THE EFFECT OF VARYING THE DAILY MILKING
FREQUENCY ON THE MILK YIELD OF THE EWE
AND EVIDENCE ON THE NATURE OF THE INHIBITION
OF MILK EJECTION BY HALF-UDDER MILKING
M. MORAG
Negev Institute
for
Arid Zone Research, Beersheva(Israel)
SOMMAIRE
Par deux
expériences,
on a déterminé l’effet de différentesfréquences
de traitejournalière
surla
quantité
de lait obtenue chez la Brebis.Expérience
1. - Des demi-mamelles sont traites r,z ou 3fois parjour, selon
unplan expérimental
en carré latin 3 X
3 (tabl. 2 ).
Le groupe bleu, de 6 brebis(tabl. i)
est trait normalement ; pour le groupe blanc(6 animaux ), la
traite est suivie de deuxinjections d’ocytocine ( 5
UI X2 )
pour extraire le lait résiduel.Il
n’y
a pas de différence entre les deux groupes pour lesquantités
de lait et de matières grasses,quand
les demi-mamelles sont traites 2et 3fois parjour (tabl.
3, 4, 5,6),
alorsqu’une
seule traite parjour
provoque une réduction d’environ 20p. 100.Le groupe trait avec
ocytocine
a uneproduction augmentée
d’environ 30 p. iooquelle que
soitla
fréquence
des traites(tabl.
5, 6).La
quantité
de laitproduite
par une demi-mamelle est influencée par lafréquence
des traites de l’autreglande
chez un même animal. Moins on trait l’une des mamelles,plus grande
est laquantité produite
par l’autre.Expérience
2. - Sur deux groupes de 4brebis(tabl.
7) on compare la traite 2et 3fois parjour, les
mamelles étant
comparées
soit entre mamelles entières, soit entre demi-mamelles, dans unprotocole
en carré latin 4 X
4 (tabl. 8).
Pour les
quantités
de lait, lesgraisses,
la matière sèche, iln’y
a pas de différence entre les semi- mamelles traites 2 ou 3fois parjour (tabl.
9). Deplus,
laquantité
de laitproduite
par une demi-ma-melle, pour une
fréquence
de traite donnée( 2
ou 3), estcomparable
à celle obtenue du côté opposétrait 2 ou 3fois par
jour. Cependant,
on observe uneproduction
réduite d’environ 20p. 100quand
unedemi-mamelle est traite seule
après
un intervalle de temps donné, par rapport à laquantité
obtenuede la même demi-mamelle
quand
on la trait en même temps que l’autre moitié etaprès
le même inter-valle de temps
(effet
de lacoïncidence) (tabl. i 5 ).
Il estpossible
que cette réduction observéequand
lesdeux demi-mamelles ne sont pas traites ensemble soit due à l’inhibition de
l’éjection
du lait. Cettehypothèse
semble confirmée par le fait que si oninjecte
del’ocytocine,
lesquantités
de lait obtenues sont les mêmes, que les demi-glandes soient traites simultanément ou non.INTRODUCTION
Evidence on the effect of
varying
thefrequency
ofdaily milking
is derived from both survey andexperimental
data.H ANSSON and BONNIER, ( 1947 )
havepointed
outthat in the former the results are biased in favour of the more
frequent milking
rou-tines. The latter
reports
are derived from whole lactation or short term studies em-ploying
variousexperimental techniques :
monozygoustwins,
and whole and half uddermilking
routines(see
reviewby E LLIOT T, 195 8).
In very
general
terms one can summarize these data based on cattle trials asshewing
that when thefrequency
ofdaily milking
is increased from twice tothrice,
there is asubsequent
andusually
immediate response in milkyield.
The increases quo- ted in variousreports
vary from6-!o
p. IOO. Onesuspects
that the increase becomes smaller as theexperimental design
becomes better. There isgeneral agreement
thatonce-a-day milking
ofdairy
cows causes a reduction of milkyield
of some 50p. 100.There is
only
onereport
on the effect of achange
inmilking frequency
on theyield
ofsheep (G AA i< 1957 ).
