duration of blind visits in minutes, 06:00 a.m. to 12:00 p.m.
3 1 25 0.9 0.04 3.1 0.9
4 4 20 1.0 0.07 4.7 1.1
5 9 92 0.9 0.07 3.6 0.8
6 9 98 0.7 0.03 2.0 0.4
7 9 46 1.1 0.06 6.5 1.2
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation, CMR = calf milk replacer
Attachment of tables 202
Table A 8: Average, minimum, maximum and standard deviation of parameters
of ingestion during weaning from the 50th to the 105th day of life
Week c n Average min max s
Mean duration of CMR-feed-intakes in minutes
8 3 50 4.6 1.6 12.8 2.1
9 5 63 6.3 1.3 12.5 2.7
10 5 53 6.3 1.4 11.5 2.4
11 4 43 4.8 1.3 11.2 2.9
12 3 35 4.0 0.1 26.7 4.3
13 2 36 4.2 1.7 8.1 1.5
14 4 37 3.5 1.5 6.6 1.4
15 2 25 4.8 1.9 8.4 1.6
Total
Average
342
4.9
0.1 26.7
2.7
Mean duration of solid-feed-intakes in minutes
8 3 160 1.1 0.07 31.4 2.8
9 5 367 1.4 0.05 36.4 3.4
10 5 363 1.3 0.16 39.7 3.1
11 4 251 1.4 0.25 24.2 2.9
12 3 222 1.5 0.11 59.7 4.0
13 2 248 2.2 0.23 37.7 5.0
14 4 338 2.0 0.27 41.2 5.2
15 2 113 0.8 0.12 15.4 1.8
Total
Average
1,062
5.0
0.05 59.7
5.5
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation, CMR = calf milk replacer
Significance: mean duration of CMR-feed-intakes in minutes up to the 49th day of life
(4,6 minutes) vs. during weaning from the 50th to the 105th day of life (4,9 minutes),
p = 0,04; mean duration of solid-feed-intakes in minutes up to the 49th day of life (2,5
minutes) vs. during weaning from the 50th to the 105th day of life (5,0 minutes), p <
0,001
Attachment of tables 203
Table A 9: Average, minimum, maximum and standard deviation of parameters
of solid-feed-intakes during weaning from the 50th to the 105th day of life
Week c n Average min max s
Mean total duration of solid-feed-intakes in hours per animal and day
8 3 10 1.1 0.5 1.8 0.5
9 5 19 1.4 0.2 2.6 0.7
10 5 19 1.3 0.04 2.7 0.8
11 4 10 1.8 1.1 3.0 0.6
12 3 11 1.7 0.3 3.3 0.8
13 2 11 2.7 1.2 3.7 0.8
14 4 16 2.4 0.5 4.1 1.4
15 2 7 0.8 0.2 1.4 0.5
Mean duration of hay-intakes in hours per animal and day
8 3 10 0.6 0.1 1.3 0.4
9 5 18 0.7 0.1 1.9 0.4
10 5 18 0.6 0.03 1.3 0.4
11 4 10 0.8 0.4 1.7 0.4
12 3 11 0.7 0.2 1.0 0.2
13 2 11 0.5 0.1 1.0 0.3
14 4 16 0.7 0.1 2.9 0.7
15 2 6 0.5 0.1 1.0 0.3
Mean duration of total-mixed-ration-intakes in hours per animal and day
8 3 10 0.5 0.2 1.3 0.3
9 5 18 0.8 0.05 2.0 0.5
10 5 18 0.7 0.04 2.0 0.5
11 4 10 1.0 0.5 1.3 0.3
12 3 11 1.0 0.1 2.6 0.7
13 2 11 2.2 1.0 3.1 0.7
14 4 16 1.6 0.2 3.6 1.1
15 2 7 0.4 0.2 0.6 0.2
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation
Table A 10: Percentage of solid-feed-intake by duration of intake during weaning
from the 50th to the 105th day of life, n = 5 calves
Week of life 8 9 10 11 12 13 14 15
Duration Percentage of solid-feed-intake by duration of intake
< 3 minutes 51% 54% 54% 48% 49% 33% 39% 67%
3 to 5 minutes 16% 18% 21% 22% 21% 21% 17% 20%
5 to 10 minutes 28% 19% 17% 24% 19% 24% 24% 9%
> 10 minutes 6% 9% 7% 6% 11% 23% 21% 4%
Attachment of tables 204
Table A 11: Average, minimum, maximum and standard deviation of parameters
of water-intakes during weaning from the 50th to the 105th day of life
