GENETIC ASPECTS OF MEAT QUALITY IN PIGS ( 1 )
P. JONSSON P. JENSEN O. K. PEDERSEN National Institute of animal Science, Department for Experiments with Pigs and Horses,
25, Rolighedsvej, 1958 Copenhagen V, Denmark
SUMMARY
Danish investigations have shown that environmental conditions like season of year, tempe-
rature during transport, and carcass weight have, on an average of a larger sample of individuals, only a slight effect at least on the variance of meat colour in the muscle.
Investigations within many breeds of pigs, including those given in tables 7, 8, and 10,
give estimates of additive gene effects of h2 = 0.3 to h2 = 0.4 for values of muscle colour and 4
5 mn pH values.
The genetic correlation between sexes in meat colour and in two other characters is given.
The phenotypic and genetic variability of characteristics related to meat quality are given together with objective carcass measurements, and the phenotypic and genetic relationship
between these are discussed.
The possibility of being able to master, through a selection program, the problem of meat quality and the problem of stress adaptability related to this, is discussed.
INTRODUCTION
Problems on meat
quality
have beenreported
asearly
as in1 88 3 ,
and in con-nection with a
pork
exhibition in Berlin HERTER and WILSDORF(IC!I4)
dealt withproblems
like meat with apale,
moist surfacecreating processing problems.
Theseauthors also dealt with breed differences in muscle colour.
Already
in theearly days
of the Danish baconproduction
the meat colour was taken intoconsideration,
as the first director of the Danish
pig
progenytesting
stationsreported
ofcomplaints
from Great
Britain, indicating
thatoccasionnally
the Danish bacon had a poor,pale
colour(BECK, 1931 ).
Aregulation
was maderequiring
that all test carcassesshould be scored for colour in m. L. dorsi. This
subjective
colourgrading
coveredan
arbitrary
five class scalegiven
on the surface of the cross section of them. L. dorsi behind the shoulder and another one in front of the ham.
(’) Invited report presented in the Study Meeting of the European Association for Animal Production, joint session of commission on Pig Production and commission on Animal Genetics, Versailles, France, July igth, 1971.
In their
description
of acute heartproblems
associated with sudden death inpigs,
FREDE( 192 6)
and HUPKA( I939 )
used thedesignation
« musculardegene-
ration » for the most
pronounced
cases of muscles with discolouration. I,uDVIGS!N( 1953
, 1954 )
described muscular discolouration in the Danish Landyace in connection withprocessing
andcanning problems
and claimed that this discolouration involved both nutritional andgenetic
factors.W I S M E R -PE D E R SE
N
( 1959 )
found aphenotypic
correlation of Yp = —0.!1 between the
pH
measurement in the m. L. doysi behind the last rib 45 minutes afterkilling
and thewater-holding capacity
of this muscle. He also found a corre-lation of rp = — 0.86 between the same
pH
value and thecorresponding
concen-tration of lactic acid. CLAUSFN and N6RTOFT THOMSEN
(ig6o)
associated ahigh
acid content of the meat with
pale
colour characteristics andreported
a correla-tion of rp = -! 0.6 between these traits.
W ISM E R -PE D E RS E
N and BRISKEY
( 19 6 1 )
were able toproduce pale,
moistmeat
by delaying
thetemperature
fall in the carcass afterkilling. They
concludedthat the fast process of
chilling
the carcasspost
mortem caused apartial
reduction inpale
colour characteristics ofporcine
muscle. HAI,I,UNG( I g6 2 )
confirmed these results.Because of the
relationship
shownby
I,uDVIGS!N( 1953 ,
1954,1955 )
betweenthe colour
changes
in the skeletal muscles and the concentration of lactic acid in the samemuscle,
CLAUSEN and NBRTOFT( 195 6)
introduced anarbitrary
colourscale of Io classes to be
given
on the cross section of the m. L. doysi cut at thetip
of the last rib. This score was introduced in
January
1954 and is still used on all testpigs slaughtered. During
theperiod 1954 - 19 65
certain environmental andgenetic
causation factors have beeninvestigated.