Ewesyielded
20 p. 100 more milk when milked thrice.This
experiment
was carried out under commercial conditions and it may well be that the results are biased in favour of the group milked thrice. In view of lack of informa- tion on thissubject
in ewes, it was decided to carry out thefollowing investigations.
EXPERIMENT I Materials and Methods
A first
experiment
into the effect of differentmilking frequencies
was carried out on twelve mut-ton breed ewes. The
sheep,
all in the first month of lactation, had beenseparated
from their lambs within six hours ofparturition.
All the ewes had suckled lambsduring
theprevious
year and none had been milkedduring
any earlier lactation. Details of theexperimental
animals aregiven
in table i. Thetwelve ewes were ranked in order of
previous yield
and then divided intopairs
in the rank order. Theewes of each
pair
were thenrandomly
allocated to one of two groups, hereafter referred to as the blue and white groups. Both groups were milked twice aday
for sevendays
and then six ewes,again
selec-ted at random, were allocated to the treatment sequences of a balanced 3 X
3 Latin
squaredesigna-
ted
Capital,
and theremaining
six ewes to the sequences of a second balanced Latin square(Cursive),
as shewn in table z. It is noted that the treatment definitions of A and a, B and b C and c
respectively,
were similar for whole
sheep
and differedonly
in respect of the half udder to which treatments wereapplied.
Eachperiod
lastedeight days
- thereadings
of the firstday
of eachperiod being
discardedin an attempt to eliminate the carry-over effects of residual milk from treatment to treatment. Total
experimental
time was 32days. made up as follows :(
I
pre-treatment X 7days)
)-( 3
treatmentperiods
Xy
discard + treatmentdays])
= 32days.
The ewes were
machine
milked usingequipment
described elsewhere(M ORAG ,
GIBB and Fox, 196 7
). The
sheep
werebrought
into theparlour
formilking
and offered concentrates in themilking
stands. The teat cups were then
applied
and when the milk flow had ceased the udders of the white group of ewes werevigorously massaged
and then machine-, andhand-stripped.
In the blue group residual milk was removedby
twoinjections
of 5i. u.oxytocin ( 1 )
which werepainlessly
administeredthrough
asemi-permanent nylon
cannula which had been inserted in thejugular
vein fivedays prior
to the
experiment.
Udderwashing
andfore-milking
were notpractised.
Themilking
times were o5.00, i3
.oo and 21.00h, at og.oo and 21.00h and at 21.00h for the thrice-, twice- and
once-a-day milking
routines. All half udder; were milked sst z r. oo h wich was ths only half udder coincidental mil-
king during
the threeperiods
of the Latin square. Udder half-milk samples were taken at every milkingand
analysed
for fatby
a modified method of Gerber as describedby
MACDONALD( 1959 ).
(Fat sam-ples
were not taken for the first fourdays
of thepre-experimental week).
The ewes were housed on slats in a closed barn illuminated
by day
andnight.
The ewes wereoffered water, grassnuts and hay ad lib. between
milkings
and were offered 3kg
of concentrates( 2 )
per
day
milking. Thisdaily
ration of concentrates was divided into a number of mealsequal
to thenumber of
milkings
which the ewe received on any oneday.
( 1
) P. 0. P. Purified oxytocin principle-manufactured by Armours Pharmaceutical Co. Ltd.
( 2
) Soya 20p. ioo, fish meal 20p. 100, rolled barley 60 p. 100, with 5 kg. bone flour and 3kg. A !- D33 pre-mix per ton.
The blood of the ewes in the blue group was
effectively anti-coagulated by
the intravenous admi- nistration of 700 !L9. ofMalayan viper.
Ancistyodon yhodosto!na( BOIE )
venom,given intravenously through
the cannula every thirdday.