Week c n Average min max s
Mean total duration of water-intakes in minutes per animal and day
8 3 8 2.4 0.8 5.2 1.5
9 5 18 5.1 0.2 12.3 3.7
10 5 17 5.7 1.0 16.3 4.0
11 4 10 10.0 0.5 18.4 6.4
12 3 9 12.7 5.5 20.3 4.7
13 2 11 10.9 4.8 23.2 5.7
14 4 16 11.0 1.8 22.5 6.9
15 2 2 7.5 4.9 9.8 3.6
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation
Table A 12: Average, minimum, maximum and standard deviation of parameters
of visits at the dispensing station without CMR-feed-intake (blind visits) during
weaning from the 50th to the 105th day of life
Week c n Average min max s
Mean duration of blind visits in minutes
8 3 20 0.9 0.02 4.1 1.0
9 5 36 1.5 0.08 4.1 1.0
10 5 54 1.2 0.06 5.1 1.1
11 4 80 1.1 0.16 5.6 0.8
12 3 72 0.9 0.05 3.0 0.6
13 2 130 1.2 0.08 7.5 0.9
14 4 124 1.1 0.10 3.8 0.7
15 2 37 1.2 0.04 3.5 0.9
Mean number of blind visits per animal and day
8 3 8 2.5 1.0 4.0 1.1
9 5 11 3.1 1.0 6.0 1.8
10 5 10 4.9 2.0 15.0 4.1
11 4 10 8.0 3.0 15.0 3.5
12 3 9 8.0 1.0 19.0 5.4
13 2 11 11.8 3.0 24.0 5.7
14 4 16 7.8 1.0 21.0 5.6
15 2 7 5.3 2.0 10.0 2.6
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation, CMR = calf milk replacer
Attachment of tables 205
Table A 13: Average, minimum, maximum and standard deviation of parameters
of calf-calf-contacts up to the 49th day of life
Week c n Average min max s
Mean duration of calf-calf-contacts in minutes
3 1 48 2.0 0.02 16.4 3.3
4 4 86 2.0 0.02 12.8 2.7
5 9 175 1.3 0.04 7.3 1.3
6 9 121 1.5 0.18 7.4 1.3
7 9 127 2.0 0.13 15.7 2.4
Mean duration of calf-calf-contacts in minutes, 00:00 to 06:00 a.m.
3 1 2 0.4 0.41 0.4 0
4 4 3 4.7 1.19 10.2 4.8
5 9 16 0.9 0.04 1.7 0.5
6 9 11 1.4 0.36 3.3 4.2
7 9 13 3.1 0.74 15.7 4.2
Mean duration of calf-calf-contacts in minutes, 06:00 a.m. to 12:00 p.m.
3 1 46 2.1 0.02 16.4 3.3
4 4 83 1.9 0.02 12.8 2.6
5 9 159 1.3 0.08 7.3 1.4
6 9 119 1.6 0.18 7.4 1.3
7 9 114 1.8 0.13 11.7 1.9
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation
Table A 14: Average, minimum, maximum and standard deviation of parameters
of calf-calf-contacts during weaning from the 50th to the 105th day of life
Week c n Average min max s
Mean duration of calf-calf-contacts in minutes
8 3 42 2.1 0.28 9.0 2.0
9 5 130 1.5 0.04 9.4 1.6
10 5 84 1.3 0.09 7.9 1.3
11 4 24 1.7 0.34 7.4 7.8
12 3 27 1.5 0.13 10.2 1.9
13 2 11 1.1 0.48 2.4 0.6
14 4 57 1.4 0.21 8.7 1.4
15 2 21 2.0 0.31 7.0 1.7
Week = week of life, c = number of calves, n = number of records, min = minimum, max
= maximum, s = standard deviation
Attachment of tables 206
Table A 15: Average, minimum, maximum and standard deviation of parameters
of locomotion and exploring behaviour (licking of objects) up to the 49th day of
life
Week c n Average min max s
Active time, total duration in hours per calf and day
3 1 5 9.4 8.0 11.4 1.3
4 4 11 8.9 5.6 15.1 3.4
5 9 3 7.8 5.0 17.3 3.2
6 9 31 8.1 2.4 15.2 3.2
7 9 31 7.5 4.8 16.8 2.9
Locomotion time total, duration in hours per calf and day
3 1 5 8.0 7.4 10.9 1.4
4 4 12 7.7 4.8 17.9 4.4
5 9 33 6.5 4.8 17.3 3.0
6 9 31 6.5 2.2 14.7 3.0
7 9 31 5.7 4.6 16.2 2.6
