ddd.uab.cat · 2014-12-17 · iv RESUMEN! # # Parapoderimplementar...
Transcript of ddd.uab.cat · 2014-12-17 · iv RESUMEN! # # Parapoderimplementar...
DEPARTAMENT DE CIÈNCIA ANIMAL I DELS ALIMENTS
CARACTERIZACIÓN MORFOLÓGICA, DE LA CANAL Y DE
LA CARNE, E INFERENCIA DE MATERNIDADES EN OVINOS
DE RAZA RIPOLLESA
CECILIA ESQUIVELZETA RABELL
Programa de Doctorado en Producción Animal
Bellaterra, julio de 2012
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Universitat Autònoma de Barcelona Departament de Ciència Animal i dels Aliments
El trabajo de investigación “Caracterización morfológica, de la canal y de la carne, e inferencia de maternidades en ovinos de raza Ripollesa’’ ha sido realizado por Cecilia Esquivelzeta Rabell1 y se presenta como requisito para optar por el grado de Doctor. Bellaterra, veinticinco de mayo de dos mil doce.
Los directores de Tesis, La doctoranda, Dr. Jesús Piedrafita Arilla y Dr. Joaquim Casellas Vidal Cecilia Esquivelzeta Rabell 1 Beca del Consejo Nacional de Ciencia y Tecnología, México (CONACyT).
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Gracias a todos los que han estado presentes y me
han ayudado de alguna u otra forma a alcanzar
esta meta, jamás encontraré la forma de
agradecer su apoyo y confianza.
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RESUMEN
Para poder implementar programas de mejora genética en el sector ovino de
carne, es imprescindible definir y evaluar los caracteres biológicos con relevancia
económica en la producción. Es por ello que la caracterización morfológica de la raza
ovina Ripollesa, la diferenciación entre rebaños, la utilización de metodologías analíticas
para evitar pérdida de información relevante y el estudio de las características de la canal
y de la carne, se definieron como objetivos del presente trabajo.
La raza ovina Ripollesa es una importante raza autóctona Española localizada
principalmente en las montañas del Mediterráneo y en llanuras cultivadas de Cataluña,
explotada bajo sistemas semi-‐extensivos de producción y actualmente destinada a la
producción de carne. La caracterización morfológica de la raza Ripollesa se llevó a cabo a
partir del estudio de ocho rebaños representativos de diferentes subpoblaciones
históricas. Un total de 224 hembras Ripollesas y 17 moruecos fueron caracterizados
registrando su peso vivo y 12 medidas morfológicas, las cuales fueron utilizadas para el
cálculo de 12 índices zoométricos. Se encontraron diferencias (p < 0.05) entre rebaños y
sexos respecto a varias medidas. El análisis de componentes principales realizado con las
medidas morfológicas reveló dos componentes principales que contribuyen con 47,6% y
12,3% de la inercia, estando relacionados con el tamaño (tamaño corporal y peso vivo) y
con la altura a la grupa y longitud de oreja, respectivamente. El análisis de cluster nos
permitió diferenciar cuatro subpoblaciones con implicaciones relevantes a ser tomadas
en cuenta en el programa de conservación o mejora de la raza. Las diferencias observadas
pueden atribuirse a la localización geográfica, historia selectiva, manejo del rebaño, y las
diferencias genéticas. Los índices zoométricos indican que la raza ovina Ripollesa es de
tamaño medio y longilínea, con una marcada orientación a la producción de carne y con
signos de adaptación al ambiente.
En los rebaños ovinos, la correcta identificación de los progenitores presenta a
menudo problemas debidos a la presencia de varios candidatos posibles tanto por vía
paterna como materna. Un ejemplo típico en donde las paternidades deben de ser
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elucidadas a partir de una lista de moruecos es en sistemas reproductivos con múltiples
machos, generalmente bajo condiciones extensivas de pastoreo. La misma situación
ocurre por la vía materna cuando se producen múltiples nacimientos en el mismo día y
uno o varios de los corderos no son adoptados por sus madres respectivas. Este
fenómeno conlleva a un porcentaje entre moderado y bajo de corderos con pedigrí
incompleto, con la consiguiente pérdida de información para el programa de
mejoramiento genético. Debido a las restricciones económicas inherentes de la industria
ovina, el genotipado sistemático de todos los corderos abandonados no es
económicamente viable para los productores y sólo un número reducido de corderos
abandonados podrían ser validados por medio de esta técnica. Dentro de este contexto,
se adaptó un modelo Bayesiano para identificar la madre en el caso de corderos
abandonados, integrando fuentes de información tanto genéticas como
medioambientales de datos fenotípicos, y así considerar la oveja desconocida como un
parámetro adicional del modelo. Éste fue minuciosamente evaluado utilizando datos de
simulación y asumiendo siete escenarios diferentes en donde de uno a cuatro corderos
abandonados tenían que ser asignados a dos ovejas candidatas. La media general de la
probabilidad de asignación materna correcta (PARD) fue 0,59. Aunque las medias de
PARD oscilaron entre 0,51 y 0,70 dependiendo del escenario de simulación, y las
estimaciones de PARD oscilaron entre 0,04 y 1,0. En un sentido similar, la sensibilidad de
esta metodología analítica varió entre escenarios de simulación, aunque la mayoría de
ellos revelan valores mayores que 0,6. La presente aproximación analítica debe ser vista
como una herramienta estadística útil para realizar una proyección preliminar de los
corderos abandonados y sus posibles madres. Es importante subrayar que cualquier
inferencia en maternidades desconocidas podría reducir consecuentes costos económicos
permitiendo enfocar el genotipado en aquellos corderos con mayor probabilidad de ser
descendientes de ovejas élite.
La preocupación e importancia que ha tomado la composición de los alimentos y
su relación con la salud humana durante las últimas décadas ha sido un punto clave para
la industria ovina actual, enfatizando el interés y la importancia por parte de los
productores para poder predecir las características de la canal y conocer las propiedades
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de sus productos. Las técnicas de ultrasonido a tiempo real resultan de gran utilidad para
predecir las características de la canal en el animal vivo de manera no invasiva. Con el
objetivo de evaluar la precisión de las medidas de ultrasonido para predecir las
características de la canal, se tomaron imágenes de ultrasonido en 124 corderos
pascuales (13-‐16 kg de peso a la canal), transversal y longitudinalmente a la columna
vertebral, y a nivel torácico (TV; entre la 12 ª y 13ª costillas) y lumbar (LV; entre la 1ª y 2ª
vértebra lumbar). El grosor de la piel, grosor de la grasa subcutánea (BFT), y profundidad
(DLD), amplitud (WLD) y área (ALD) del músculo Longissimus dorsi fueron obtenidos con
ayuda del programa ImageJ 1.42q. Después del sacrificio, BFT (TV, 2,30 ± 0,06 mm; LV,
2,46 ± 0,06 mm), DLD (2,47 ± 0,03 cm; 2,48 ± 0,03 cm), WLD (4,50 ± 0,04 cm; 4,60 ± 0,04
cm) y ALD (9,96 ± 0,12 cm2; 10,19 ± 0,13 cm2) fueron medidos directamente sobre la
canal del cordero. Las correlaciones entre las medidas de ultrasonido y de la canal fueron
superiores a 0,61 para DLD, WLD y ALD (p < 0.05), mientras que fluctuaron entre 0,32 y
0,60 para BFT (p < 0,05); por otra parte, las correlaciones fueron significativamente (p <
0,05) mayores para las vistas transversales que para las longitudinales. En un sentido
similar, los análisis de regresión lineal sugirieron una moderada subestimación para BFT y
DLD lumbar utilizando tecnologías de ultrasonido, mientras que WLD, ALD y DLD a nivel
torácico fueron sub y sobreestimadas para valores pequeños y grandes de caracteres de
la canal, respectivamente. Después de descomponer el error de predicción (MSPE) para
las diferentes medidas de ultrasonido, encontramos que el error aleatorio es el que más
contribuye al MSPE, seguido del error de tendencia central y el error debido a la
regresión. El error estándar de la predicción (SEP) también fue calculado como un
indicador de precisión adicional, obteniendo estimaciones inferiores a las reportadas en
estudios previos con corderos más grandes. En conclusión, las medidas de ultrasonido
transversales tanto a nivel torácico como lumbar pueden ser una herramienta útil para
predecir DLD, WLD y ALD en corderos ligeros, y en menor medida BFT. Esta información
puede ser de relevancia especial para productores de corderos ligeros de todo el mundo,
con un énfasis especial en la zona del Mediterráneo en donde este tipo de sistemas de
producción contribuyen con un porcentaje importante a la industria ovina.
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La carne de cordero es una fuente importante de diferentes ácidos grasos con
numerosos beneficios para la salud del consumidor. El objetivo del cuarto estudio fue
caracterizar caracteres de la canal y de la carne en 14 corderos pascuales (peso a la canal,
13 a 16 kg) Lacaune, 77 Ripolleses y 33 cruzados Lacaune×Ripollesa utilizando medidas de
ultrasonido y de la canal, y análisis de laboratorio de las muestras de carne.
Específicamente, se tomaron imágenes de ultrasonido y medidas sobre la canal del
Longissimus dorsi (LD) entre la 12ª y 13ª costillas y entre la 1ª y 2ª vértebras torácicas,
caracterizando el grosor de la piel, grosor de grasa subcutánea, y profundidad, amplitud y
área del LD. Después del sacrificio, se tomaron los pesos de las diferentes partes del
despiece comercial de la canal (i.e., pierna, costillar, espalda y cuello), mientras que una
muestra del músculo LD fue analizada para composición de ácidos grasos. Las medidas de
ultrasonidos y canal no difirieron significativamente entre razas, siendo 0,16 ± 0,004 cm y
0,23 ± 0,01 cm para BFT, 2,40 ± 0,02 cm y 2,47 ± 0,03 cm para DLD, 4,53 ± 0,05 cm y 4,50
± 0,04 cm para WLD, y 9,79 ± 0,11 cm2 y 9,97 ± 0,12 cm2 para ALD, respectivamente. El
rendimiento a la canal fue mejor en la raza Lacaune seguido de la Ripollesa (p < 0,05); los
pesos del despiece comercial no mostraron diferencias significativas, aunque también
estaban correlacionados (0,2-‐0,4) con las medidas de ultrasonidos. El análisis de ácidos
grasos (FA) mostró que los FA mayormente representados en el tejido muscular fueron:
oleico (C18:1n-‐7, 38,8%) siendo mayor para Lacaune, palmítico (C16:0, 21,7%) sin
diferencias entre razas, y esteárico (C18:0, 14.3%) con menor porcentaje para Lacaune. Se
encontraron diferencias entre razas (p < 0,05) para los porcentajes de FA saturados (SFA;
39.0-‐41.2%), monoinsaturados (MUFA; 48.1-‐49,6%) y poliinsaturados (PUFA; 6,0-‐7,5%).