As earlier
mentioned,
WISMER-PEDERSEN( 1959 )
found aphenotypic
correla-tion of rp = —
0 . 71
between the
pH
value in the m. L. dorsi behind the last rib 45 minutespost
mortem and the waterholding capacity
in the same muscle. Fur- thermore he found Yp = — 0.86 between this 45 minutespH
value and the corres-ponding
concentration of lacticacid ; consequently
the 45 minutespH
value was measured on all Danish testpigs
from195 8
to19 6 2 ,
but due tomissing
dataonly
the material from the test year
I g 5 8- 5 g
could beanalyzed.
Table I shows the
relationship
between the Danish colour score and the 45 mi- nutespH,
both taken at thetip
of the last rib.ENVIRONMENTAL FACTORS AFFECTING THE MEAT COLOUR
I,UDVIGS!N
( 1954 )
and WISMER-PEDERSEN and RIEMANN( 19 6 0 )
discussedthe
importance
ofpreventing
thepigs
fromfighting
andbiting during transport
to the bacon
factory
in order to reduce the incidence ofpale,
moist muscles. In Ig599 the Danish Meat Research Institutedeveloped
a halter to beplaced
on thepigs
before
delivery
at the baconfactory (W ICHMAN - J6 R GE NS E N
1959,I9 6 I ).
This halterwas tested on
pigs
from the three progenytesting
stations fromJanuary
toJuly 19
6 0
,
and caused ageneral improvement
in the mean of the muscle colour score of the testpigs. (« Sjaelland
» : P ! 0.20,« Fyn
« : P ! o.ooi, and« Jylland
» P <0 . 20 ).
Using halter,
thephenotypic
variance within the sameday
ofdelivery
wasalso
decreased,
but most in the uncastratedanimals,
thegilts (table 2 ).
It is
clearly
shown that theprocedure
ofusing
halter decreases the variance betweenpigs
delivered on the samelorry,
and that thegilts
are the most sensiblesex, their variance in meat colour score
being
app. io per centlarger
than thatof the castrates. A remarkable effect of the distance
(km
to baconfactory)
on theintra week variance in meat colour is demonstrated.
To
repeat
this examination of the effect oftransport
stress on both the colourscore and the 45 minutes
pH value,
anexperiment
was carried out with 54 test groups(each
test group consisted of two castrates and twogilts)
to be delivered from thenewly
established fourth test station to the baconfactory, driving
distancebeing
0.5 km. Onegilt
and one castrate from each of the 54test groups were killed in their individual pens andtransported
dead to the baconfactory,
whereas the othergilt
and castrate litter mates weretransported
alive to thefactory (N 6RTO rr THOMS!rr, 10 6 1 ).
The results are shown in table 3.The muscle colour mean was
improved
in both the castrates and thegilts
killed in the pen. The standard deviation within litter and sex decreased in both sexes, but most in the
gilts. However, only
the standard deviation of the 45minutespH
value in the muscle differedsignificantly
in thegilts
killed in the pen from that of thegilts
killed at thefactory (P < 0 . 04 8,
table3 ).
To
investigate
the effect oftransportation
on the meatquality
in order tostandardize the treatment
during transport
of notonly
the testpigs,
but also ordi- nary baconpigs
from commercialproducers,
the Danish Meat Research Institutehas set into
operation
anexperiment
in 1971(W I C HMANN -J 6R GE N SE N , 1971 ).
The effect of season on both the meat colour score and 45 mn
pH
in them. L. dorsi is
investigated examining
their variances. The effect of the 3months-season isapproximately
5 per cent of thephenotypic
variance. In table 4 isgiven
therelative variance of the month and the
day
ofdelivery
and of the individual testpig
for each of six station-sexsubgroups
within the test year195 8-59.
It is
clearly
shown that8 5
to Ioo per cent of thephenotypic
variance is deuto the difference between the individual reaction among
pigs
delivered to the baconfactory
on the samelorry
on the sameday.