The
experiment
was carried out at Lane End Farm,University
ofReading,
fromApril
ioth toMay
x4th 1964.Results
The
weekly yields
of left andright
udder halves of milk and fat in the blue and white groups wereanalyzed separately according
to thefollowing
model:When [1. = the mean
daily yield
in a treatmentperiod
Hs = the effect due to the ith half udder i = I, 2, 3 P
j = the effect due to the
jth period j =
1, 2, 3Tx = the effect due to the kth treatment k = 1, 2, 3
The half udder treatment means for the two latin squares and the two groups with the relevant standard errors are
given
in tables 3 and 4. The milkyields
of theudder halves
during
thepre-experimental
week aregiven
in table i. It is noted that in tables 3and the
means for the threemilking frequencies
arequalified by
the mil-king frequency
on the other half udder of the ewe. The mean udderyields
of milk andfat in the blue and white groups,
regardless
of themilking frequency
on the other sideof the
udder,
have been calculated and appear in table 5. The effect of themilking frequency
on theopposite
side of the ewe isgiven
in table 6. No standard errors aregiven
in tables 5 and6 ;
this is because the treatment means arecomposed
of estimates derived from different latin squares and thus nomeaningful
standard error could becalculated.
Discussion
The
experimental
resultspresented
shew that there is no difference between theyield
of milk or fat when a ewe is milked twice or three times aday. (see
table 5 andf
i g. z).
This would appear to be true whether or not the residual milk was removed atmilking (compare
the means in tables 3 and4 ).
None of the communications ofcomparisons
between twice and thricedaily milkings, including
a trial carried out on alarge
number of ewes(GnAr&dquo; ig 57 ), reported
that the milkyield
did not increase inresponse to the extra
daily milking. Furthermore,
thedrop
in milkyield following
achange
to asingle daily milking
- 24p. ioo and i8 p. 100in the blue and white groupsrespectively
- was far smaller than the 50p. Ioo that one wouldexpect
on the basis of cattleexperiments. (C LAESSON , H ANSSON ,
GUSTAFSSONand BxaNNArrG,I959)!
The
experiment provided
a further demonstration of thegalactopoeitic
effect ofthe
oxytocin
removal of residual milk in the ewe as describedby
D!rrAMUR and iVInx-TINET
,
10 6 1
and MopAG andFox, 19 66. During
the three weeks of the latin square the blue group(oxytocin) yielded
29p. 100more milk and 32 p. 100 more fat than did the white group(without oxytocin).
The effect on the
yield
of a half udder of themilking frequency
on the other side is seen in table 6 andfigure
2. It woutdappear
that the lessfrequently
theopposite
sideis milked the
higher
theyield
becomes. Apossible explanation
tothis,
could be that it is due to some obscure intra-uddercompensatory
mechanism such as had beensuggested
in other contexts in cattle
by
GARRISON and TURNER( 193 6) and by GRIFFIN (i 9 66).
In view of the
inability
to demonstrate theexpected
response to a thirddaily
mil-king
and in order toclarify
the effect ofapplication
of differentmilking frequencies
tothe half udders of the same ewe a further
experiment
was initiated.EXPERIMENT 2 Materials and Methods
The effect of twice and
thrice-a-day milking
estimatedby
half and full uddertechniques
wasinvestigated using
8 mutton breed ewes (see table7 ). The
8 ewes were ranked in order of theirprevious yield
and were then divided in the rank order into twoyield
blocks of 4ewes each. The ewes of each block were thenrandomly
allocated to the treatment sequences of a ¢ X 4Latin square (see table8).
Each
period
lasted 7days
and thereadings
of the firstday
of eachperiod
were discarded in an at- tempt to eliminate carry-over effects of residual milk. Totalexperimental times
was 28days
made up :Milking
was carried out in theparlour
described above. Ewes not scheduled to be milked at anyparticular
hour werekept
in theyard
out of earshot of thepulsators
until themilking
session wasover. The ewes to be milked were
brought
into theparlour
and offered concentrates in themilking
stands. The teat cups were then
applied
and when the milk flow had ceased the udders werevigo- rously massaged
and then machine-, andhand-stripped.