En cuanto a la proporciones para parámetros nutricionales, solo se encontraron
diferencias (p < 0,05) para la proporción PUFA/SFA, siendo mayor para la raza Lacaune
(0,19), seguido por la Ripollesa (0,16) y los individuos cruzados (0,15). Los valores
nutricionales para n-‐6/n-‐3 variaron desde 6,27 a 6,96, y desde 7,85 a 8,57 para C18:2 n-‐
6/C18:3 n-‐3. Toda esta información es esencial para los productores de cordero ligero, un
producto ganadero que tiene una alta demanda en el mercado Mediterráneo.
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SUMMARY
To be able to implement genetic selection programs in meat-‐producing sheep
breeds, it is essential to define and evaluate biological traits with economic relevance in
the production system. Within this context, the morphologic characterization of the
Ripollesa sheep breed, the differences between flocks, the use of analytical
methodologies to avoid the loss of relevant pedigree data, and the study of carcass and
meat trais, were defined as objectives of the present work.
The Ripollesa sheep breed is an important Spanish local breed mainly located in
the Mediterranean mountains and cultivated plains of Catalonia, exploited under semi-‐
extensive production systems and currently intended for meat production. The
morphologic characterization of the Ripollesa breed was carried out from the study of 8
flocks representative of different subpopulations. A total of 224 Ripollesa ewes and 17
rams were recorded for their live-‐weight and 12 morphological measures, which were
used for calculating 12 zootechnical indexes. Differences (p < 0.05) between flocks and
sexes were revealed for several measures. A principal component analysis was performed
on the morphological measures and revealed two main components accounting for 47.6%
and 12.3% of the inertia, being related to the frame (body size and live-‐weight) and to
withers height and ear length, respectively. The cluster analysis allowed differentiating
among four subpopulations with relevant implications to be taken into account for the
breed conservation program. The observed differences could be attributable to
geographic location, selective history, flock management, and genetics. The estimated
zootechnical indexes classified the Ripollesa sheep breed as a medium-‐sized and long-‐
shaped body frame, with a marked orientation to meat production and with signs of
adaptation to the environment.
In sheep flocks, lamb’s parents can be misidentified or confounded due to the
plausibility of two or more potential sires or dams. A typical example where paternities
must be elucidated from a list of several candidate sires is in multiple-‐sire mating systems
under pastoral conditions. The same situation occurs when two or more ewes gave birth
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the same day and one or more lambs are not adopted by its corresponding mother. This
phenomenon leads to a moderate-‐to-‐low percentage of lambs with missing pedigrees,
with the subsequent loss of information for the corresponding selection schemes. Given
the economic restrictions inherent to the sheep industry, systematic genotyping of all
abandoned lambs is not affordable by stockbreeders and only a much reduced number of
lambs could be validated by this technique. Within this context, we adapted a Bayesian
model to ascertain the dam of abandoned lambs by integrating both genetic and
environmental sources of information from phenotypic data and modeling the uncertain
dam as an additional unknown parameter. The model performance was thoroughly
evaluated by using simulation data and by assuming seven different scenarios where one
to four abandoned lambs had to be assigned to two candidate ewes. The overall average
probability of assignment to the right dam (PARD) was 0.59, although PARD averages
ranged between 0.51 and 0.70 depending on the simulation scenario, and raw PARD
estimates ranged between 0.04 and 1.0. In a similar way, sensitivity of this analytical
methodology varied across simulation scenarios, although most of them revealed values
larger than 0.6. This analytical approach must be viewed as a useful statistical tool for
performing preliminary screenings on abandoned lambs and their candidate dams. Note
that any inference on uncertain dams would reduce further economic costs by focusing
genotyping efforts on those lambs with higher probabilities to be offspring from elite
ewes.
Concern about meat composition and its relationship with human health has
increased during the last few decades has been a key point for the actual sheep farming
industry, emphasizing the interest and importance for producers to be able to predict
carcass traits and know the properties of their products. Real-‐time ultrasound techniques
are of great utility to predict carcass characteristics in the live animal in a non-‐invasive
way. With the aim to assess the accuracy of the ultrasound measurements for predicting
carcass traits, ultrasound images were taken in 124 Spanish ‘pascual’-‐type lambs (13 to 16
kg of carcass weight), transversal and longitudinal to the vertebral column, and at
thoracic (TV; between 12th and 13th ribs) and lumbar (LV; between 1st and 2nd lumbar
vertebrae) locations. Skin thickness, subcutaneous back-‐fat thickness (BFT), and depth
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(DLD), width (WLD) and area (ALD) of Longissimus dorsi were obtained by the ImageJ
1.42q software. After slaughtering, BFT (TV, 2.30 ± 0.06 mm; LV, 2.46 ± 0.06 mm), DLD
(2.47 ± 0.03 cm; 2.48 ± 0.03 cm), WLD (4.50 ± 0.04 cm; 4.60 ± 0.04 cm) and ALD (9.96 ±
0.12 cm2; 10.19 ± 0.13 cm2) were directly measured on the lamb carcass. Correlations
between ultrasound and direct carcass measurements were larger than 0.61 for DLD,
WLD and ALD (p < 0.05), whereas they fluctuated between 0.32 and 0.60 for BFT (p <
0.05); moreover, correlations were significantly (p < 0.05) higher for transversal than for
longitudinal views. In a similar way, linear regression analyses suggested a moderate
underestimation for BFT and lumbar DLD when using real-‐time ultrasound technologies,
whereas WLD, ALD and thoracic DLD suffered from under-‐ and over-‐estimation for small
and large values of carcass traits, respectively. After decomposing the error of prediction
(MSPE) for the different ultrasound measurements, we found that the error due to
disturbance contributed most to the MSPE, followed by the error of central tendency and
the error due to regression. The standard error of prediction (SEP) was also calculated as
an additional precision indicator, obtaining estimates lower than that in previous studies
with larger lambs. In conclusion, transversal ultrasound measurements at both thoracic
and lumbar levels could be a useful tool for predicting DLD, WLD and ALD in light lambs,
perhaps suffering from worse prediction properties when focusing on BFT. This
information could be of special relevance for light lamb producers worldwide, with a
special emphasis in the Mediterranean basin where this kind of production system
accounts for an outstanding percentage of the sheep industry.
Lamb meat is an important source of different fatty acids with numerous health
benefits for the consumer. The aim of the fourth study was to characterize carcass and
meat traits in 14 Lacaune, 77 Ripollesa and 33 Lacaune×Ripollesa ‘pascual’-‐type lambs
(carcass weight, 13 to 16 kg) by using ultrasound and direct measurements on carcasses
and laboratory analyses on meat samples. More specifically, ultrasound images and direct
carcass measurements of the Longissimus dorsi (LD) were taken between the 12th and
13th ribs and between the 1st and 2nd lumbar vertebrae, characterizing skin thickness,
subcutaneous back-‐fat thickness, and depth, width and area of LD. After slaughter,
standard commercial joints were weighted (i.e., leg, rack, shoulder/foreshank and neck),
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whereas a sample of the LD muscle was analyzed for fatty acid composition. They were
not found significant differences between breeds for ultrasound and carcass measures,
being 0.16 ± 0.004 and 0.23 ± 0.01 cm for GSC, 2.40 ± 0.02 and 2.47 ± 0.03 cm for PM.
4.53 ± 0.05 and 4.50 ± 0.04 cm for AM, and 9.79 ± 0.11 and 9.97 ± 0.12 cm2 for ARM,
respectively. Carcass yield was better for Lacaune followed by Ripollesa; weight of carcass
pieces did not show significant differences but were also correlated (0.2-‐0.4) with
ultrasonic measurements. The meat fatty acid (FA) analysis showed that the FA mostly
represented in muscular tissue under examination were: oleic (C18:1n-‐7, 38.8%) being
greater for L sheep breed, palmitic (C16:0, 21.7%) without differences between breeds,
and stearic (C18:0, 14.3%) lower percentage for L breed. Differences between breeds for
saturated (SFA; 39.0-‐41.2%), monounsaturated (MUFA; 48.1-‐49.6%) and polyunsaturated
FA (PUFA; 6.0-‐7.5%) proportions were found. Differences for nutritional ratios were only
found for PUFA/SFA ratio, being greater for L breed (0.19), followed by R (0.16) and RL
(0.15) breeds. Nutritional ratios for n-‐6/n-‐3 varied from 6.27 to 6.96 and from 7.85 to
8.57 for C18:2 n-‐6/C18:3 n-‐3 ratio. All of this information becomes essential for light lamb
producers, a livestock product that has high market demand in the Mediterranean basin.
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ÍNDICE
CAPÍTULO 1. Introducción 1 La importancia de los ovinos 3 Los ovinos en el mundo, en la Unión Europea y en España 4
Razas ovinas en España 9 Sistemas de producción 10
Extensivo tradicional 11 Semi-‐extensivo 12 Semi-‐intensivo 13 Intensivo 14
Producción de carne 15 Situación del sector ovino 19
CAPÍTULO 2. Objetivos CAPÍTULO 3. Revisión bibliográfica
La raza ovina Ripollesa Origen de la raza Descripción de la raza Principales características reproductivas y productivas Sistemas de producción Sistema tradicional Sistema extensivo de montaña Sistema semi-‐extensivo Asociación de criadores y situación actual
Zoometría Instrumentos y consideraciones de medición Medidas habituales y su importancia Índices zoométricos Estudios zoométricos en la raza Ripollesa
Aprovechamiento de datos biométricos Mejora genética Genética cuantitativa BLUP Asignación de paternidades
Ecografía Definición de la ecografía Funcionamiento de la ecografía Tipos principales de equipos de ultrasonido
A-‐mode B-‐mode
Ultrasonido a tiempo real TM-‐mode
Consideraciones de las medidas de ultrasonido y su relación con la predicción de la composición de la canal
21
25 27 27 27 30 32 32 32 32 33 35 35 36 37 40 41 41 41 42 43 45 45 45 47 47 48 48 49
50
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Características de la canal Canal ovina.