Licking of objects total, duration in minutes per calf and day
3 1 6 11.7 6.4 15.5 3.7
4 3 7 15.5 4.5 30.2 11.0
5 2 5 11.4 2.1 39.1 15.6
6 1 2 6.5 5.4 7.7 1.6
7 5 10 9.3 1.3 21.7 7.2
Week = week of life, c = number of calves, n = number of records, min = minimum, max
= maximum, s = standard deviation
Table A 16: Average, minimum, maximum and standard deviation of parameters
of games up to the 49th day of life
Week c n Average min max s
Mean total duration of games with ball, brush, chains and head butts, in minutes per calf
and day
4 4 14 9.2 0.6 23.0 6.7
5 9 31 7.6 0.5 24.0 6.1
6 9 26 12.6 0.2 36.1 8.4
7 9 28 11.5 0.6 36.1 9.1
Mean duration of games with ball, brush, chains and head butts in minutes
4 4 105 1.2 0.1 5.2 0.6
5 9 217 1.1 0.1 7.0 0.4
6 9 244 1.4 0.1 9.1 0.6
7 9 226 1.4 0.04 7.9 0.6
Week = week of life, c = number of calves, n = number of records, min = minimum, max
= maximum, s = standard deviation
Attachment of tables 207
Table A 17: Average, minimum, maximum and standard deviation of parameters
of games by type of toy as well head butts up to the 49th day of life
Week c n Average min max s
Mean duration of games with ball in minutes
4 4 70 1.3 0,1 5,2 1,2
5 9 97 1.2 0,1 7,0 1,1
6 9 91 1.4 0,2 5,5 1,1
7 9 110 1.5 0,04 7,9 1,5
Mean duration of games with brush in minutes
4 4 5 1.0 0,4 1,4 0,4
5 9 53 0.8 0,1 2,8 0,6
6 9 44 1.3 0,1 4,0 1,0
7 9 60 1.4 0,2 6,7 1,3
Mean duration of games with chains in minutes
4 4 21 0.7 0,3 1,4 0,3
5 9 40 1.0 0,2 2,6 0,6
6 9 82 1.2 0,2 9,1 1,2
7 9 28 1.2 0,1 3,3 0,8
Mean duration of head butts in minutes
4 4 9 2.0 0,3 3,7 1,4
5 9 27 1.3 0,1 3,9 0,9
6 9 27 1.5 0,2 5,3 1,3
7 9 28 1.3 0,2 6,1 1,4
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation
Attachment of tables 208
Table A 18: Percentage of games with toys as well head butts by calves up to the
49th day of life
Calf / Round Percentage of games
Ball Brush Chains Head butts
2 / 1 91
% 0
% 0
% 9
%
3 / 1 91
% 0
% 0
% 9
%
4 / 1 86
% 10
% 0
% 4
%
5 / 2 63
% 1
% 33
% 3
%
6 / 3 23
% 21
% 41
% 15
%
7 / 3 - - - 100
%
8 / 3 34
% 25
% 26
% 15
%
9 / 3 41
% 27
% 21
% 12
%
10 / 4 35
% 28
% 28
% 9
%
11 / 5 50
% 9
% 27
% 14
%
12 / 5 38
% 15
% 26
% 21
%
13 / 5 34
% 36
% 23
% 7
%
Total 52.9
% 18.5
% 18.3
% 10.3
%
Table A 19: Average, minimum, maximum and standard deviation of parameters
of games during weaning from the 50th to the 105th day of life
Week c n Average min max s
Mean duration of games in minutes per calf and day
8 3 70 1.4 0.3 5.0 0.6
9 4 144 1.7 0.1 9.2 0.8
10 4 165 1.8 0.02 10.2 0.9
11 3 105 1.8 0.2 11.8 1.0
12 3 74 1.4 0.1 10.2 0.6
13 2 110 2.0 0.2 18.5 1.2
14 3 112 1.5 0.2 9.1 0.6
15 2 37 1.3 0.2 5.8 0.5
Mean number of games per calf and day
8 3 10 7.0 3 11 2.2
9 4 12 6.7 2 14 4.5
10 4 13 8.8 1 19 6.3
11 3 9 9.1 1 14 4.6
12 3 10 7.4 3 14 4.1
13 2 11 10.0 3 20 5.8
14 3 15 7.5 1 13 2.9
15 2 7 5.1 3 7 1.7
Week = week of life, c = number of calves, n = number of records, min = minimum,
max = maximum, s = standard deviation
Attachment of tables 209
Table A 20: Average, minimum, maximum and standard deviation of parameters