The influence of outside
temperature
on the muscle colour score was estimatedas the linear
regression
of the mean muscle colour score on thetemperature,
measu- red outside the test stations at noon in Co for all test stations and sexes over fouryears. The mean
temperature
of year fluctuated from 7.2 Co to II.OC ° ,
and theregression
estimates fluctuated from-f-
o.ooo 02 to - 0.02. 24 out of 28regression
coefficients showed a
negative
influence of the outdoortemperature,
thedelivery
distance of 14 kmshowing
thestrongest
influence.Eftect of
chilled carcassweight
on muscle colourand 45 mrc
PH
in m. I,. dorsiTable 5 shows the
repeatability
of theconsistently negative
effects of chilledcarcass
weight
on both muscle colour andpH though
these effects areonly slight, being
of the order between o.3per cent and4 .8
per cent of thephenotypic
variance.Similarity
inrelationship
to other charactersfor
the 4.5 mnPH
valueand
for
the meat colour both in the m. I,. dorsi at thetip of
the last rib In theprevious
tables 2, 3, 4, and 5 it is shown that these two characters have reactedquite similarly against
the causative effects mentioned. The reactions of these two characters with a third one are also very similar which is shown in table 6.The correlation of the residual sector is a measure of the covariation within litters after the elimination of the additive gene effect. The effects of the inter-
(epistatic)
and intra-
(dominant)
allelic gene action are included in this correlation.Meat colour and
pH
in the muscle arenegatively
correlated with fastgain
andpositively
with feedconsumption
rate. At constantgain
the covariation between feedconsumption
and these two characters is notchanged. Length
of thepig
andall fat measurements are
positively
correlated with colour andpH
in the muscle whereas meat content isnegatively
correlated. None of these correlations arestrong.
The intra-station
phenotypic
correlation between the two characters was esti- mated at + 0.68 and + 0.71 for castrates andgilts, respectively. However,
it isof more
importance
that in the same material of 17IS castrates and i 8mgilts,
thegenetic
correlation was estimated at rG =-f-
0.86( 4 8 0
d. f. forsires).
H!RITABIIrITY ESTIMATES FOR POINTS FOR MEAT COLOUR AND
45
MN PH,BOTH IN THE M. L. DORSI AT THE TIP OF THE LAST RIB
The
partitioning
of thephenotypic
variance of the meat colour score and two otherimportant
carcass characteristics are based on the data from195 6
to1965 (table 8).
This should include a sufficient number of test year - test station - sexsubgroups,
one year of testcomprising
material from four teststations,
and thereforeeight subgroups including
app. godegrees
of freedom for sires.The hierarchical structure of these estimates from
195 8
to19 65
is : two testpigs
of the same sex per test group, five testpigs
of the same sex per sire half-sibfamily,
and 12 testpigs
of the same sex perbreeding
centre(elite herd).
Within the elite herds
relationships
of y.5 p. Ioo were found between dams mated to the samesire,
and 2.6 p. IOO between siresstanding
at the samebreeding
centre.
Thus the data in tables 7 and 8 are corrected for the effect of the
relationship
between sires and for that between
dams ;
further for the influence of chilled carcassweight,
seasons of year, progeny teststations,
and year of test.The
pH value,
measured 45 minutes afterkilling,
is included in the interna- tionalrecognized
criteria forclassifying
apig
carcass to be either normal orPSE
(e.
g. DuTSOrr etal., 1971 ).
In table 7 aregiven
the results of theonly complete
test year in
respect
to data setincluding
this criterion for meatquality.
This is thesame set of data as used for the results in table 6.
An essential fact in table 7 is the characteristic
higher heritability
in bothcriteria for the
gilts
than for the castrates in thissample. However,
this dramatic difference between sexes isundoubtedly
due tosampling
variation as it is not foundover the 9 years
period 195 6- 19 6 5
for the meat colour score(table 8 ;
castrates : h2 =
0 . 27 J; 0 . 0 6, gilts :
h2 =0 . 3 6 J; 0 . 0 6).
As an average between the two sexes, an estimate of h2 = 0.32 for the 45 mn
pH
value(table 7 )
is amoderately high heritability, indicating
that the additive gene effect iscontrolling
the structural conditions in thepig ;
WISMER-PEDERSENhaving
demonstrated afairly high phenotypic
correlation of Yp = — 0.71 between this 45 mnpH
value and the waterholding capacity.
The best estimate of the
partitioning
of thephenotypic
variance in the DanishLandrace
pig
in theperiod prior
to19 6 7
isgiven
in table 8 for three characters which areimportant
for the selection of bacontype.
PE D E RS E
N
( 19 6 4 )
found that the m. L. doysi area controls app. 25 p. 100 of the lean meat content in the carcass( Y p (castrates)
= + 0.44 andY p (gi l ts)
=+ o.aC!),
whereas he found that the side fat measurement controls 50 p. Ioo of the lean meat content in the carcass
(Y P (castrates)
_ - 0.71 andYp(!,it.)