Calidad de la canal Peso de la canal Grado de engrasamiento de la canal Conformación de la canal Composición de la canal Composición regional Composición tisular Composición química
Clasificación de las canales Características de la carne
Definición de carne Composición química de la carne Perfil de ácidos grasos Calidad nutritiva de la carne
CAPÍTULO 4. Characterization of the Ripollesa breed Introduction Material and methods The Ripollesa breed
Sheep samples Morphological measurements Zootechnical indexes Statistical analyses
Results and discussion Morphological diversity between sexes and flocks Principal components analysis Discriminant and cluster analyses Zootechnical indexes
CAPÍTULO 5. Bayesian inference of maternities Introduction Material and methods Rationale Bayesian inference for uncertain maternities Illustration with the simulated data Results and discussion
CAPÍTULO 6. Real-‐time ultrasounds in light lambs Introduction Material and methods Data source Animals and management Weight and Real-‐Time Ultrasound Measurements Slaughter Procedure and Cold Carcass Measurements
51 51 51 52 52 53 53 54 54 54 54 59 59 59 60 61
65 67 68 68 70 73 74 75 76 76 79 82 86 89 91 92 92 93 95 99
109 111 112 112 112 112 115
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Statistical Analysis intensity of the relationship between Analysis of Ultrasound Reliability Residual Decomposition of Ultrasound Measurements Results and discussion Average Performances of Carcass Traits in Light Lambs Reliability between Ultrasound and Direct Carcass Measurements Correlation Coefficients Linear Regression Analyses Residuals for Ultrasound Data Residual Decomposition Standard Error of Prediction CAPÍTULO 7. Carcass traits and meat composition of light lambs Introduction Material and methods
Animals and management Ultrasound measurements and image analysis Slaughter procedure and carcass trait measurements Intramuscular fatty acid composition analysis Statistical analysis
Results and discussion Ultrasound and direct measurements on the Longissimus dorsi muscle
Carcass quartering Intramuscular fatty acid composition CAPÍTULO 8. Discusión Discusión general
Ventajas de incluir el uso de la técnica de ultrasonido a tiempo real en el programa de selección y mejoramiento genético de la raza ovina Ripollesa
CAPÍTULO 9. Conclusiones Bibliografía
115 115 116 117 117 120 120 121 127 127 130
131 133 134 134 135 136 136 137 137 137 140 141
147 149
158
161
167
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ÍNDICE DE FIGURAS CAPÍTULO 1. Introducción Figura 1.1 Distribución del censo mundial ovino por continentes (FAO,
http://faostat.fao.org/faostat/). Figura 1.2 Efectivo ganadero en España por especie (×1.000; MAPA, 2010). Figura 1.3 Distribución del censo ovino por comunidades autónomas en España
(MAPA, 2010). Figura 1.4 Explotación extensiva Figura 1.5 Explotación semi-‐extensiva Figura 1.6 Explotación intensiva Figura 1.7 Distribución de la producción de carne de cordero por comunidades
autónomas (MAPA, 2010) CAPÍTULO 3. Revisión bibliográfica Figura 3.1 Oveja Ripollesa Figura 3.2 Cordero de raza Ripollesa Figura 3.3 Corderos en el aprisco Figura 3.4 Esquema de selección de la raza ovina Ripollesa (modificado por
Casellas (2006) a partir de Guillaumet y Caja (2001)) Figura 3.5 Imagen ecográfica A-‐mode Figura 3.6 Imagen ecográfica B-‐mode Figura 3.7 Imagen ecográfica a tiempo real Figura 3.8 Imagen ecográficas TM-‐mode CAPÍTULO 4. Characterization of the Ripollesa breed Figure 4.1 Grazing ewes of the Ripollesa sheep breed Figure 4.2 Geographical location of the flocks (1: Cal Terrisco flock, 2: SGCE
flock, 3: Las Parras de Martín flock, 4: Mas Muxach flock, 5: Montseny flock, 6: Cal Sabaté flock, 7: Mas Ros flock, 8: SEMEGA flock)
Figure 4.3 Zoometric measures. HL: head length; HW: head width; EL: ear length; CD: chest depth; CW: chest width; CG: chest girth; WH: withers height; HB: height at the middle of the back; BL: body length; RL: rump length; RW: rump width; CC: Cannon circumference
Figure 4.4 Ram of the Ripollesa sheep breed Figure 4.5 Ewe of the Ripollesa sheep breed Figure 4.6 Representation of active individuals on the principal components
(PC1 and PC2) space (1: Cal Terrisco flock, 2: SGCE flock, 3: Las Parras de Martín flock, 4: Mas Muxach flock, 5: Montseny flock, 6: Cal Sabaté flock, 7: Mas Ros flock, 8: SEMEGA flock)
CAPÍTULO 5. Bayesian inference of maternities Figure 5.1 Number of right (black columns) and wrong (white columns) dam
assignments (left Y axis), and their subsequent sensitivity (right Y axis), for single-‐birth abandoned lambs with two single-‐birth candidate dams (scenario SS2). Note that lambs were classified depending on the ewe
5
5
7 12 13 14
17
29 30 31
34 47 48 49 49
69
71
74 78 79
80
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with highest probability of dam assignment (X axis) Figure 5.2 Number of right (black columns) and wrong (white columns) dam
assignments (left Y axis), and their subsequent sensitivity (right Y axis), for abandoned lambs with one single-‐birth and one twin-‐birth candidate dam and without additional abandoned lambs (scenario ST1). Note that lambs were classified depending on the ewe with highest probability of dam assignment (X axis)
Figure 5.3 Number of right (black columns) and wrong (white columns) dam assignments (left Y axis), and their subsequent sensitivity (right Y axis), for single-‐birth (a; scenario ST2S) or twin-‐birth (b; scenario ST2T) abandoned lambs with one single-‐birth and one twin-‐birth candidate dam; both scenarios assumed two abandoned lambs and one mothered lamb. Note that lambs were classified depending on the ewe with highest probability of dam assignment (X axis)
Figure 5.4 Number of right (black columns) and wrong (white columns) dam assignments (left Y axis), and their subsequent sensitivity (right Y axis), for single-‐birth (a; scenario ST3S) or twin-‐birth (b; scenario ST3T) abandoned lambs with one single-‐birth and one twin-‐birth candidate dam when all three lambs were abandoned. Note that lambs were classified depending on the ewe with highest probability of dam assignment (X axis)
Figure 5.5 Number of right (black columns) and wrong (white columns) dam assignments (left Y axis), and their subsequent sensitivity (right Y axis), for abandoned lambs with two twin-‐birth candidate dams when two (a; scenario TT2), three (b; scenario TT3) and all four lambs (c; scenario TT4) were abandoned. Note that lambs were classified depending on the ewe with highest probability of dam assignment (X axis)
CAPÍTULO 6 Real-‐time ultrasounds in light lambs Figure 6.1 Transversal real-‐time ultrasound image Figure 6.2 Longitudinal real-‐time ultrasound image Figure 6.3 Linear regression of ultrasound (US) back-‐fat thickness (independent
variable) against direct carcass back-‐fat thickness (dependent variable) at the thoracic (a) and lumbar (b) levels. Ultrasound images were taken on the Longissimus dorsi muscle, transversal (black) and longitudinal (grey) to the longitudinal axis of the lamb. The discontinuous thin black line characterizes the hypothetical perfect fit (unity line)
Figure 6.4 Linear regression of ultrasound (US) depth of Longissimus dorsi (independent variable) against direct carcass depth of Longissimus dorsi (dependent variable) at the thoracic (a) and lumbar (b) levels. Ultrasound images were taken on the Longissimus dorsi muscle, transversal (black) and longitudinal (grey) to the longitudinal axis of the lamb. The discontinuous thin black line characterizes the hypothetical perfect fit (unity line)
Figure 6.5 Linear regression of ultrasound (US) width of Longissimus dorsi (independent variable) against direct carcass width of Longissimus dorsi (dependent variable) at the thoracic (black) and lumbar (grey) levels.
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123
124
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Ultrasound images were taken on the Longissimus dorsi muscle, transversal to the longitudinal axis of the lamb. The discontinuous thin black line characterizes the hypothetical perfect fit (unity line)
Figure 6.6 Linear regression of ultrasound (US) area of Longissimus dorsi (independent variable) against direct carcass area of Longissimus dorsi (dependent variable) at the thoracic (black) and lumbar (grey) levels. Ultrasound images were taken on the Longissimus dorsi muscle, perpendicular to the longitudinal axis of the lamb. The discontinuous thin black line characterizes the hypothetical perfect fit (unity line)
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ÍNDICE DE TABLAS CAPÍTULO 1 Introducción Tabla 1.1 Censo ganadero en la Unión Europea (miles de animales) en 2009
(FAO, http://faostat.fao.org/faostat/) Tabla 1.2 Distribución del número de ovinos según tipo productivo en las
comunidades autónomas españolas (MAPA, 2010) Tabla 1.3 Estimación del censo ovino Español según raza Tabla 1.4 Censo y producción de carne de ovino en la Unión Europea
(EUROSTAT, http://epp.eurostat.cec.eu.int) CAPÍTULO 3. Revisión bibliográfica Tabla 3.1 Medias reproductivas y productivas de la raza ovina Ripollesa (Caja et al., 2009). Tabla 3.2 Clasificación de la canal ovina respecto a su conformación (Reglamento
CEE 1278/94) Tabla 3.3 Clasificación de las canales ovinas respecto al grado de engrasamiento
(Reglamento CEE 1278/94) Tabla 3.4 Clasificación de las canales ovinas de menos de 13 kg Tabla 3.5 Ácidos grasos mayoritarios en la grasa de origen animal CAPÍTULO 4. Characterization of the Ripollesa breed Table 4.1 Description of the flocks and number of animals measured in each
flock Table 4.2 Definition of the zootechnical indexes calculated for each Ripollesa
individual Table 4.3 Mean, coefficient of variation (C.V.), minimum value (Min.) and
maximum value (Max.) of the morphometrical measures Table 4.4 Average of the morphological measures and live weight of Ripollesa
ewes of flocks of different origins Table 4.5 Eigenvectors from the principal components analysis performed on
the morphological measurements Table 4.6 Number of animals and corresponding percentage classified into each
flock from the discriminant analysis Table 4.7 Means of the variables studied in females of each group of Ripollesa
sheep breed Table 4.8 Mean, standard deviation (S.D.), coefficient of variation (C.V.),
standard error (S.E.), minimum value (Min.) and maximum value (Max.) of the calculated zootechnical indexes
CAPÍTULO 5. Bayesian inference of maternities Table 5.1 Assumed values for systematic effects (excluding year of lambing) and
variance components for the simulation of birth weight data of Ripollesa lambs
Table 5.2 Probability of assignment to the right dam in the different scenarios. Table 5.3 Minimum values of assignment probabilities to reach a sensitivity of
6
8 10
16
31
56
57 58 61
72
75
77
81
82
83
85
87
98 99
xix
99% and 80%, and percentage of assignments agreeing with an 80% sensitivity or larger (Proportion80).