of sucking sessions activities by weeks of life in the 06th to 15th week of life, n =
13 examined calves
Week C / S
Number of sucking sessions Duration of sucking sessions
n /
night d max/
d mean min max s
6 9 / 1 1 / 0 1 1 9.7
7 9 / 1 3 / 0 3 1 3.1 2.1 4.3 1.1
9 4 / 1 2 / 0 1 2 1.1 0.9 1.2 0.2
11 3 / 1 1 / 0 1 1 1.4
13 2 / 1 3 / 0 3 2 1.3 0.9 1.6 0.3
14 4 / 1 1 / 1 1 1 1.9
15 2 / 1 8 / 1 4 5 2.6 0.6 4.5 1.6
Week = week of life, C = number of examined calves, S = number of suckers,
n = number of records, night = at night, d = number of days, min = minimum,
max = maximum, s = standard deviation
Table A 21: Average, minimum, maximum and standard deviation of parameters
of sucking sessions by calves in the 5th and 15th week of life, n = 13 examined
calves
Calf Round
Number of sucking sessions Duration of sucking sessions
n /
night d max/
d Average min max s
1 1 9 / 2 5 5 2.5 0.6 4.5 1.5
3 1 2 1 2 1.1 0.9 1.2 0.2
5 2 4 3 2 1.3 0.9 1.6 0.3
9 3 4 4 1 4.7 2.1 9.7 3.4
n = number of records, night = at night, max = maximum, d = number of days,
min = minimum, s = standard deviation
Attachment of tables 210
Table A 22: Average, minimum, maximum and standard deviation of parameters
of sucking sessions at calf feeder, percentage at night as well visits at dispensing
station up to 5 minutes before sucking in the 5th and 15th week of life
Calf /
Round
Suction sessions
Number Durationr
n d max
/
d
Mean duration perc.
CMR
/
BV
before sucking
Average min max s
4/ 1 40 10 10 1.4 0.04 8.1 1.5 9 / 21 %
5 / 2 31 5 12 1.3 0.1 6.6 1.3 20 / 8 %
6 / 2 31 7 8 1.4 0.2 3.5 0.8 35 / 4 %
7 / 3 11 3 8 1.3 0.6 2.9 0.8 0 / 11 %
8 / 3 20 5 10 1.5 0.03 12.4 2.7 28 / 28 %
9 / 3 21 5 10 1.0 0.3 3.7 0.7 17 / 17 %
Total 154 35 12 1.3 0.03 12.4 1.4 22 / 15 %
n = sucking sessions, d = days, max = maximum, CMR
/
BV = CMR-feed-intake
/
blind
visits, before sucking = up to 5 minutes before sucking,
min = minimum, s = standard deviation, perc. = percentage
Table A 23: Average, minimum, maximum and standard deviation of parameters
of sucking sessions at suction dummy next by dispensing station, percentage at
night as well visits at dispensing station up to 5 minutes before sucking in the 5th
and 6th week of life
Calf /
Round
Suction sessions
Number Duration
n d max
/
d
Mean duration
perc.
at night
perc.
CMR
/
BV
before
sucking
Average min max s
11 / 4 3 2 2 0.7 0.6 0.9 0.1 18 % 20 / 20 %
12 / 5 5 4 3 0.9 0.3 2.4 0.9 0
% 0 / 25 %
13 / 5 8 6 2 0.6 0.2 1.2 0.3 12
% 25 / 0%
Total 16 12 3 0.7 0.2 2.4 0.5 8.3 % 23 / 23 %
n = sucking sessions, d = days, max = maximum, at night = 00:00 - 06:00 a.m., CMR
/
BV =
CMR-feed-intake
/
blind visits, before sucking = up to 5 minutes before sucking,
min = minimum, s = standard deviation, perc. = percentage
Attachment of tables 211
Table A 24: Average, minimum, maximum and standard deviation of parameters
of sucking sessions at suction dummy at gate, percentage at night as well visits
at dispensing station up to 5 minutes before sucking in the 5th and 6th week of
life
Calf /
Round
Suction sessions
Number Duration
n d max
/
d
Mean duration
perc.
at night
perc.