_ -0.68).
That is the reason
why
these two characters are included in table 8together
with the meat colour score.
Difference between sexes is not found in the colour score mean like in the two other
characteristics,
but thephenotypic
standard deviation as well as thegenetic
one differ between the two sexes in all characters as demonstraded in the tables 7 and 8. This difference in variance is about 8 p. Ioo, and a similar difference is found in the score for nasal alterations :
Meat colour score : .-
1956-60
S2 intra litter
(gilts)/s 2 tntra
Utter(castrates)
=0.235/0.217
= 1.08.Nasal alterations
(yhinitis score) :
.-1956-60
s2 (gilts) IS (castrates)
=0 .8 24/0 .7 35
= 1.12.In table 8 it is
clearly
demonstrated that under asystem
of test, where it is necessary to restrict the material so thatonly
a little more than fivepigs
per sirehalf-sib
family
andonly
12pigs
per herd isobtained,
it is necessary to include anumber of test years to
get
sufficient unbiased estimates of thepopulation
para- meters in the breed. This agrees with thetheory given by
RoBER’rsorr( I9 6 0 )
aboutexperimental design
on the measurement of heritabilities.Besides
having
a sufficient number of individuals persubgroup
toget
unbiased estimates of the different intra-classcorrelations,
the years must cover some siregenerations
because thesample
ofpaternal
half sib groups sent to the test station per year is notnecessarily representative
for thepotentialities
ofzygotes
from thebreeding
centres as a whole.Sex differences in the heritabilities are not demonstrated in the muscle arae
and the side fat measurement in the overall estimates within test stations and years.
In the
points
for meat colour it should be concluded that the sirecomponent
esti- mated from the castrates data tends to be decreasedand, therefore,
the « litter environment » and the error variance iscorrespondingly
increased. In abreeding
program it
should, therefore,
be more efficient to base the selection on data from uncastrated animals.It has been shown
previously
that the effects teststations,
seasons of year, and chilled carcassweight
areaffecting
the meat colouronly slightly (tables
5 and7 ).
Table 8 shows that the
only
two causations which matter for the meat colour is theheritability
and the residual error. Ifonly gilts
are included in the selection program, it is realistic to work with aheritability
of 0.4 and a residual error of o.6.If both sexes are included in the test group, the
heritability
is 0.3 and the residualerror is o.7.
A rather
strong
maternal effect is found in the muscle area. This could to some extent be due tomothering
abilities ofprenatal
nature.GENETIC COVARIATION BETWEEN
THE TWO SEXES IN POINTS FOR MEAT COLOUR
AND TWO OTHER TRAITS
The test groups of the litters from the state
recognized breeding
centres consistof 2castrated males and 2females. Because of the
uncertainty
of thegenetic
varianceof the meat colour in the castrates and also because the
genetic improvement
ofa character is increased per year when
only using
2 insteadof 4
litter mates dueto the increased selection
intensity,
it was of interest toinvestigate
thegenetic
correlation between the two sexes with
respect
to theirperformance
in the threecarcass characteristics
(table 9 ) (J ONSSON ,
1071b).
The variance
components
for the interaction between sire half-sib families and the two sexes were very small in the meat colour score and in the side fat measure-ment, 0.003
4 points 2
and o.0028 cm 2 , respectively ;
theF-quotients
were 1,09 and 1.IS,respectively.
This is the reasonwhy
thecorresponding genetic
correlationsare not
unity.
In the muse.Long.
doysi area,however,
theF-quotient
was consis-tently
beneathunity
in the different testyear-test
stationsubgroups,
so no sire-sexinteraction is found in this character.
PHENOTYPIC AND GENETIC VARIATION AND COVARIATION IN SOME IMPORTANT
CARCASS AND MEAT CHARACTERISTICS IN THE DANISH LANDRACE PIG
The two Danish carcass evaluation centres were started in
19 6 7
toinvestigate
the new characters for carcass
quality
which were to be recorded at these centres.The material
comprised
1 403gilts
and 1 400 castrates. As this firstinvestigation
on the new carcass characteristics was
planned
so that sires would be tested on at least two test groups(two gilts
and two castrates per test group ; it was notpossible,
as
originally planned,
to set the limit at three testgroups),
it wasimpossible
toinclude the variation between
breeding
centres in the hierarchical classification. So any carry-over effect frombreeding
centres will be included in the sirecomponent.
