Responses of dairy
ewesbefore and after parturition
to
different nutritional regimes during pregnancy
II.
-Energy intake, body-weight changes during lactation
and milk production
D. STERN J. H. ADLER H. TAGARI E. EYAL I nstitute of Animal Science, Agricultural Research Ovganizatio!z,
The Volcani Center, Bet Dagan, Israel (
*
) Dept. of Physiology, Hadassah-Hebrew University Medical School, Jea!xssalena, Israel (
**
) Dept. of Animal Sciences, Faculty of Agriculture,
The Hebrew University of Jerusalem, Rehovot, Israel
Summary
Some effects of nutritional levels during pregnancy on subsequent lactation were studied
in dairy ewes. Three basic (maintenance) rations and three steaming-up levels were compared
in a factorial experiment. All ewes were fed 0.4 kg of hay daily and pellets ad Libitum. Ewes
entering late pregnancy in heavy body condition due to a high basic ration consumed zi p. 100 less feed in early lactation than those that had received the lower rations. Reduced appetite
was more marked in single- than in twin-bearing ewes.
Steaming-up levels did not have any effect on ad lib. feed consumption during lactation.
Twin-bearing ewes consumed, on the average, 9.6 p. ioo more feed than single-bearing ones.
There was a negative correlation between body weight change during lactation and weight change during pregnancy.
Overfatness of ewes was followed by low milk yields. In ewes which had been on constant
steaming-up levels during the last two months of pregnancy, milk production was significantly higher in twin- than with single-bearing ewes. There was no such difference in the ewes whose feed allowance during pregnancy had been gradually raised according to the free fatty acidconcen- tration in their blood plasma.
There was no correlation between feed intake and body weight change during lactation
on the one hand, and body weight change and milk production on the other hand. There were
highly significant correlations between feed intake and milk yield.
Fat percentage in milk was not affected by feeding level during pregnancy. Average fat
level was 6.7 7-0.2 p. ioo and 7.5 J: 0.2 p. 100 in the milk of twin- and single-bearing ewes,
respectively.
Introduction
The effect of nutrition
during gestation
on theembryo’s development
andon the
well-being
of the ewe and of the newborn lamb has been studied in detail and describedby
numerous authors(e.g., R EID , r 9 6 3 ;
GORDON andTRIBE,
Ig5r;DO N A L
D and Russ!!&dquo; zg7o; STERN et
al., ig 7 8).
Some recent work indicates thatfeeding
levelsduring
pregnancy may also affect thesubsequent
lactation of theewe
(B U TTE RWORTA
and BLORE,19 6 9 ;
Gaxnvnx andHOGUE, ig63;
PEART,1967;
TREACHER,
1970,1971). Furthermore,
there seems to exist an interaction betweenbody
condition(degree
offatness)
of the eweduring
differentstages
ofgestation
and the level of extra feed allowance
during
late pregnancycommonly
referredto as &dquo;
steaming-up
&dquo;(P EART
, 19 68,
1970; andunpublished
results of our labo-ratory).
Allpublished
literature on thissubject
deals with milkproduction
asa
major
factoraffecting
lambproduction.
To the best of ourknowledge,
thereare no
reports
ondairy sheep.
Thepresent
communication ispart
of astudy
toclarify
the interaction ofbody
condition andfeeding
levelduring
theperiod
bet-ween
conception
and thesubsequent parturition,
insheep
which arekept
forhigh
levels of
production
of both lambs and milk. A detaileddescription
of theexperi-
ment which was carried out, and of the effects of the various
feeding regimes
onlamb
production,
is included in aprevious
communication(S TERN et al., 197 8).
Materials and methods
General
Forty-two
Awassi and Assaf ewes were block randomized into nine treatment groups(3 X 3 factorial).
Theexperimental
treatments included three sequences of basic dietsthroughout
pregnancy:i) flushing
andhigh
maintenance(fh); 2 ) flushing
and low maintenance(fn); 3 )
noflushing
and low maintenance(nn);
andthree levels of
steaming-up: high (H),
low(L),
andadjusted according
to bloodplasma
freefatty
acid concentration(A).
Each ewe waskept
in theexperiment
45
days post-partum.