CMR
/
BV
before
sucking
Average min max s
10 / 4 2 1 2 0.6 0.6 0.7 0.1 46
% 0 / 0 %
11 / 4 3 3 1 1.3 0.4 2.6 1.2 0
% 0 / 0 %
13 / 5 8 7 2 1.2 0.2 2.9 1.0 31.7
% 0 / 9 %
Total 13 11 2 1.1 0.2 2.9 0.9 19.0
% 0 / 9 %
n = sucking sessions, d = days, max = maximum, at night = 00:00 - 06:00 a.m.,
CMR
/
BV = CMR-feed-intake
/
blind visits, before sucking = up to 5 minutes before
sucking, min = minimum, s = standard deviation, perc. = percentage
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336 Seiten, 91 Abbildungen, 74 Tafeln
ISBN 3-932227-53-0 12,50 €
Bd. 19 Fechner, J., Fock, T.: Analyse des landwirtschaftlichen Fachkräfte- und Bildungs-
bedarfs im Land Brandenburg. 2002
115 Seiten, 24 Abbildungen, 71 Tabellen
ISBN 3-932227-54-9 8,00 €
Bd. 20 Autorengruppe des durch die Deutsche Bundesstiftung Umwelt (DBU) geförderte
Forschungsvorhabens: DEICHRÜCKBAU UND SALZGRASLAND AN DER SÜD-
LICHEN OSTSEEKÜSTE. 2003
219 Seiten, 17 Abbildungen, 54 Tabellen, 5 Karten
ISBN 3-932227-55-7 12,50 €
Bd. 21 Lührs, H., Spitzner, P.: Freiraumplanung und Pflege des Denkmals Klosterberg
Broda. Neubrandenburg, 2004
129 Seiten, 51 Abbildungen, 42 Fotos
ISBN 3-932227-64-6 8,00 €
Bd. 22 Fock, T., Schwertassek, S.: Die Stellung der Landwirtschaft in der Region Meck-
lenburgische Seenplatte. 2005
98 Seiten, 9 Abbildungen, 43 Tabellen
ISBN 3-932227-66-2 7,50 €
Bd. 23 Dehne, P. (Hrsg.): Gedanken zur Entwicklung ländlicher Räume in Mecklenburg-
Vorpommern. 2006
156 Seiten, 22 Abbildungen, 7 Tabellen
ISBN 3-932227-73-5 8,00 €
Bd. 24 Fock, T., Tillack, J.: Landwirtschaftliche Kleinerzeugung in der Region Mecklen-
burgische Seenplatte. 2006
108 Seiten, 3 Abbildungen, 44 Tabellen
ISBN 3-932227-75-1 8,00 €
Bd. 25 Hirte, K., Walter, J.: Handlungsstrategien und Werte – zwischen Leitbildern und
Leistungsvorstellungen im ökologischen Landbau in Ostdeutschland. 2006
120 Seiten, 28 Diagramme
ISBN 3-932227-76-X 7,50 €
Bd. 26 Behrens, H., Kaether, J., Stöckmann, M.: Freiwillig Selbstkontrolle Nachhaltigkeit
(FSK-N) in Region und Stadt). 2006
239 Seiten, 53 Abbildungen, 17 Tabellen
ISBN 3-932227-77-6 10,00 €
Innerhalb der Schriftenreihe erscheinen noch:
Reihe A – Fachbereich Agrarwirtschaft und Landschaftsarchitektur
Reihe B – Fachbereich Bauingenieur- und Vermessungswesen
Reihe C – Fachbereich Soziale Arbeit, Bildung und Erziehung
Reihe D – Fachbereich Technologie
Reihe E – Interdisziplinäre Schriften
Reihe F – Allgemeine Schriften
Reihe G – Fachbereich Gesundheit, Pflege, Management
Reihe H – Multimedia
Reihe J – Fachbereich Landschaftswissenschaften und Geomatik (neu seit Mai 2013)