Udderwashing
andfore-milking
were notpractised. Milking
times were at 0.800, 16.oo and 24.oo and at 12.oo and 24.oo h on the thrice andtwice-a-day milking
routinesrespectively.
The fat and total solids content ofdaily composite
sampleswere carried out
by
the modified method of Gerber(MnCDOrrnLn, ig 5 g)
andby
the method of GOL- DING
(i 934 ), respectively.
Feeding, housing
andgeneral
management were as described in theprevious experiment.
Results and Discussion
The average
daily
half-udderyields
for eachperiod
ofmilk, fat,
total solids and solids-not-fat wereanalyzed according
to thefollowing
model :when [1. = the mean
daily yield
of a half udder in aperiod
Si = the effect due to the ith. ewe i =i, 2...
8Hss = the effect due to the
jth.
half udder of the ith.ewe j =
1,2 Pt = the kth.period
k = I, 2, 3, 4Tim = the effect due to the Ith. treatment of two or three times
a-day
mil-king
and the mth. treatment of two or threetimes-a-day milking
on the otherside of the udder I = 1, 2 and m = 1, 2.
The means and relevant standard errors are
given
in tables 9 and 10.During
thehandling
of the field data it was noticed that themidnight yield
were oftenhigher
than the
day
timeyields.
The differences between theyield
atmidnight
and the othermilkings,
corrected for the differences in intervallengths (see
footnote table9 ),
weretherefore also
subjected
toanalysis according
to the above model.(The
fatyields
could not be
analyzed
in this way as the fat content had been determined indaily
com-posites).
The data of this
experiment
confirm thefindings
of theprevious experiment
andshewed that a
change
from two to threedaily milkings
had no effect on theyield
ofmilk in the ewe.
Furthermore,
there was asuggestion (significant only
at a level ofio p.
100 )
that a non coincidentalmilking
routine had caused a reduction in milkyield.
Only
the waterphase
wasactually
affected as theyield
of total solids shewed no such trend(see
interactions means for milk and total solids in table9 ).
The reduction inyield
at a non-coincidentalmilking
ascompared
to a coincidental washighly signifi-
cant
(see
interaction means for the difference betweenmilkings
atmidnight
and atother times in table
9 ).
Examination of the data from individual ewes shewed that the
slope
of the lac- tation curve was a characteristic of each ewe and that the decline wasindependent
oftreatment. This is shewn in table ii. and
figure
3 which compare theslope
of thedaily yield
within eachperiod
with that betweenperiods
in individual ewes. The linearrelationship
of theplots
in thefigure
illustrates thispoint clearly.
Now the model used for theanalysis
takes out bias due to time as a« period
o effect(with
3degrees
of free-dom)
and is based on theassumption
that the decline inyield
with the advance oflactation is uniform in all
subjects.
As this was not so in thesedata, they
were reana-lyzed using
thefollowing
linearmodel; (for general
method seeCox, 195 8, 19 6 2 ) :
when bi is the
regression
ofyield
of time in the ith. ewe(other
nomenclature as inthe previous model).
This model takes out the bias as an individual ewe «
slope
effect with 8degrees
offreedom. The decline in
yield
with time for the 16 half udders hadpreviously
beenplotted
out, and it was seen that theslopes
of the decline of the two half udders in anyone ewe were
similar,
and for this reason theregressions
were calculated on a ewe ra-ther than a half udder basis.
The covariance means and the relevant standard errors are shewn in table 12. It can be seen that the
similarity
between theyields
to twice- andthrice-a-day
mil-king
remains as in theprevious analyses.
The reduction in milk(that
is of the waterphase)
on thestaggered milking
routineshas, however, disappeared.
This indicates that the reduction shewn in the earlieranalysis
was dueentirely
to aspurious
latinsquare effect and was not, in
fact,
ofbiological origin.
An additional set of
6 4 analyses
were carried out on the differences inyield
bet-ween the
midnight
anddaytime yield
of milk in each half udderperiod, according
tothe
following
model :!t = the mean difference between the
midnight
and thedaytime yield
in anudder half.