CAPÍTULO 6. Real-‐time ultrasounds in light lambs Table 6.1 Mean, standard deviation (SD), maximum and minimum of transversal
ultrasound measurements at both thoracic and lumbar levels (n = 124; average BW = 26.89 ± 1.24 kg).
Table 6.2 Mean, standard deviation (SD), maximum and minimum of longitudinal ultrasound measurements at both thoracic and lumbar levels (n=124; average BW = 26.89 ± 1.24 kg).
Table 6.3 Mean, standard deviation (SD), maximum and minimum of transversal carcass measurements at both thoracic and lumbar levels (n=124; average BW = 26.89 ± 1.24 kg).
Table 6.4 Correlation coefficient between direct carcass and ultrasound measurements in light lambs.
Table 6.5 Residual decomposition of transversal ultrasound measurements compared to carcass measurements at both thoracic and lumbar levels.
Table 6.6 Residual decomposition of longitudinal ultrasound measurements compared to carcass measurements at both thoracic and lumbar levels.
CAPÍTULO 7. Carcass traits and meat composition in light lambs Table 7.1 Least square means ± SD of transversal ultrasound measurements at
thoracic and lumbar levels for purebred (Ripollesa and Lacaune) and crossbred lambs.
Table 7.2 Least square means ± SD of longitudinal ultrasound measuresments at thoracic and lumbar levels for purebred (Ripollesa and Lacaune) and crossbred lambs.
Table 7.3 Least square means ± SD of direct carcass measurements at thoracic and lumbar levels for purebred (Ripollesa and Lacaune) and crossbred lambs.
Table 7.4 Mean ± SD of weight and carcass traits for purebred (Ripollesa and Lacaune) and crossbred lambs
Table 7.5 Light lambs Longissimus dorsi muscle intramuscular fat, fatty acid composition in percentage by weight of total identified fatty acids, and nutritional ratios
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Capítulo 1
INTRODUCCIÓN
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ts LO� � Aé� f hj � gf � f hz �
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r m 3 � r cTs � P� gPA� f � hj �
� � t� � gf � gh� A� AA�
� s s LO� � áhA� y(éj j � ggP� gz (áéP�
� s ctr T� � f (gzy� g(z f P� zhy� á(f z j �
� OLs � � LO� s � f á(j f h� P(Pj g� Py� z (Aj g�
� R er O� � gPé� g(f zP� áá� g(Pj P�
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r cLs u� t� � (0� áj j A.� � Om Oc�� t� � (0� áj j A)(� �� l L t�C �� er � nc u L t�� �� ncs � r � � OvL � � �
� s c� cs u� u� er � us L � �L O3 � u� Cvl L u� , � L s � 3 r utc�L � � ts u� LOl u� � � � ns uO� OvL � Tc�u �0 �
n cs � � � O� Os L � 3 L t � u� � � Tc�L � O3 ns ct�L � O�� � s Ls � c0� � t c3 OL �c � , � ns � c� nc � � Oc0� L �
� � us � � � s u� ns uO� u� � �L � O� �t s u� �� u � � OvL0� �� � s LNs c3 �� OvL � , � � s 3 nsuO� OvL � � � �� � �L � � , �
� � � � � �c L 0� n�c �� ns � c� s Tc�c � t L c� ncs � r � ts u� � NOLO� s u� , � � �c �� t cOx�� s u� n�c �� r L �� 3 Cs c�
�� nt� � OvL � , � l � s c�� OvL � ns c� n�c t � � � s u� � s Lur 3 O� s c u(� �
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l �L � 3 s � ONO� � L � s 0� s u� ncs � r � ts c u� u � l L � s � OT�� s u� �� �� �n t�c u � �� uts u� � �3 � Osu� , � � r 3 n Oc�
� s L � � u� mn � t�t Ol �u (� � ut�� uOtr �� OvL � R�� R � Rs � er � s u� ncs � r � ts c u� u � OLt c u L � 3 �u � ns c� �u �
� � c� � t cDutO� � u� � � �� � �L � 0� � � �� � � cL � , � ur � nc � O� tO� O O� �� � L � �L O3 � u� l Ol s u(� � �
r tc�u s LO� s � �� tO 3 ns � c � � u� r L �� tI � LO� �� L s � � utcr � tOl �� � s L � �� er � u � nr � � � r �L tONO� �c � �
� u�c cs s � 3 r u� r �c � , � � � s u� t CO� s u� Tc�u s u� L � � � LO3 � � L � l Ol s 0� �� 3 �u � � � nc � � Oc� �u �
� � c� � t cDutO� � u� � � �� � �L � � 1� O � 0� gPyj )(� � � r us � � � �� � s Tc�ND�� n�c �� utO3 �c � � Tcs us c� � � � �
Tc� u� � ur � � r t�L �� , � � �c �� � � 3 au� r s � � s cu� � ut�� � O L � � s � r 3 L t�� s � us � c � ts � s � L �
l �� r L s � � � � �c L � , � ns c� OLsu� 1� ccOLT� t� � (0� gPPh.� � s r TRts L � , � � r c OLTts L0� gPPá)� , �
� s LutOtr , � r L �� R cc�3 O L t�� O3 ns ct�L t � n�c �� � 3 Cs c�3 O L ts � T L I tO� s � � � �u �
� � c� � t cDutO� �u � � � �� � �L � � L � ut�u � un � O u(� � r Ler � L � s l OLsu� s u� r tc�u s LO� s u� R� L �
t LO� s � r L � r us � O3 Ot�� s � 1� s nÓOLu0� gPPj )0� � � cD�L � sNc � c� � u� 3 Ou3 �u � s ns ctr LO� �� u� n�c ��
� r � L tONO� �c � �� � s LNs c3 �� OvL � �L O3 � � � � r L �� 3 �L c�� s � C tOl �( � �u � l L t�C �u � � � ut � 3 I ts � s �
u � � �u � L � ncOL � On� 3 L t � L � er � u � �n O� �� L � �L O3 � u� l Ol s u0� s u� � �c �� t c u� 3 � O� s u�
tO L L � r L �� 3 s � c�� �� �� � t�� R c � �� O O� �� � , � r L � � s ut � c �t Ol �3 L t � � �C s � L � � s 3 n�c �� OvL �
� s L � s tcs u� 3 I ts � s u� n�c �� nc � � Oc� �� � s 3 nsuO� OvL � � � �� � �L � 0� � s 3 s � �� ts 3 s Tc�ND��
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� � I ut�� t � L s s TD�� R�� �, r � �� s � �� s u� ncs � r � ts c u� �� s � t L c� �L O3 � u� � s L � 3 �, s c� � �L tO� �� �
� � 3 au� r s � , � 3 L s u� Tc�u � � 1� � Rp i c c� t� � (0� gPPy.� � p � s 3 � t� � (0� áj j á)(�
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�T cs n � r � cO�u 0� ns c� � cD�� � s L l L � Os L � � s � 3 � O�L t � OLT LO cD�� T L I tO� �0 � � s LutOtr , � r L �
n� tcO3 s LOs � 3 r L � O� � � � l � s c� OL � � � r �� (� �� � Ol cuO� �� � T L I tO� �� er � u � L � r L tc��
�� tr � 3 L t � L � �u � c�x �u � � � �L O3 � u� � s 3 I utO� s u� n c3 Ot � �� s u� T�L �� cs u� u � � Os L �c �
ns � �� Os L u� r � s � t L c� L r l �u � c�x �u � er � c uns L � � L � �� s u� � �3 � Osu� � � 3 � Os � �3 � O L t 0� ��
s u� L r l s u� � s Ls � O3 O L ts u� us � c � �u � L � uO� �� u� � � L r tcO� OvL � Rr 3 �L �0 � �� �u � �3 L �x �u � � �
LN c3 � �� u0� �� �� uOtr �� OvL � � � 3 c� �� s � , � �� �u � L � uO� �� u� us � O� u� 1 � � 0� áj j é)(� � �
OL � c 3 L ts � � � �� ns � �� OvL � Rr 3 �L � � , � � � � c � O L t � � 3 �L � �� � � � O3 L ts u� t�L ts � � � s cOT L �
l T t� 0� � s 3 s � � � s cOT L � �L O3 � 0� R�� R � Rs � er � ns � s � �� ns � s � s u� T�L �� cs u� l �, � L � s nt�L � s �
ns c� OLt LuONO� � c� ur u� mn s t�� Os L u� � OLtcs � r � Oc� c�x �u � L r l �u � , � un � DNO� �u � n�c �� � � � � � tOns � � �
ncs � r � � OvL(� � ut�� u � � OvL � R�� ncs l s � �� s � r L � � � c 3 L ts � L � � r us � � � c�x �u � L s �
un � O� Ox� � �u 0� ns LO L � s � L � n OTcs � ur � mOut L � O�� , � ns c� L � � �� mOut L � O�� � � �u � c�x �u �
�r tv � ts L � u� 1 �c � D� 0� gPhj .� � � L � cs Ó0� gPPj )� � s 3 s � �� � Ons u�( � �u � c�x �u � � � T�L �� s �
�r tv � ts L �u � ns u L � c� uTsu� l � Osusu� � s 3 s � �� c uOut L � O�� �� � O ct�u � LN c3 � �� u0� Tc�L �
N ctO O� �� 0� � r L �u � � r � O� �� u� 3 �t cL �u 0� �t cO� r ts u� aLO� s u� � � s u� ncs � r � ts u0� s LT l O� �� � , �
�� �n t�� OvL � � � 3 � Os 0� L tc � s tc�u (� � �c �� l Ot�c � �� nI c� O� �� � � c � r cusu� T L I tO� s u� �L O3 � u0� � �
� � � c � s 3 O L � �� ut�� � c� ncs Tc� 3 � u� � � � s Lu cl �� OvL � , � 3 �L t LO3 O L ts � T L I tO� s � �� LOl �
3 r L � O� (� �u � � OutOLt�u � N� u u� � � ncs Tc�3 � � � s cc uns L � L � �� �� � u� cOn� OvL � T L c� � � � ��
ns � �� OvL0� �� ur � l x� � Ol O� O� � � L � � ON c L t u� �n �c t�� s u� L tc � s u� er � u � L � r L tc�� ��
� u� cOn� OvL � � � �u � � �c �� t cDutO� �u � c� � O� u� , � ncs � r � tOl �u � � � OLt cI u0� � � c� � t cOx� � OvL � c�� O� 0�
ncs Tc�3 �� � � � s Lu cl �� OvL � T L I tO� �� � � � � � � � 0� ncs Tc�3 �� � � � s Lu cl �� OvL � T L I tO� �� � � � � � � � � , �
ncs Tc�3 �� � � 3 Cs c�� T L I tO� �0 � n�c �� � � r � � u� L � u�c Os � � s Ls � c� �u � � �c �� t cDutO� �u � � � ��
c�x �� n�c �� ns � c� c � Ox�c � r L �� u � � OvL � �� � r �� � � � � s u� �L O3 � u� c ncs � r � ts c u� , � � �
c ns uO� OvL0� � n L � O L � s � � � s u� s � C tOl s u� � � u � � OvL(�
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OLO� O� tOl �u � n� c�� l Ot� c� � � � �O3 O L ts � � � �� �� tOl O� �� � � � �u � mn s t�� Os L u0� � s t�L � s �u � � �
r L � � utcr � tr c�� ncs � r � tOl �� l O�� � , � �u Tr c�L � s � �� ur � l x� � c l s � T L c�� Os L � (� � L � ut �
u L tO� s 0� �u � �u s � O�� Os L u� T�L �� c�u � us L � r L � l RD� r s � O� vL s � n�c �� �� s nt�c � ut � tOns � � �
OLO� O� tOl �u 0� �u � � r � u� u � l L � OL � L tOl �� � u� ns c� s u� ncs Tc�3 �u � � � �, r � �u � ut�� � O� s u� ns c� �
� � � � � c ts � gj zGáj j h� 1� � � 0� áj j h)� , � ns ut cOs c u(� � L tc � s u� c er OuOtsu� ut�� er � �u �
�T cr n�� Os L u� � s LutOtr O� �u � � � c�L � � s L t�c � � s L � � O3 LuOvL � ur NO� O L t � n�c �� s c� L �c � �� s N ct��
� s L � l Out�u � �� �� 3 Cs c�� � � �� � s 3 c� O� Ox�� OvL � , � �� � � c � r � � OvL � � � � s ut u0� , � L � ur � � �u s 0� �� ��
ncs 3 s � OvL � , � r us � � � � r � � s � � � s u� c � r cusu� L �t r c� u� 3 � O�L t � uOut 3 �u � � � n�u ts c s �
tc� � O� Os L � (� � �3 � OI L � u � uO L t�L � �u � � � u u� n�c �� �� s nt�c � �u � �� � Os L u� 3 �u � �� � r �� �u � � s L �
l Out�u � �� � � � s L � L tc�� OvL � 3 nc u�c O� 0� � s L � �� OLtcs � r � � OvL � � � � cOt cOsu� 3 s � cLs u� � �
s cT�L Ox�� OvL � , � T utOvL0� �u D� � s 3 s � n�c �� T�c �L tOx�c � �� �� � r �� �� Ns c3 �� OvL � � � s u� tOtr �c u� � �
� u� mn s t�� Os L u� , � �� un � O� Ox�� OvL � � � �u � ncs � r � � Os L u� �� tc�l I u� � � �u � nc�� tO� �u � � �
tOnONO� �� OvL0� � � c�� t cOx�� OvL � � � s u� ncs � r � ts u� , � � � � � ncs � r � � OvL � s cO L t�� �� � � 3 c� �� s (�
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�� T�L �� cD� � � �u �� �� L � �u � c�x �u � �r tv � ts L �u 0� � s LutOtr , � r L �� ncs � r � � OvL � 3 r , �
l � Osu�� � u� � 3 r � Rs u� nr Lts u� � � l Out�0 � t�L ts � � s Lv3 O� s u� � s 3 s � us � O� u� , �
3 � Os �3 � O L t� u(� �� ncOL � On� � O3 ns ct� L � O�� � � �u � c�x �u � �r tv � ts L �u � u� er � us L � r L � c � r cus �
T L I tO� s � Nr L � �3 L t� � er � u� L � u�c Os � � s Lu cl �c 0� uOL � OTLs c�c � ur � Tc�� s � � � �� �n t�� OvL � ��
� u� � s L � O� Os L u� 3 � Os �3 � O L t� u(� � s c� s 0� � 3 �L t LO3 O L ts � � � �� � �� � o�� T�L �� c��
�r tv � ts L � � u� l Ot� � n� c�� �� � s Lu cl �� OvL � � � � s uOut 3 �u � � � n� uts u0� er � � � s tc�� 3 �L c�� u �
� Tc�� �c D� L � , � u � n c� cD� L (� � �3 � OI L � u� r L �� R cc�3 O L t�� n�c �� �� NOC�� OvL � � � �� ns � �� OvL � � �
3 � Os � cr c� � � � �3 n O�u � xs L � u0� ns c� s � er � u� O3 ns ct�L t � �, r � �c � �� er � u � ncs � r x� �L �
3 Cs c�u � L � �u � � s L � O� Os L u� � � l O� �0 � ns c� 3 � Os � � � �� 3 Cs c�� T L I tO� �� � � � � c�x �� , � tI � LO� �u �
� � 3 �L Cs .� � � ut� � 3 �L c�� u � nr � � � s Lu Tr Oc� OL � c 3 L t�c � �� � � O� �� � � � s u� ncs � r � ts u� , �
ns c� s � t�L ts � � � s 3 n tOtOl O� �� (
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Capítulo 2
OBJETIVOS
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Capítulo 3
REVISIÓN BIBLIOGRÁFICA
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Capítulo 4
CHARACTERIZATION OF THE RIPOLLESA BREED
CHARACTERIZATION OF THE RIPOLLESA BREED
67
Morphologic analysis and subpopulation characterization of Ripollesa
sheep breed
Introduction
Livestock farming based on local breeds constitutes a very valuable animal
industry from the economic, social and environmental points of view. Local breeds have
remarkable special characteristics like resistance to prevailing diseases, fertility, maternal
ability, longevity, adaptation to the environment and unique attributes of their final
products, among others (García, 1980). Nevertheless, the increasing demand for animal
products has led to an intensification of production systems and the subsequent
restriction of the livestock industry to a few specialized breeds. This practice has reduced
the use of local breeds and put their survival in danger (Oldenbroek, 1999), which is also
the case of the Ripollesa breed. To overcome this problem, the Food and Agriculture
Organization (FAO; http://www.fao.org) of the United Nations recommended establishing
conservation programmes for the maintenance of animal genetic resources. Note that
these programmes include, among other actions, the characterization of these local
breeds.
Most of the native breeds of Spain included in the inventory of Spanish livestock
breeds belong to the ovine species (Spanish Real Decreto 2129/2008,
http://www.boe.es/boe/dias/2009/01/27/pdfs/BOE-‐A-‐2009-‐1312.pdf). The Ripollesa is
the most abundant local sheep breed in Catalonia (Milán, Arnalte and Caja, 2003; Caja et
al., 2010), an Autonomous Community located in the NE of Spain. The breed is usually
considered as being moderately prolific and is exploited for the production of “pascual”-‐
type lambs (Guillaumet and Caja, 2001; Caja et al., 2010). Indeed, previous studies
reported both appealing productive (Torre et al., 1989; Torre, 1991; Casellas et al., 2007c)
and reproductive (Casellas et al., 2007a, 2007b) performance within the context of a
remarkable adaptability to the geoclimatic conditions of the Mediterranean area
(Guillaumet and Caja, 2001). Nevertheless, additional efforts to typify this breed become
CHARACTERIZATION OF THE RIPOLLESA BREED
68
necessary, given that there is a substantial phenotypic heterogeneity that could even lead
to different subpopulations (Torre, 1991).
Although the existence of different Ripollesa subpopulations was advocated by
stakeholders for decades and suggested by different authors (Torre, 1991; Guillaumet and
Caja, 2001), this hypothesis has never been tested. If true, this breed could suffer from a
severe genetic structure (bottleneck) with important consequences on its effective
population size and, by extension, on its conservation or even selection programme.
These hypothetical subpopulations may have originated in different geographic areas and
could differ in both, production and morphological traits, the latter being the main
objective of our study.
In the present work, an attempt has been made to characterize the morphologic
diversity of the Ripollesa sheep breed, both between and within flocks. This research was
performed on the basis of standardized morphological measurements and the
subsequent calculation of zootechnical (ethnological and functional) indexes.