The hierarchical structure of the
analysis
was as follows :The standard errors for the
heritability
estimates werecomputed according
tothe method
given by
B. Woo!,x(FALCONER, 19 6 3 )
as shown inJorrssorr (i g y a).
The per cent of lean meat in the entire carcass side
(character 9 )
ispredicted by
io individual carcass measurements andweights, including
cold carcassweight
and sex. The side fat measurement is a
prominent
x-variable in theprediction equation, controlling
app. 50per cent of the lean content in the entire side. R =0 .8 7 (C
LAUSEN
e1 al., 19 68).
Table Igives
thephenotypic
andgenetic population parameter
estimates in the Danish Landracepig
for six traditional and six new carcassquality
characters introduced at the two carcass evaluation centres.
For the average backfat thickness the
magnitude
ofheritability given
in table Ioseems to be more
reasonable,
and for the m. L. dorsi area the valuegiven
in table 8seems more reasonable. An estimate of h2 ! 0.62 seems too
high
for the area ofm. L.
dorsi ;
this should also be the case for characters nr.6,
8 and 9 in table io.One reason for this must be the lack of
including
the class for« breeding
centres »in the
hierarchy
and the lack ofhaving
corrected for seasonal differences. Theperiod
of
investigation
isperhaps
also a little short. The elite breeders havegiven
consi-derable attention to the three characters m. L. dorsi area, side-fat measurement and per cent lean meat in the entire side from
19 6 7
and onwards. This means that theeffect of selection is included in the differences between sires’ half-sib groups.
Having
included more years and corrected for effects from seasons and years, and
having
included the classification
breeding
centres in thehierarchy,
aprobable upward
bias will be corrected for.
But the estimates for the additive gene effect in table io show that no lack of additive
genetic variability
in the breedecists,
which is also confirmedby
the esti-mates
given by
STAUN( 19 68)
and STAUN andJE NS E N ( I970 )
for the same breed.Their
heritability
estimates for the samecharacters,
estimated from the data from thepig
progenytesting stations,
rank fromo. 4 6
to0 . 7 8.
The
magnitude
of thegenetic
standard deviation and the coefficient of variation for theimportant
characters isstriking.
Heritability
estimates for colour valuesreported
from other breeds range from0
. I
8 ::!::
0.06 forNorwegian
Landrace(I, ANG FI O I, Z , 19 66)
over0 . 25 ::!::
o.09 forLa y ge
White
(PEASE
andSMITH, 19 65), 0.28 !
0.15( JE NS E N e1 al., 19 6 7 )
as an estimatewithin the breeds
Duroc, Yo y kshi y e, Hccmpshire,
PolandChina,
andSpotted
Swinebreeds,
to0.38 !
O.IO for Deutsches veredeltes Landschwein(F!.ocx, 19 68).
Theseestimates are similar to the
present
estimategiven
for the character 6.In Denmark cross
breeding experiments
between thepig
breedsLarge
Whiteand Danish Landyace are
planned.
One of the mainsubjects
to beinvestigated
isthe
hypothesis
of dominant gene effect on meatquality
characteristics. SYBFSMA( I970
)
hassuggested
that Crossbreeding
different breeds is a verypromising
meansof
improving
meatquality.
WrsM!R-P!D!RS!rr
( 1959 )
found a correlation of yr = o.y between 45 mnpH
and 24hours waterholding capacity,
both charactersgiving
a reasonable accurate measure of structural conditions in the tissue. W!NiG!x et al.( 1970 , reported
fromW E I SS
,
19 6 7 )
foundheritability
estimates of h2 =0.37 !
0.14 for WHC(centrifug- ing)
and h2 =0 . 19 ::!::
o.io for 45 mnpH
value. The estimates ofheritability given
among others
by
WENiG!R et al.( I970 )
for the WHC of h2 =0.37 !
0.14 and that for the 45mnpH
value of h2 = 0.32 from table 7aresupporting
thehypothesis
that structural conditions in the muscle as a meat
quality
criterion is controlledby
additive geneeffect, and, therefore,
can be included in a selection program forimprovement
of meatquality.
In table m, the traditional meat and carcass
quality
characters aregiven.
The
genetic
correlations aregiven
below thediagonal
and thephenotypic
correlationsare
given
above thediagonal.