Theexperiment
was described in detailby
STERN et al.( 197
8). Only
detailspertaining specifically
to the lactationperiod
will be des- cribed below.Feeds and
feeding
Ewes were maintained in individual pens. From
parturition
onwards eachewe received o.4
kg hay daily
and concentratepellets containing
r4.5 p. 100 fiber and 15.5 p. 100 crudeprotein
ad libitum in individual self-feeders. Self-feeder balance was done every week and from this average,daily consumption
was cal-culated. Feed
composition
wasgiven by
STERN et al.( 197 8).
Weighing of
e¡cesEwes were
weighed
on themorning
of the thirdday !ost-partum
and at fort-nightly
intervals thereafter.Suckling o/
lambs and milkrecording
Each ewe suckled
only
one lamb. Litter mates from twin ortriplet
birthswere taken away
4 8
hpost-partum
for artificialrearing.
From the fourthday
on,the
remaining
lamb wasput
with its mother for r5 minutesonly
after themorning
and
evening milkings
to suck whatever residual milk there was in the udder. Milkyields
were recorded everyfortnight.
Onrecording days
lambs wereweighed before
and aftersuckling
to estimate thesuckling yield (SuY);
totalyield (TY)
was estimated
by adding
SuY to the milked ordairy yield (DY).
See MoRAGet al.
( 1973 )
for detailed definitions of milk fractions. Percentdry
matter(DM)
in milk was determined
by drying
in an oven at io5°C; percent
fatby
the Gerbermethod;
andpercent
solids not fat(SNF) by subtracting
fatpercent
from DM.Aliquots
of themorning
andevening milkings
were combined foranalysis
to re-present
thecomposition
of the DI’.Determinations of the
composition
of the suckled milk were done twiceduring
the
¢ 5 -day suckling period
for each ewe. To obtain the residualmilk,
five inter- national units ofoxytocin
wereinjected
into thejugular
veinimmediately
aftermilking.
Residual milk was thenmilked, weighed,
andsampled
foranalysis.
To calculate the energy value of
milk,
it was assumed that each gram of fat wasequivalent
to 9.4 and each gram of SNF to 4.9 calories. Combined fat or SNFyields
of thedairy plus
the suckled milk were used to calculate thepercent
of these fractions of the total milkyield
and thus the total milk energy.Results
Feed intake
Prepartum
nutritional level affected feed intakeduring
lactation(Table I ).
The reduction in
appetite
as a result of thehigh
basic level(fh) (ewes entering
latepregnancy in fat
condition)
wasparticularly
marked. Loss ofappetite
was moresevere in
single-bearing
than intwin-bearing
ewes. The effect of the&dquo; steaming-
up &dquo;
regimes
on feed intakeduring
lactation was small andstatistically
notsigni-
ficant.
The fact that in
twin-bearing
ewes intake wasgreater (by
an average of9 .6
p.ioo)
than insingle-bearing
ewes isnoteworthy.
This difference between the ewesof the two birth
types
was smallest in theadjusted steaming-up
groups(A).
Allthese differences
persisted
over the first 6 weeks oflactation,
when intake increasedcontinuously
in all the groups.Body weight changes
There was a noticeable difference at
mating
and inearly
pregnancy betweenbody weights
of twin- andsingle-bearing
ewes. The former were heavier than the latter(see
STERN etal., rg 7 8).
This differencedisappeared
almostcompletely during
the last two months of pregnancy.At
lambing, twin-bearing
ewes(who
either lost more orgained
lessweight)
were heavier
by only
3 p. 100, on the average, thansingle-bearing
ewes.On the other
hand, significant
differences inweight
remained between eweswhich had been on different basic rations. The same holds true for ewes on differ- ent
steaming-up
levels.During
the first 6 weeks of lactation(when
ewes were fed adLibitum)
therewas a
negative relationship
betweenprogressing body weight changes during
thisperiod
and those associated withadvancing gestation.
The correlation coefficients(r)
were - 0.727(P
<o.oi)
for ewes fed the constantsteaming-up regimes (H
and
L)
and —0 . 3 6 9 (NS)
for theadjusted
group(A). Fig.
i shows the relativebody weight changes
of ewesduring
the first six weeks of lactation(weight
atmating
= 100 p.100 ).