Material and Methods
The Ripollesa breed
The Ripollesa sheep were described as a medium-‐sized breed with convex profile
and characteristic pigmentation with black or brown spots on the head and legs (Sánchez-‐
Belda and Sánchez-‐Trujillano, 1986; Guillaumet and Caja, 2001; Caja et al., 2010). This
breed belongs to the Spanish medium-‐fine (i.e., “entrefino”) wool-‐type trunk with white
wool and packed fleece (wool fibre diameter, 23 to 26 μm; Sánchez-‐Belda and Sánchez-‐
Trujillano, 1986) (Figure 4.1). The breed is characteristically Mediterranean exploited
under semi-‐extensive production systems in the mountains and cultivated plains of
Catalonia and currently intended for meat production (Guillaumet and Caja, 2001; Caja et
� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �
� ) : � �� �
�P � n� 1Gí Gpd� � L � v� �i H� vBPC� TBi � vP�s EI eHi � � v� l 6�v � s �P uhek6H� ,11� eB� 18� FE� PZtHho HZEI eg� o ZeI �
�� Bs e� í 1� FE� � �i � �v v� o HZEI epn� s vs �P Pk� CZiH� ePk� � k� eI H� vI HH6� Bo SHiv� eB� eI H� PB� �P � � s e� I Hi vd� �
�
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�
�
� Bve� � Z6BPPHv�� TPB� Fv� �i H� PB� �e HC� ZS � eI H� 6iBtZS� Hv� BT� � �i � HPBS�� �S C� � ZiBS �� , � Bi iHn�
í : : í g� � ZP�S � He� �P � n� 1GG9g� � � �� He� � P� n� 1Gí Gpn� � BtHiZSE� �� o ZCH� i �S EH� BT� HStZiBSL HSev� TiBL �
6P�Z Sv� eB� � B� ve�P � L Bs Se�Z Svd� � I Zv� I HeHiBEHSHZekn� �S C� HtHS� EHBEi �6 I Z� � ZvBP�e ZBSn� L �k � I �t H�
PHC� eB� eI H� HtBPseZBS� BT� eI H� � i HHC� eBo �i Cv� CZTTHiHSe� vs � 6B6sP�e ZBSvd� � PeI BsEI � eI Zv� FZSC� BT�
6B6sP�e ZBS� vei s � es iH� I �v � SHtHi � � HHS� 6iB6HiPk� vesCZHC� ZS � eI H� � Z6BPPHv�� � i HHCn� 6s e�e ZtH�
vs � 6B6sP�e ZBSv� I �t H� � HHS� vs EEHveHC� �� � Bi CZSE� eB� EHBEi �6 I Z� � PB� �e ZBSv� , � Bi iHn� í : : í g�
� s ZPP�s L He� �S C� � � � n� 1GGí g� � Bi iHvn� 1GGzpd� �
�
� ZTTHiHSe� S �L Hv� BT� eI HvH� vs � 6B6sP�e ZBSv� �i H� CHiZtHC� TiBL � 6�i eZ� s P�i � PB� �P ZeZHv� ZS �
eI H� � i H�n � eI HZi � L �Z S � TH�e s iHv� � HZSE� eI H� TBPPBo ZSE� , � �� PH� 8dí pb� �
�
,í p � BvBPHS� � n� TiBL � � DvBP� , � �i � HPBS�p n� I Bi SHC� Ho Hv� o ZeI � P�i EH� � BCk� vZfHd�
,1p Es � P� CZS � n� TiBL � Es �P �C �� , � �i � HPBS�p n� SBShI BiSHC� Ho Hv� o ZeI � v� �i � H� o BBP� BS � eI H�
� HPPkd�
,9p Ps � SHS� � n� TiBL � eI H� Ps �S Av� , � �i � HPBS�p n� L HCZs L � � BCk� vZfHd�
CHARACTERIZATION OF THE RIPOLLESA BREED
70
(4) Queralpina, from Queralbs (Girona), smaller and less prolific sheep with longer and
the finest wool fibres.
(5) Hilarenca, from Sant Hilari Sacalm (Girona), the largest rams with long horns and
presence of abundant hair in the wool in rams and ewes. Lambs are markedly hairy
at birth.
Main differences among subpopulations should be expected with regard to body
size and characteristics of wool and horns (Guillaumet and Caja, 2001).
Sheep samples
Morphological measurements were taken between September 2007 and June
2008, on adult Ripollesa animals (more than two years old) belonging to eight flocks
registered in the official Flock-‐Book of the breed. Flocks were distributed across the
geographic areas where the Ripollesa sheep are located (Figure 4.2), providing samples
from the subpopulations described by Torre (1991) (Table 4.1).
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� z í � �� �
� � � � � � � M� � � � HBEi �6 I Z� � P� PB� �e ZBS� BT� eI H� TPB� Fv� ,í b� � �P � � Hi i Zv� B� TPB� Fn� 1b� � � � � � TPB� Fn� 9b� �v �� �i i �v � CH� � �i eOS� TPB� Fn� 8b� � � v� � s c�� I � TPB� Fn� 0b� � BSevHSk� TPB� Fn� ) b� � �P � � �� � eR� TPB� Fn� zb�� �v � � Bv� TPB� Fn� ; b� � � � � � � � TPB� Fpd�
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o I HiH�v � TPB� Fv� z � � SC� ; � o HiH� Hc6HiZL HSe�P � T�i L v� s SCHi � ZSeHSvZtH� � BSCZeZBSvd� � I HH6�
v� L 6PHC� TiBL � H�� I � TPB� F� � BSvZveHC� BT� 18� eB� 9G� SBSh6iHES�S e� Ho Hv� �S C� G� eB� 8� i �L v� e�F HS�
�e � i �S CBL � TiBL � eI H� BPCHiheI � Sheo Bhki � � i HHCZSE� veB� Fg� SBeH� eI �e � i �L v� o HiH� 6iBCs � HC� ZS �
eI H� v�L H� TPB� F� Bi � �� 7s ZiHC� TiBL � eI H� SHZEI � Bs i � TPB� Fvd� � � eBe�P � BT� 18í � vI HH6� ,118� Ho Hv� �S C�
í z � i �L vp� o HiH� ZS � PsCHC� ZS � eI H� vesCkn� �v � CHe�Z PHC� ZS � � �� PH� 8dí d� �
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CHARACTERIZATION OF THE RIPOLLESA BREED
72
CHARACTERIZATION OF THE RIPOLLESA BREED
73
Morphological measurements
Morphological variables were measured by the same trained operator in the
mornings, before the animals left the shelter to graze to avoid undesirable variations due
to changes in live-‐weight and rumen volumes. On the basis of Aparicio (1944), the
following 12 morphological measures were taken on each animal (Figure 4.3):
(1) head length (HL), frontal distance from mouth to poll,
(2) head width (HW), maximum distance between zygomatic arches,
(3) ear length (EL), distance from the base to the tip of the right ear, along the dorsal
surface,
(4) chest depth (CD), vertical distance from the top of the withers to the xyfoid process
of the sternum,
(5) chest width (CW), maximum intercostal diameter at the level of the 6th rib, just
behind the elbows,
(6) chest girth (CG), perimeter of the chest at the level of the 6th rib,
(7) withers height (WH), height from the top of the withers to the ground,
(8) back height (BH), height at the middle of the back, between the thoracic and the
lumbar vertebrae,
(9) body length (BL), distance from the manubrium of the sternum to the pin,
(10) rump length (RL), distance from hip to pin,
(11) rump width (RW) maximum distance between left and right hurls,
(12) cannon perimeter (CP), perimeter of the right foreleg, between the knee and the
pastern.
These variables were obtained using a ribbon measuring tape and a measuring
stick for horses (Hauptner and Herberholz, Solingen, Germany). Additionally, live weight
was recorded using fixed (SR2000, Tru-‐Test, Auckland, New Zealand) or portable FX1
(Iconix NZ, Oamaru, New Zealand) electronic scales.
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CHARACTERIZATION OF THE RIPOLLESA BREED
75
Table 4.2 Definition of the zootechnical indexes calculated for each Ripollesa individual.
Indices Type † Calculation
Cephalic Ethnological HW / HL x 100
Thoracic Ethnological CW / CD x 100
Pelvic Ethnological RW / RL x 100
Corporal Ethnological BL / CG x 100
Dactyl-‐Thoracic Functional CC / CG x 100
Dactyl-‐Costal Functional CC / CW x 100
Relative Depth of Thorax Functional CD / WH x 100
Transversal-‐Pelvic Functional RW / WH x 100
Longitudinal-‐Pelvic Functional RL / WH x 100
Balance Functional (RW x RL) / (CD x CW)
Length Functional BL / WH
Cumulative
Functional
(W / �) + Balance
Index + Length Index
HL: Head length; HW: Head width; EL: Ear length; CD: Chest depth; CW: Chest width; CG: Chest girth; WH: Withers height; HB: Height at the middle of the back; BL: Body length; RL: Rump length; RW: Rump width; CC: Cannon perimeter; W: Body weight; �: estimated average for W. † Ethnological indexes contributed general information about breed characteristics whereas functional indexes contributed information about the type, purpose and performance of the breed.
Statistical analyses
All measurements and indexes were separately analyzed under the following
linear model,
ijkjiijk SFY +++=
where ijkY was the dependent phenotypic record, was the population mean, iF was
the effect of the flock (1-‐8), jS was the effect of the sex (1 or 2), and ijk was the residual
term. Flock effects were contrasted by a one-‐way ANOVA test followed by the Student
Newman-‐Keuls multiple comparison test, implemented in the General Linear Model
procedure of SAS 9.1 (SAS Institute, Inc., Cary, NC, USA).
CHARACTERIZATION OF THE RIPOLLESA BREED
76
The residuals obtained from the previous analyses on morphological
measurements were used for a principal component analysis (PCA) performed by the
SPAD 5.5 (2002) software (Coheris, Suresnes, France). For all subsequent analysis, only
data from ewes were used due to the small number of rams contributing to this study.
This analysis allowed us to understand the differences among flocks with regard to
morphological measures, where only principal components accounting for more than 10
percent of the phenotypic variance were retained.
On the basis of the estimates from the PCA analysis, a discriminant analysis (SPAD
5.5) was conducted to estimate the proportion of animals that were properly classified
into their own flock. This was an additional way to evaluate within-‐flock resemblance and
the degree of differences between flocks. As a final step, a cluster analysis (SPAD 5.5) was
performed for classification of data to establish the optimum number of groups (i.e.,
subpopulations) based on the number of principal components accounting for at least 80
percent of the total inertia. Between-‐group differences of morphological measures were
compared with a Student Newman-‐Keuls multiple comparison test (SAS 9.1).
Results and Discussion
Morphological diversity between sexes and flocks
According to the values reported in this study, the Ripollesa breed showed a
marked sexual dimorphism, as rams (Figure 4.4) were larger than ewes (Figure 4.5) for
almost all measurements (P < 0.05), except ear length and rump width (Table 4.3). Sexual
dimorphism is a fundamental morphological characteristic of most ungulates (Andersson,
1994), and has important consequences for ecology, behaviour, population dynamics and
evolution (LeBlanc et al., 2001).
CHARACTERIZATION OF THE RIPOLLESA BREED
77
Table 4.3 Mean, coefficient of variation (C.V.), minimum value (Min.) and maximum value (Max.) of the morphometrical measures.