It is
doubtful,
whether thepositive genetic
correlation between colourbright-
ness and muscle size still exists in the Danish Landrace
pig,
asreported by J ONSSON
(i g
y a).
Thischange
insign
could have been forcedby
achange
in genefrequency
in the Danish Landyace
pig
because of a more direct selection for meat contentduring
the recent years. Table igives
the correlation values as follows : Correlation between rp rG- --
Points for meat colour in m. L. dorsi
X m. L. dorsi area - 0.08 - o.2g This correlation is
slight,
but indicates anegative
trend.The
negative
trend between muscle colour and size of muscle agrees withestimates from other
breeds,
e. g. with thatgiven by
F!ocx( 19 68),
who estimatedr
p = — 0.28 and rG = —
0 . 5 6
between these two characters in the German vere-deltes Landschwein breed.
DISCUSSION
In tables 2, 3, !, and 5 it is demonstrated that the average environment has
only
aslight
effect on the meat colour in the carcass. For the Danish Landyacepig
table 7, 8 and io show
together
with estimates from other breeds thatbrightness
of colour is influenced to a moderate
degree by
additive geneaction, heritability
estimates
being
of amagnitude
of h2 ! 0.3 for colour values.The
high genetic
correlation between sexes in meat colour as well as in other charactersimportant
for value of the carcass tells thatincluding only
one in thefamily
selection for carcass and for meat evaluation ensuresequal genetic
progress in the other sex.However,
theproblem
of structure in meat tissue is not connected with musclecolour. It
is, however, significantly
related to the stresssyndrome
asreported
e. g.by
LUDVIGSEN( 19 68
a and19 68 b), JUDGE ( 19 6 9 ),
STEINHAUF el al.( 19 6 9 ),
BRISKEYand LISTER
( 19 68),
HAASE and STW rrHaux( 1971 ),
STAUN( 19 68),
and many others.Undoubtedly,
it isonly
a matter oftechnique
to be able to obtain sufficient accurate andrepeatable
WHC values to ensurereasonably high heritability
estimatesto be included in selection programs for
improvement
of the meat structure, gene-tically.
But all these characters have
only
asecondary
effect on theadaptability
ofthe
pig
to environmental stress conditions. The mainproblem
of the future must be tolay
open the characteristics of the stresssyndrome
in thepig
and thegenetic
effect behind these
adaptation
characters. In this way thequestion
can beanswered,
whether the
problem
ofquality
and deathlosses,
asreported
from many countries(e.
g.by
WENIGER e1al., 1970 )
can befought against by
means of selection within the breedpopulations and/or by
means ofcrossing
between breedsutilizing
a pro- bableheterosis,
or may be be masteredthrough
environmental measures.GERRiTS ! at.
( I9 6 9 )
have shown that intense selection for meatiness inpigs
has a
significant
correlated response togrowth
hormone concentration. Considerable need exists for more selectionexperiments
like this cited tolay
open theseproblems.
Reçu pour publication en se!tembre 1971.
RÉSUMÉ
ASPECTS
GÉNÉTIQUES
DE LAQUALITÉ
DE LA VIANDE CHEZ LE PORC Les recherches effectuées au Danemark ont montré que l’influence des conditions ambiantes, telles que la période de l’année, la température pendant le transport et le poids de la carcasse,sur un assez grand nombre d’animaux examinés est faible, du moins sur la variance de la couleur de la viande dans le muscle.
Les recherches à l’intérieur de nombreuses races porcines, y compris les recherches effectuées
au Danemark sur les animaux de la Landvace danoise, ont permis d’évaluer à h2 = 0,4 et h2 = 0,3 les actions additives de gènes pour les valeurs du pH après 45 minutes et celles de la couleur de la viande, toutes les deux mesurées sur les muscles dorsaux à l’extrémité de la dernière côte.
Le matériel animal suivant a été employé pour ces recherches : matériel de 1958-59 pour les valeurs du pH après 45 minutes et le pointage de la couleur de la viande, matériel de 1956 à 19
6
5 pour le pointage de la couleur de la viande et matériel de 1967-69également pour le poin- tage de la couleur de la viande.
On donne la corrélation génétique entre les sexes en ce qui concerne la couleur de la viande et deux autres caractères.