Thefollowing
facts were evident:a)
The smallestchange
frommating
tolambing
occurred in theadjusted
group. Differences between twin- and
single-bearing
ewes were also smallest in this group.b)
There was a marked difference in the course ofweight change during early
lactation between twin- and
single-bearing
ewes. Twinners lostweight
orgained only
alittle; singlers
lost little orgained
at anoticeably greater
rate. There wasno correlation between
body weight changes
and feed intake.Milk
yields
Average daily
milkyields
in the first six weeks of lactation forsheep
whichhad received different rations
during
pregnancy are shown in Table 2.It seems evident that
overfattening during gestation (basic
levelfh)
led toreduced milk
yield, especially
insingle-bearing
ewes.Also,
the milkyield
of twin-bearing
ewes on constant&dquo; steaming-up &dquo; regimes (H
andL)
washigher
than thatof
singles. Furthermore,
ewes on theadjusted regime
showed no suchdifference,
and there was even atendency
toward ahigher
milkyield
insingle-bearing
ewes.Figure
2 shows thatyields
increased until the4 th
week of lactation and then stabilized orbegan
to decrease.In ewes on constant
steaming-up regimes,
the difference between twinners andsinglers
was maintainedthroughout
the first 6 weeks of lactation. In theadjusted steaming-up feeding (A),
atendency
was found for twin-bearers’ milkyield
to rise moresteeply,
butby
the 6th week both groups hadequal yields.
No correlation was found between
body weight changes during
lactationand milk
yield.
The correlations between milkyield
and feed intake weresigni-
ficant at each of the three
points
that were testedduring
the first 6 weeks of lacta- tion. The correlation coefficients are shown in Table 3.Milk
fat percentages
The effect of
prepartum
nutritional levels on thepercentage
of milkfat wassmall
(Table 4 ), yet
differences found between varioussteaming-up regimes
weresignificant (P
<0 . 05 ).
It appears that low(L) level, especially
in the case offat ewes
(basic
levelfh),
led to arelatively
lowpercentage
of milk fat.The most
prominent
differences wereagain
observed between twin- andsingle- bearing
ewes, with the fat content of the latter’s milkbeing consistently higher.
It is
interesting
to note that this state exists notonly
in the constant&dquo; steaming-
up
&dquo;
regimes (H, L),
where the milkyield
ofsingle-bearing
ewes waslower,
butalso in ewes on the
adjusted regime (A),
where the milkyield
ofsingle-bearing
ewes was
high.
In
figure
3, it can be seen that the differences betweensingle.
andtwin-bearing
ewes were maintained
during
the first 6 weeks of lac tation. Within eachregime,
there was an inverse relation of milk
yield
to fatpercentage.
Total milk energy
The
higher
level of milk fat insingle-bearing
ewes was notenough
to make up for their milkyield
incomparison
with thetwin-bearing
ewes on constantsteaming-
up
regimes
H and L, when milkyield
wasexpressed
as energy(Table 5 ).
In theadjusted steaming-up regime (A),
there was ahigh percent
of milk fat insingle- bearing
ewesalong
with ahigher
milkyield, compared
withtwin-bearing
ewes.Therefore,
the differences between ewes of the two birthtypes
in thisregime
areeven
greater
whenexpressed
in energy than whenexpressed
in milkyield.
The differences between energy
yield
of ewes on the various&dquo; steaming-up
&dquo;regimes
were found to besignificant (not
so for the milkyield expressed
inkg).
The differences arise
mainly
from thesingle-bearing
ewes.Single-bearing
eweswhich were on the
adjusted feeding
beforelambing produced
both more SNF andmore fat than those with a constant
steaming-up (P
<0 . 05 ).
Intwin-bearing
ewes an
opposite
trend wasnoted,
but differences were small andstatistically
notsignificant.
Discussion
Effect of !ye!aytum
nutyition onfeed
iyatalzeduring
lactationThe
&dquo; steaming-up &dquo; regimes
hadonly
a small effect on feed intakeduring
lactation.
Actually,
aslightly higher
intakecompared
with that of othersteaming-
up groups was noted in
single-bearing
ewes in theadjusted (A) feeding regime
alone.Even
there,
thepositive
effect was limited tosheep
on thehigh
basic levels(fh, fn). Sheep entering
thesteaming-up period
in a fat condition suffered frominap- petance
towards the end ofgestation (S TERN et al., 197 8)
and it seems that thiseffect was carried over into the lactation.
Contradictory
results exist in the literature on thissubject.