Variables Ewes Rams
Mean C.V. Min. Max. Mean
Head length, cm 23.7a 4.4 21 28 26.8b
Head width, cm 13.8a 4.5 11 17 15.4b
Ear length, cm 13.5a 7.7 10 16 13.2a
Chest depth, cm 30.7a 6.9 20 42 34.1b
Chest width, cm 18.8a 10.0 12 28 21.2b
Chest girth, cm 90.4a 5.1 75 108 100.9b
Withers height, cm 69.4a 4.4 58 80 75.9b
Height at the middle of the back, cm 68.2a 4.3 56 77 74.5b
Body length, cm 75.6a 5.3 43 85 84.1b
Rump length, cm 24.2a 6.9 15 29 26.5b
Rump width, cm 21.8a 6.5 16 29 22.5b
Cannon perimeter, cm 8.6a 6.1 7.5 10 10.0b
Weight, kg 51.4a 12.9 25 80 75.1b
Means with the same superscript did not differ significantly (P > 0.05) between sexes.
It is expected that all animal populations have certain variability between groups
or subpopulations, these differences being of great importance to improve performance
by selection. In our study, a moderate variability (CV ranging between 4 and 13 percent)
in all the morphological measures considered was observed for the whole ewe sample
(Table 4.3). The highest CVs were for live weight (12.9 percent), chest width
(10.0percent), ear length (7.7 percent), chest depth (6.9 percent), rump length (6.9
percent) and rump width (6.5 percent). Despite this, the greatest differences between
average estimates for each flock with respect to the overall means were reported for
withers height (65.3 to 73.2 cm; 11.4 percent), chest depth (26.6 to 32.9 cm; 20.5
percent), rump width (19.8 to 22.9 cm; 14.2 percent), head width (12.5 to 14.7 cm; 15.9
percent) and ear length (12.5 to 14.5 cm; 14.8 percent). These differences could be
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CHARACTERIZATION OF THE RIPOLLESA BREED
81
CHARACTERIZATION OF THE RIPOLLESA BREED
82
Table 4.5 Eigenvectors from the principal components analysis performed on the morphological measurements.
PC 1† PC 2
Head length, cm 0.26 0.09
Head width, cm 0.22 0.25
Ear length, cm 0.15 0.22
Chest depth, cm 0.25 -‐0.03
Chest width, cm 0.24 -‐0.42
Chest girth, cm 0.40 -‐0.40
Withers height, cm 0.33 0.44
Height at the middle of the back, cm 0.33 0.46
Body length, cm 0.28 -‐0.06
Rump length, cm 0.27 -‐0.03
Rump width, cm 0.26 0.05
Cannon perimeter, cm 0.27 -‐0.11
Live weight, kg 0.36 -‐0.35
† PC: Principal component; PC1 accounted for 47.6 percent of the variance and PC2 accounted for 12.3 percent of the variance.
Discriminant and cluster analyses
The results from the discriminant analysis (Table 4.6) confirmed this partial
overlapping as the percentage of individuals correctly classified into their own flock
ranged between 70 and 100 percent, with only 18.7 percent of the animals, on average,
being classified in wrong flocks. The high percentages of matching suggested a marked
degree of differentiation among flocks, being the greatest in the SEMEGA (#7) flock in
which 100 percent of their own ewes were assigned correctly and only 2 ewes more from
other flocks were incorrectly assigned to this flock.
CHARACTERIZATION OF THE RIPOLLESA BREED
83
CHARACTERIZATION OF THE RIPOLLESA BREED
84
On the other hand, the cluster analysis defined four sheep groups, each
represented mainly by the following flocks (number of ewes and percentage of
matching):
(1) Group 1 (19.6 percent of the ewes): Mas Muxach (24 ewes, 89 percent) and
SGCE (13 ewes, 45 percent), characterized by being heavy and large framed
animals.
(2) Group 2 (17.9 percent of the ewes): Montseny (13 ewes, 54 percent) and Las
Parras de Martín (13 ewes, 45 percent), characterized mainly by being tall
animals.
(3) Group 3 (40.6 percent): Cal Sabaté (20 ewes, 67 percent), Cal Terrisco (20 ewes,
67 percent) and Mas Ros (14 ewes, 52 percent), characterized by including
animals with intermediate frame and long ears.
(4) Group 4 (21.9 percent): SEMEGA (24 animals, 86 percent), with the smallest
body size (Table 4.7).
This grouping can be justified in part by the different subpopulations that
contributed to the studied flocks. Although the Montseny and Las Parras de Martín flocks
must have come from the same original subpopulations as do Mas Muxach and SGCE
(Hilarenca and Igualadina subpopulations), they are characterized by having a greater
height than the general average. Even assuming a common origin, departures between
Group 1 and Group 2 must be probably linked to variable contributions from the
Hilarenca and Igualadina historical subpopulations and an independent evolution during
several years or decades. Focusing on Group 3, it was expected that animals from Cal
Sabaté flock had a medium size, since this flock derived from original Hilarenca and
Queralpina subpopulations, although differing from flocks assigned to Groups 1 and 2; in
a similar way, the Cal Terrisco flock (Lluçanenca subpopulation) and Mas Ros flock
(Queralpina subpopulation) were assigned to this Group 3. Nevertheless, there are also
animals from the original Queralpina subpopulation in the group of SEMEGA ewes, the
smaller individuals of the Ripollesa breed reported.
CHARACTERIZATION OF THE RIPOLLESA BREED
85
Within this context, the morphological characteristics of the historical
subpopulations are still present in current morphological groups, although influences
from more than one subpopulation could be found in the same flock and independent
evolutionary patterns would have led to different subpopulations although departing
from the same original genetic background. This makes evident the moderate to low
genetic flow between flocks, where some specific morphotypes were preferentially
preserved in each flock. In consequence, a substantial genetic structure could be
anticipated in the Ripollesa populations, increasing the complexity of a conservation
program.
Table 4.7 Means of the variables studied in females of each group of Ripollesa sheep breed.
Group 1 Group 2 Group 3 Group 4
Number of animals 56 53 87 28
Head length, cm 23.6a 24.9b 23.3a 22.2c
Head width, cm 14.2a 13.9a 13.9a 12.5b
Ear length, cm 13.2a 14.0b 13.7b 13.0a
Chest depth, cm 32.4a 32.3a 29.8b 26.6c
Chest width, cm 21.5a 18.8b 17.5c 17.3c
Chest girth, cm 97.7a 89.8b 88.6b 82.7c
Withers height, cm 70.0a 71.9b 68.8a 65.3c
Height at the middle of the back, cm 68.7a 70.5b 67.8a 64.3c
Body length, cm 76.5ab 78.0a 75.1b 71.1c
Rump length, cm 25.1a 25.7a 24.1b 19.9c
Rump width, cm 23.4a 21.4b 21.6b 19.8c
Cannon perimeter, cm 8.8a 8.7ab 8.5bc 8.4c
Weight, kg 59.9a 54.9b 48.2c 37.9d
Means with the same superscript did not differ significantly (P > 0.05). Group 1: Mas Muxach and SGCE flocks; Group 2: Montseny and Las Parras de Martín flocks; Group 3: Cal Sabaté, Cal Terrisco and Mas Ros flocks; Group 4: SEMEGA flock.
CHARACTERIZATION OF THE RIPOLLESA BREED
86
Differences among flocks were also found by Avellanet (2006) in ewes of the
Xisqueta breed, where the values of the morphological measures varied depending upon
the location of the flocks, management, and feeding, among others. Also Kunene et al.
(2007) found that the location of the flocks had a significant effect on the morphological
measures of Zulu ewes. In this study, however, it is possible that the differences among
flocks could also have a genetic origin because some of the differences among the
hypothetical subpopulations of origin are retained when the flocks are reared in other
areas.
Zootechnical indexes
The calculation and analysis of the different zootechnical indexes allowed us to
ethnologically classify the Ripollesa sheep breed (Table 4.8). On the basis of the cephalic
index, the Ripollesa showed to be a dolichocephalic breed, since the length of the head
predominated over the width. Both the thoracic and the length indexes allowed us to
classify the breed as long-‐shaped, whereas the pelvic index indicated that it is a convex
breed, with the rump length predominating in relation to its amplitude. The breed had a
medium frame according to the dactyl-‐thoracic and dactyl-‐costal indexes. These results
agreed with the visual valuations of Torre (1991) and Guillaumet and Caja (2001). The
relative depth of the thoracic index indicated that the breed is suitable for meat
production, lower values indicating better meat aptitude (Aparicio, 1944), in agreement
with Daza (1997), Guillaumet and Caja (2001) and Milan et al. (2003). This index also
indicated the length of the legs suggests that the breed had good adaptation to the
environmental conditions under which it is raised. The transversal pelvic and longitudinal
pelvic indexes also provided information about the aptitude of the animal, supporting the
Ripollesa as a sheep breed for meat production, which reinforces the results obtained
from the calculation of the cumulative index.
Given that some of the indexes gave the same information, we suggest simplifying
the number of analyzed indexes for the future to the following ones: thoracic, cephalic,
pelvic, dactyl-‐thoracic, and relative depth of thorax. These subsets of indexes are enough
CHARACTERIZATION OF THE RIPOLLESA BREED
87
to ethnologically classify the breed and also to provide information about the productive
purposes of the Ripollesa breed.
Average differences between flocks and subpopulations of Ripollesa relied on
sheep size, weight and height. The cluster analysis suggested four differentiated groups,
although there were ewes that could not be assigned to a particular group. The results of
the analyzed indexes allowed us to classify the Ripollesa sheep breed as medium-‐framed
and convex, predominantly suitable for meat production.
Table 4.8 Mean, standard deviation (S.D.), coefficient of variation (C.V.), standard error (S.E.), minimum value (Min.) and maximum value (Max.) of the calculated zootechnical indexes.
Indices Mean S.D. C.V. S.E. Min. Max.
Cephalic 58.4 3.7 6.3 0.2 48.0 76.2
Thoracic 61.5 8.1 13.2 0.5 41.4 87.1
Pelvic 90.8 9.8 10.8 0.5 73.1 129.4
Corporal 83.9 6.1 7.3 0.3 47.8 101.2
Dactyl-‐Thoracic 9.6 0.7 7.3 0.0 7.8 11.7
Dactyl-‐Costal 46.6 6.1 13.0 0.3 30.8 69.2
Relative depth of Thorax 44.2 3.7 8.3 0.2 29.4 60.3
Transversal-‐Pelvic 31.4 2.5 8.0 0.1 26.4 38.7
Longitudinal-‐Pelvic 34.9 3.1 8.9 0.2 23.8 43.3
Balance 0.9 0.1 15.6 0.0 0.5 1.4
Length 1.1 0.1 5.8 0.0 0.6 1.3
Cumulative 3.0 0.2 7.2 0.0 2.0 3.6
Note that our results contribute essential information to characterize this meat-‐
type breed following FAO recommendations, and has become a relevant source of basic
information to support the conservation and selection program of the Ripollesa sheep
breed.