On donne en outre la variabilité phénotypique et génétique des caractéristiques relatives
à la qualité de la viande ainsi que les mensurations objectives de la carcasse, et on examine la relation phénotypique et génétique existant entre ces caractéristiques.
On étudie également la possibilité de venir à bout, au moyen d’un programme de sélection
approprié, du problème de la qualité de la viande ainsi que de celui de l’adaptabilité aux stress, qui s’y rattache.
RÉFÉRENCES BIBLIOGRAPHIQUES
B
ECK N., 1931. ig. beretning om sammenlignende forsog med svin fra statsanerkendte avlscentre 139
. beretning fra forsôgslaboratoriet, Kôbenhavn, 184 pp.
B
RISKEY E. J., LISTER D., 1968. Influence of stress syndrome on chemical and physical characte- ristics of muscle post mortem. In : D. G. TOPEL, The Pork Industry, Problems and progress. Iowa State University Press, Ames, Iowa. XIV, 236 pp.
CL AU S
EN H., NBRTOFT THOMSEN R., 1956. 44. beretning om sammenlignende forsog med svin fra statsanerkendte avlscentre. 288. beretning fra forsôgslaboratoriet, Kôbenhavn, 15o pp.
C
LAUSEN H., NÔRTOFT THOMSEN R., 1960. 48. beretning om sammenlignende fors&g med svin fra statsanerkendte avlscentre. 317. beretning fra forsôgslaboratoriet, Kobenhavn, 127 pp.
C
LAUSEN H., NÔRTOFT THOMSEN R., PEDERSEN O. K., 1968. 56. beretning om sammenhgnende forsog med svin fra statsanerkendte avlscentre. 364. beretning fra forsügslaboratoriet, Kobenhavn’
177 PP.
D
UTSON T. R., PEARSON A. M., MERKEL R. A., KOCH D. E., WEATHERSPOON J. B., 1971. Histo- chemical activity of some lyso3omal enzymes in normal and in pale, soft and exudative pig muscle.
J. Animal. Sci., 32, 233-238.
FALC
ONER D. S., 1963. Quantitative inheritance. In W. J. BURDETTE : Methodology in mammalian genetics, 193-216.
F
LOCK D. K., 1968. Farbhelligkeit im musculus longissimus dorsi als Selektionsmerkmal beim Schwein.
Die Fleischwirtschaft, 48, 1362-1365.
F
REDE W., 1926. Toxische Herzerkrankung bei Schweinen. Deutsche tierürztliche Wochenschrift, 34, 6
5 8-659.
GE
RRITS R. J., HETZER H. O., RICHARDSON G. V., 1969. Growth hormone potency in lines of swine selected for high and low backfat. Abstract, J. Animal Sci., 28, 873.
HA
ASE S., STEINHAUF D., I97I. Effects of stress on some oxygen metabolism parameters in boars. 2. In- ternational symposium on condition and meat quality in pigs given at the R.esearch Institute for Animal Husbandry, « Schoonoord », Zeist, March 22-24, 1971.
H
ALLUND O., 1962. Eôdfarvens afhaengighed af kâleforlâbel. Not published research data, Danish Meat Research Institute, Roskilde.
H
ERTER M., WILSDORF G., 1914. Die Bedeutung des Schweines für die Fleischversorgung. Deutsche Landwirtschaftgesellschaft, Berlin, XVII + 381 pp.
H
UPKA E., 1939. Seuchenhaft auftretende hühnerfleischâhnliche Muskelentartungen unter den Schwei-
nen. Deutsche tierârztliche Wochenschrift, 47, 242-244.
J
ENSEN P., CRAIG H. B., ROBISON O. W., 1967. Phenotypic and genetic associations among carcass
traits of swine. J. Animal Sci., 26, 1252-1660. Jo
N
ssoN P., I97Ia. Population parameter estimates of the Danish Landrace Pig. Acta Agr. Scand.
21, II-16.
J
ONSSON P., 1971 b. Model computation on future expectation of genetic gain. Acta Agr. Scand., 21, 17-25.
J
UDGE hi. D., 1969. Environmental stress and meat quality. J. Animal Sci., 28, 755-759.
L
ANGHOLZ H.-J., 1966. Das züchterische Hilfsmittel der stationâren Nachkommenprüfung beim Schwein III. Heritabilitaten und genetische Korrelationen beim norwegischen Landschwein. Acta Agr. Scand., 16, 97-114.