TREACHER( 1970 )
found that the nutritional level in late pregnancy had no influence on feed intake
during
lactation. PEART(ig6 7 ),
on the otherhand,
did find such an effect. It is worthnoting
that in PEART’S work the ewes’ ration in late pregnancy was ad-justed according
to FFA concentration in the blood. It may besuggested
thatincreased
feeding
at a constant levelduring
the late months of pregnancy isapt
to increase
appetite
inearly lactation,
whengiven
to lean ewes. For an averagepopulation,
agradually
increased diet may beadvantageous. Overfattening
would
probably
induceinappetance.
Relatiovcship
betweenfeeding levels, body weight changes
and milhyield High
correlation coefficients were found between feed intake and milkyield,
as in other studies
(PE AR T, 19 6 7 ; F ORB E S , ig6g; T R E A C H ER, 1970). However,
there was no correlation between feed intake and
body weight change during lactation,
nor betweenbody weight change
and milkyield.
Feed intake wasinfluenced to a certain
degree by
the ewes’body
condition at the end of pregnancy.Thus,
lean ewes consumed more than those inheavy
condition under the ad libitumfeeding
in thisstudy. They
consumed well in excess of their maintenanceplus
lactation
requirements.
This resulted in an inverse relation between rate ofbody weight change during gestation
and thatduring
lactation. An identical resultwas recorded
by
TREACHER(rg!o).
The maintenance levels andsteaming-up
levels alike were found to have
virtually
no effect on feed intake and milkyield
in the
subsequent
lactation oftwin-bearing
ewes. On the otherhand, they clearly
affected
single bearers,
which wereseverely
affectedby
thehigh
maintenance level(obesity)
butparticularly
benefited from the FFAadjusted feeding (t1).
It isnoteworthy
that the latterfeeding regime
alsoproduced
the heaviest lambs.Twin-bearing
ewes consumed more food!ost-pavtum
andput
on moreweight
than
singlers.
In the constant(both high
andlow) steaming-up regimes,
twinnersproduced
more milk as well. This canperhaps
beexplained by
the fact that whenon a constant
pre-partum
nutritionallevel, twin-bearing
ewes come much closerto a state of energy
deficiency
than do thesingle-bearing
ewes,by
the end of ges- tation. PEART( 19 68)
showed that lean ewes(in
thepresent
case,twin-bearers)
utilized their food more
efficiently
for theproduction
of milk than did fat ewes.It
is, however, possible
that the very hormonal differencesresponsible
for differentovulation rates at
mating
also affect milkyield.
The literaturesupports
theconjecture
that these same hormones act onfertility
and milkproduction (see,
for
example, HUNTER, 19 68).
The difference in feed intake cannot be related tobody weight differences,
since the latterdisappeared
almostcompletely
betweenmating
andlambing.
Thetwin-bearing
ewes were on the average 3 p. 100heavier thansingle
bearers(not significant). However,
the average difference in feed intake was of the order of 10p. 100(p
<o.oi).
The
only comparisons
in the literature between thetype
of birth and milkyields
indairy sheep
were those of GooT(ig66, Ig 74 ).
He found differencesclosely resembling
those describedherein, although
of a much smallermagnitude.
Many
researchers found ahigh
level of milkproduction
intwin-bearing
ewes(WAI,I,AC!, zgq8; BARNICOAT,
LOGAN andGRANT,
1949;Ai,Ex.KNDER
andDAVIES,
1959;HADJIPIERIs
andHO LMES , 19 66;
PEART,19 6 7 ),
but none of these studiesare
comparable
to ours, sincethey
dealt with ewes which bore and also suckledtwins,
and were not milked at all. In thisstudy, twin-bearing
ewes suckledonly
one lamb and the
suckling
consistedonly
of milk left over aftermilking.
For theadjusted feeding
groups(A)
the difference inbody
condition and energy balance in late pregnancy between twin- andsingle-bearing
ewes was very small.However,
also
here,
trends in feed intake andbody weight changes
resembled those in otherregimes.
On the otherhand,
the milkyield
ofsingle-bearing
ewes washigher
thanor
comparable
with that of twinners. This indicatesprimarily
that theadjusted feeding (A)
led tohigher efficiency
notonly
in lambproduction (STE RN et al., rg 7 8),
but also in milk
production.
Inaddition,
it seems thattwin-bearers,
for the samereasons,
perhaps,
thatthey produced relatively
small lambs(STERN
etal., 197 8)
were also unable to attain
high
levels of milkproduction.