Capítulo 5
BAYESIAN INFERENCE OF MATERNITIES
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Capítulo 6
REAL-‐TIME ULTRASOUNDS IN LIGHT LAMBS
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ñqqRB� �s � � cm�d l � i � � � �l � geds � � � c� 1s � � � 1l � i i �s � �s � �i � � c2� � 1l � i � 1b b � � 1� l � mg� �� l � i �
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i � i � cc1es � b � l e� eme 1� c� yi eu1� � � �� y i � m1b 1s �i g� � �i �� l � i 1o�l 1es � e� � l � � �� � di �� g� e� �
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Capítulo 7
CARCASS TRAITS AND MEAT COMPOSITION OF LIGHT LAMBS
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� EG� E3� Gf� HE� C� Bt rH� � ED� � GD� �� E1B� N � �B � � EN FEHCBCED� �D � � CBH� G� I�B CEDH CF� WCB � 1N �D �
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EDH� B� E� � � C�� � B� Hf� �D � � E� � HCBt � m� ED � � B� �I df� ( kkzwf� �D � � CB� �H � G� 3� �I � � � �� IEW� G� I� 3� I� E� �
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FI� B EG�� E� � � � � BEGH� B �B � CD� I1� D� � � I�N � � N � �B � L1�I CBt � m� �M1� Ef� � �D � � h� �D � � � I� EDHEf� ( kk) wd�
� CH� CH� E� � HF � � C�I � G� I� 3�D � � � CD� IC B� I�N � � FGE� 1� BCED� HtHB� NH� W � G� � ICBBI� � CH� TDEWD� �� E1B�
B � � � EN FEHCBCED� E� � I�N � � N � �B � �B � � �G It � � � Hd�
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N � Dt � � � � CB� GG�D � �D � � E1DBGC� H� m� �M1� E� � B� �I df� ( kk) wf� ICBBI� � CH� TDEWD� �� E1B� � �G � �H H�
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� CFEII� H�� � G� � � f� WCB � �D � � FFGEi CN �B � � � � DH1H� E� � 2vfvvv� � W� H� CD� B � � DEGB � �H B� GD� G� CED� E� �
� F � CD� m� �H � II� H� � B� �I df� . vvz � wd� � CB CD� B CH� � EDB� i Bf� B CH� N �D 1H� GCFB� � E� 1H� H� ED� B � �
� � G�� B� GCh�B CED� E� � � �G � �H H� �D � � N � �B � � EN FEHCBCED� BG�C BH� � EG� IC B� I�N � H� � GEN � H � � F � � G� � � H�
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H � � F � � G� � � f� Cd� df� B � � � CFEII� H�d � � �G � �H H� BG�C BH� W� G� � � 3�I 1�B � � � � t � � EB � 1IBG�H E1D� �
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éj j á;� n� � � � yíyÉ� �� yíqá� o�r �� � N� �r � �� � � N� t e� l N9� � � � � � � � � � � � � � � � , � � r1�1 � � e� � u� �Nfí� xyyáz�
� 9 �r uní� éj j qz� � � r� � r� � u� �Nfí� éj j áz� � 9l 1 � n� � I í� éj j y;f� � eu9� el I� r� � s l � � � 1� u9� 9e� � NN9e�
e� � o9� rS�� 9� u� 1� r� r� eol � eu�� � 1� uI� �� �� N�� e� N� � � IL1í� �e � �t 1� �o r9a� � D�1 � 9� N�e � a� 1u�U �e �
� � rIa�� �e � � � � N�� I oN� � 1u�� IL1� � � � N�� � a�Nl �� IL1� � � � N�e � � �r �� u� rSeuI� �e � � � � N�� � �1 �N �
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� u� �Nfí� éj j ( z� � INNI� e� � u� �Nfí� éj j ) z� � ee� 1� � u� �Nfí� éj j Éz� � INe91� � u� � Nfí� xyyy;í� N� � � 9 o9eI� IL1�
n� � �NI� �� � � � � N�� Ie � � , � �b �� � u� �Nfí� xyy) ;í� �� � � e� � � � I� � 1uI� I� �r � � � 1� I� �u 9e� �� e� N� � � IL1� n�
l e� r� N�� I1� 9r �� IL1� 9� u� 1I� �� � 1� N9e� or9 r� �e � � � � � U9r� I� 1u9� � 1� uI� 9f� � 9e�
� 9 o91� 1u� e� � � � N� � � �1 �N � � � � � rS�1 � � � � e� r� � 91eI� � r�� 9e� Ul 1u9� � 91� 9ur9e� � �r �� u� r� e�
or9� l � uIa9e� � � � o� e9� � � 91L I� 9� r� N� a�1 u� e� �� � � e� � � � N�� I1� 9r �� IL1� � � N� r� 1� I I� 1u9� � � �
� r� � I I� 1u9í� � a�Nl � � IL1� aIel �N � � � I1� 9r �� IL1� � � N� o� � I rS� �N � u9 �r � � � � IeI91� e� � � � e� N� � � IL1�
� 1� N9e� r� � �4 9ef� � �� I oN� � 1u�� IL1� � � � N� � u� � 1I� �� � � � l Nur�e 91I� 9� �� uI� o9� r� �N � o�r ��
or� � � � Ir� N� e� � �r �� u� rSeuI� �e � � � � N�� � �1 �N � n� � � � N�� � �r 1� í� o� r IuIr�� �� 9ru�r � � N� uI� o9� � � � N�e �
orl � � �e � � � � or9 � 1I� � o�r � � � �r �� u� r� e� � � � N�� � �1 �Ní� r�� I�e � �� s l � � � e� l 1�� u� � 1I� �� 19� I1a�e Ia��
s l � � e� � ol � � � � r� �NIi �r � � 1� � N� �1 I �N � � 1� aIa9� n� N9e� � �u 9e� � � � N� � � �1 �N � ol � � � 1� e� r� � 9N� � u�� 9e�
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xyyé;f�
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n� � l �1 � 9� N�� D� r� � �� INI� �� � � � � N9e� Ie 9e� e� �� �n 9r� �� � � r9f� � eu9í� �� el � a� i í� o� r Iu� � �� I1� r�
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e� N� � � I91�r � �� N9e� � U9r� e� � �1 � I� �u 9e� o�r �� � 1ur�r � �N � � � 1ur9� � � � u� eu�U � � � � � 9rl � � 9e� � � � N��
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9 � 1u9� N�� � U9r�� � � � N� � or9NI� I� I� �� í � �e S� � 9 9� � N� �1 u� 1I I� 1u9� � � � N�e � � �r �� u� rSeuI� �e �
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ur� � �U9� e91� l n� I o9ru�1 u� e� o�r �� N�� r�i �� 9aI1�� � Io9NN� e�í � n�� s l � � r� or� e� 1u�1 � N��
o9eI� INI� �� � � � � 9� u� 1� r� l 1� or9 r� e9� � 1� uI� 9� � 1� � l � 1u9� �� � �r � � u� r� e� or9� l � uIa9e� ,If� fí�
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� 1� l � 1ur�1 � N9e� r� � � 49e� � � � r�i �� 9aI1�� � Io9NN� e�í � o�r uI� l N� r � 1u� � N9e� s l � � � eu�1 � I1e� rIu9e�
� � 1ur9� � � N� or9 r� �� � � � � 91ur9N� � � � r� 1� I I� 1u9e� � � � � � � f � N� I1u� r� e� � 1� � eu� � e� 1uI� 9�
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� � u� r I1�r � 1� � N� e� 1uI� 9� � 1� � N� � l �N � � � � � � Ir� N� � e� N� � � IL1í� � 91e� ra�1 � 9� N�� � � I� I� 1� I�� � l 1� I91�N�
� � � N9e� �1 I �N � e� � � � � � l � r� 9� �N � � I9� � 1� � N� � l �N � D�� Iu�1 � , � 9� 9� � u� �Ní� xyyá;f�
�
Capítulo 9
CONCLUSIONES
� � � � � � � �
� t j p� �� �
� � � � � � � � � � � �
� � � � � I � L� � �� � U� L� UI � S�� U� � 1OI � US U� � � � � � � OL� U� � S� � SL�� �� a� � �U � OL� � � � O�� � U�
� � � � I U� � � U� � � � � �U � � � � US� � �� � � � U� � � U�L L � � �� �U � � � � � U� � �O � SI � U� OL� � U� U � � � �U �
U� � I � � � S� Uc�
� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �
�
t u � �U � � � � � L� � � � �U � � L� � C� � � � U� � � SL� � L� � �� U� 6� UI � O � � �� � � � U� � � � � �� L�3 ��
� � �� � � O � � � U�a � U� � � � � � � � OL� � � � O�� � � � S� � �� � S�� � � � � � � � �U � � � �U a� O� U � 6�
�� SI L�u �
xu � U� L� UI � S�� U� � � � � U� � � � � � � U� �� �� � 3�� U� � U� O� L� � S� � � � � �U � � � � �L � �� � �� L�3 ��
� � �� � � O � � � U�� � � � � � � S�� �� � � � � � � 6� � � � 1 a� OL� � � � � �� S� � � � S� �
� � � � �� �� � � OL � I � � � C� � � � � � �L � � u�
pu � �� � � �U � � � � �� � C� � � � USCL� � � � � � � � U� � � US� � S U� � � S� O U� � UI � O � � �� � � � U� U� �
UI � � � L� � � � U� �U �� �a � � US�� � � � �� O � � �� � C� � �� SI �� � � L� �� �� �� � � �U � O L�
� I � U� � � S� O U� NI � � � � L� �� � � � � � �� � � 3� � �� � � � � �6 L� � � � � L� � � � � � �� � � � � U�
� � S� O U� �� S� � I Uu�
�
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� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �
�
nu � �U � � �S � L� � � �� � U� � � � � � LS�U � OI � � � � � U� L� � � � � �� �U � � �� � � � � NI � U�
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