Da = the effect due to ith.
day
i = I, 2, ... 6T
j
= the effent due tojth.
time ofday j
= 1, 2.The bias in favour of the
midnight milking
wassignificant (P
<0 . 01 )
in 32cases,that is in all cases that the
daytime milkings
were vcon-coincidental andnon-significant
in the 32 cases zvhen both
day
andnight milkings
were coincidental.As half udder
techniques
have beenwidely
used in cowstudies,
a check was madeto see if a similar effect was
produced by
the non-coincidentalmilking
of half uddersin cattle. Data taken from an earlier half-udder
experiment by
Er,rro’r2( 195 8)
wereinvestigated
and a similar effect to that shewn in thepresent work,
albeit of lessmagni- tude,
was observed.(see
l!IoRnG,ig66).
A
suggested explanation for
theeffect of
non-coincidenceof milking
based on a
reanalysis of
the dataof Experiment
7The reduction in half udder milk
yield
at a non coincidentalmilking
couldpossi- bly
be due to an inhibition ofejection
for reasons which arequite
obscure. It can beargued
that if this was so there should be no reduction when half udders are milkednon-coincidentally
with the aid of an exogenousoxytocin.
To test thishypothesis
theoriginal
data ofExperiment
I wereanalyzed
for differences between theyield
ofmilk at 2r.oo h and the other
milkings
in the blue and the white groups.Analysis
ofthe first week’s results gave the difference between two coincidental
milkings,
and any bias thatappeared
canonly
be attributed to some diurnal effect. Theanalysis
carriedout on the differences
during
theperiods
of the latin squares(i.
e.during
the2 nd, 3
rd
and4 th experimental weeks)
deals with the above biasplus
the effect of non-coincidence of
milking. (All yields
of milkduring periods
in which half udders weremilked
only once-a-day have,
of course, been discarded from theseanalyses).
Table i4 shews that in both the blue and white groups there was a
significant
bias in milk
yield
in favour of themorning milking, (the
difference in fatyield
wassig-
nificant
only
in the blueoxytocin group) during
the firstweek,
when allmilking
was coincidental. This is in
general agreement
with thehypothesis
ofS EM j AN ( 19 6 2 )
that the rate of milk secretion in the ewe is
higher during
thenight.
Oneis, however,
unable to discern any diurnal effect of the mechanism ofejection,
as didSemjans
since the differences in
yield
between theevening
andmorning milkings
in both the blue and white groups appear to be similar.Table i5,
however, presents
a differentpicture. During
theperiods
of the latin square there was asignificant
bias in theyields
of milk and fat in favour of the eve-ning
i. e. coincidentalmilking.
The reduction inyield
at a non-coincidentalmilking
discerned in
Experiment
2is thus alsoclearly
demonstratedby
this furtheranalysis
of data from the first
experiment.
These data do notonly
confirm thisfinding
but alsopresent strong
evidence that the reduction inyield
at a non-coincidentalmilking
is at leastpartially
due to an inhibitionof
milkejection.
This can be deduced since the bias in favour of a coincidentalmilking, although
stillsignificant,
becamefar
smaller whenoxytocin
wasgiven
at allmilkings.
This is seenby comparing
thegrand
means in theblue
(oxytocin)
and the white(without oxytocin)
groups(table z 4 ).
The two
experiments
described in this paperconsistently
shew that there is noincrease in milk
yield
when ewes are milked thricedaily
either with or without the aid ofoxytocin.
It would appear that this is animportant species characteristic,
whenone compares the lack of response in the ewe with the many
reports
of an increase in milkyield
from a thirddaily milking
derived from cow studies. An additionalspecies
characteristic is the
relatively
small reduction in milkyield
causedby
achange
fromtwo to a
single daily milking,
which was far smaller than thatreported
in cows.Re f
u pour
publication
enjanvier
1968.SUMMARY
Two
experiments
were carried out to determine the effect of differentdaily milking frequencies
on the milk
yield
of the ewe.In the first
experiment
ewe half-udders were milked once, twice or thricedaily, using
a 3 X 3 Latin squaredesign.