Under the
present conditions,
when initialfeeding
was adlibitum,
there wasno benefit from the reserves built up in the ewe’s
body
from thehigh steaming-up regime (H). However,
the situation may be different in the case of a rationeddiet,
where atemporary
energy deficit is liable to appear, at least inearly
lactation.Support
for thehypothesis
that ewes may benefit from abuild-up
ofbody
reservesin pregnancy may be found in the
reports
of PEART( 1970 )
and TpEACHER(1971),
who found that ewes fed more in late pregnancy were able to maintain a
higher
milk
yield
over a shortperiod
when thefeeding during
lactation was deficient.These differences in the lactation response to different
pre-partum feeding
levels mayexplain
at least some of the contradictions found in the literature on thissubject.
That fat
percentage
falls as milkyield rises, particularly
on a concentratedration,
is known from the literature ondairy
cows(STORR!1·, ig 7 o).
The nutritionin late pregnancy
(steaming-up levels)
seems to have affectedsingle-bearing
ewes,in
agreement
with SPRAY( 1950 )
fornon-ruminating animals,
and with TREACHER(
1971
)
forsheep.
Theseinvestigators
determined thatmilk-producing
femalescan draw energy from
depots
formed in thebody
late ingestation.
Single-bearing
ewes of theadjusted regime
had ahigher
fatpercentage
intheir milk
despite
the fact that in this treatmentthey
alsoproduced
more milk.When this is added to other
evidence,
it leads us to the conclusion that thepattern
at which a ration is
being
increasedduring gestation
may affect thepartition
ofmetabolites in the animal’s
body
both before and afterparturition.
Differences in SNF concentration weresmaller,
butthey
showed similar trends.Accepté pour publication en Nvrier T978.
Acknowledgements
This study has been supported by a grant from the Israeli Sheep Breeders Association.
We also wish to express our gratitude to blr. A. LAW and the team of workers caring for the ex- perimental flock at Newe Ya’ar for the good care of the animals, and to Mr. Y. AST, Mrs. J. VER- Ti;stt-I,
FV y
, Mr. 5. lBSV-Yomnn and Mr. I,. T!r,By for technical assistance.
Résumé
Réponses
de brebis laitières avant etaprès l’agnelage
à
différents
niveauxd’a!ports
alimentairespendant
lagestation
II. - Co!asommation
d’énergie
et variation depoids pendant la lactation,
et
!yoduetion
laitièreOn a étudié certaines répercussions des niveaux d’apports alimentaires distribués pendant
la gestation sur le déroulement de la lactation suivante chez des brebis laitières. Trois séquences d’apports alimentaires de base (fh, fn et nn) et trois niveaux de supplémentation à la fin de la
gestation (H, L et A) ont été comparés dans une expérience factorielle. Toutes les brebis ont reçu
chaque jour o,4kg de foin et de l’aliment concentré aggloméré ad Li6itxsm.
Les brebis grasses à l’approche des deux derniers mois de gestation (séquence fh) ont moins
consommé au début de la lactation (— 2I p. 100) que les brebis plus restreintes avant la fin de la gestation (séquences fn et nn). Cette diminution des quantités ingérées était plus marquée
chez les brebis ayant mis bas des agneaux simples que chez celles ayant mis bas des jumeaux.
Les niveaux de supplémentation à la fin de la gestation n’affectent pas les quantités ingérées
au début de la lactation. Les brebis mères de jumeaux consomment 9,6 p. 100 de plus que les brebis mères d’un unique agneau.
On observe une corrélation négative entre la variation de poids pendant la lactation et celle pendant la gestation.
Un état d’engraissement excessif des brebis se traduit par une faible production laitière
ultérieure. Lorsque la supplémentation était constante pendant la fin de la gestation (niveaux
H et L), la production de lait des mères de jumeaux était significativement supérieure à celles
des mères d’un seul agneau. Cette différence n’existait pas pour les brebis dont la supplémentation
était progressivement augmentée en fonction de la teneur plasmatique en acides gras libres
(niveau A).
Il n’existait pas de liaison entre les quantités ingérées et les variations de poids pendant la
lactation d’une part, et entre les variations de poids et la production laitière d’autre part. Par contre, une corrélation hautement significatiwe existait entre les quantités ingérées et la production
laitière.
Le taux de matières grasses du lait n’a pas été affecté par les niveaux d’apports alimentaires distribués pendant la gestation. Il a été en moyenne de 6,7- 0,2 p. 100et 7,5 . o,2p. 100pour les mères de jumeaux et d’un seul agneau respectivement.
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