One group of six ewes was milked as in normalpractice
whilst another group 6 ewes was milked with the aid ofoxytocin.
Within each group the milk and fatyields
of udder halves milked twice and thrice were identical, whilst asingle daily milking
caused a reduction of some 20p. ioo. The group milked with the aid ofoxytocin out-yielded
the other ewesby
over 3o p. 100on all
milking frequencies.
Theyield
of agiven
udder half was shewn to be influencedby
the mil-king frequency
on the other udder half : the lessfrequently
the other side was milked the greater theyield
of thegiven gland.
The second
experiment compared
twice and thricedaily milking
on a whole and a half udder ba-sis in
eight
ewesusing
a 4 X 4Latin squaredesign.
Theyields
of milk, fat and total solids of half ud- ders milked thrice wasagain
found to be identical to that of half udders milked twice. Furthermore theyield
of an udder half on agiven milking frequency
were similar when theopposite
side was milkedtwice or thrice
daily.
A reduction of over 20p. 100inyield
was however, detected when a half-udderwas milked alone after any
given
interval(non-coincidentally)
ascompared
to theyield
of the sameudder half when milked
together
with toopposite gland (coincidentally)
after the same interval. Itwas
suggested
that this reduction inyield,
when agland
was milkednoncoincidentally,
was due toan inhibition of milk
ejection. Support
for this was found in that whenoxytocin
wasgiven
at bothcoincidental and non-coincidental
milkings
theyields
were similar.ACKNOWLEDGEMENTS
This work was undertaken with a grant from the Miriam Sacher Charitable Trust. The author
was in
receipt
of theHuntley
and PalmerResearchship.
Theoxytocin
wasspecially prepared
anddonated
by
Mr. W. F. TICEHURSTof Armours Pharmaceuticals Co. Ltd. The statisticalanalyses
werecarried out
by
Dr.J.
L. BROOKHOUSEand Miss F. N. NuTT. The chemicalanalysis
was doneby
Dr. BALCH.
Help
with themilking
wasgiven by
Mr. C. S. SIMMOMS, Mr. M. FRANKS and Miss E. BROA- DBENT. I record my
appreciation
to all concerned. I am likewise indebted to Dr. D. ROBERSTON SMITH of theRoyal
BerkshireHospital,
to Dr.J.
MARTINET and Mr. S. Fox for their comments and to Mr. PARAN,lately Agricultural
Counsellor at the IsraeliEmbassy,
London, for his kind interest in this work.REFERENCES C
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, !., HANSSON A., GUSTAFSSON 5., BRANNANG E., 1959 Studies on monozygous cattle twins.
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Cox C. P., ig58. The analysis of latin squares with individual curvatures in one direction. Jl. R. stalist.
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ENAMUR R., MARTINET J., ig6i. Action de l’oxytocine sur la secretion du lait de brebis. Ann. Endocr., 22
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OTT G. M., 958. The effect of milk accumulation in the udder of the dairy cow upon milk secretion rate its bearing on certain milking practices. Ph. D. Thesis. University of Reading.
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L M., 1967. Milking ewes three times daily on a large farm in Hungary (in Hungarian).Allattanyyesztes, Budapest, 6, 61-64.
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OLDING J., r934. The rise of the air-damped balance for the determination of total solids in milk. Analyst Lond., 59, 468-74.
GRIFFIN T. K., 1966. Personal communication.
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ANSSON A., BONNIER G., 1947. Studies on monozygous cattle VIII. Amount and composition of the
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D F. J., r959! A neutral reagent for the routine determination of fat in milk and milk production.
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RAG M., 1966. A reduction in milk yield with certain half-udder milking techniques. J. Dairy Res., 33, 223-226.
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EM]AN S., 1962. Residual milk of sheep. XVI. Int. Dairy Congr., 1, I7-24·