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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

Transcript of ddd.uab.cat · 2014-12-17 · iv RESUMEN! # # Parapoderimplementar...

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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.  

103            

103              

104              

105    

       

106      

113  114  

       

 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)  

   

   

125            

126  

                                                                       

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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  

 

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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  

 

 100  

       

118      

119      

119    

120    

129    

130          

138      

139      

140    

141      

143  

   

   

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Capítulo  1  

INTRODUCCIÓN  

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gPhj .� � OLOut cOs � � � � TcO� r  tr c�0 � � u� �� , � �  O3 L t�� OvL0� � � � � 0� áj j y)(� � L �  � �3 � Ots �

� s Lv3 O� s 0�  �u � l L t�C �u � � s 3 c� O�   u� �  � u � ts c� s l OLs � c�� O� �L � L �  �� � �n �� O� �� � � �

s � t L � OvL � � � r L � � �3 n  O�� T�3 � � � � ncs � r � ts u0� � s 3 s � er us u0� � �c L � , � nc L � �u � n�c � � l utOc�

� �  �u � OL � r utcO� u� t mtO  � , � n   t c�0 � L tc � s tcs u(� � � 3 �u 0�  � T�L �� s � s l OLs � u� r L �   3 L ts �

Nr L � �3 L t�   � L �  �u � � s Ls 3 D�u � � � ur � uOut L � O�0 � Nc L � L � s �  � � � uns �  �� OvL � Rr 3 �L �� �  �

� � 3 ns � , � �, r � � L � s � �   � 3 �L t LO3 O L ts � �  � 3 � Os � cr c�   � 1� �x � 0� áj j á)(� � L �  s u� a  tO3 su� �o s u� u �

R� � c TOutc�� s � � r L � � � ncs Tc uOl �� � ncsN uOs L �   Ox�� OvL � �  � u � ts c� 1� � � � 0� áj j é)0�  � � � r �  0�

� n L � O L � s � � �  � �  s � �   Ox�� OvL � � �  �u � mn  s t�� Os L u� , �  �� s N ct�� , �  �� � 3 � L � �� � �  s u�

ncs � r � ts u0� R� � �R s L � �� s � L �  �� un � O�  Ox� � OvL � ncs � r � tOl �� n�c � � � � cL 0�   � R � s �  �L � 0�

uOtr �� OvL � er � � s L     l �� er �  s u� T�L �� cs u� � NOL � L ur u� s � C tOl s u� n�c ��  �� u   � � OvL � � �  s u�

s l OLsu(�

� � � � � � � � � � � � � � � � � � � � � i� � � � � � � � � � � � � � � � � � � � � L � � � � � � � � � � �

�  � � Lus � 3 r L � O�   � s l OLs � � u� O L � � �� g(j éé� 3 O    s L u� � � � �� x�u 0� � � �� r c� s � � s L �  �u �

L � r ut�u � c �   Ox�� � u� L �  � �o s � áj gj � ns c�  �� � cT�L Ox�� OvL � � �  �u � � �� Os L u� � LO� �u � n�c ��  ��

� TcO� r  tr c�� , � �  O3 L t�� OvL � 1 � � 0� RttnqGGN�s ut� t(N�s (s cTGN�s ut�t G)(� � � I uts u0�  � gá6 � 1gf á�

3 O    s L u� � � � �� x�u )� u � � s L � L tc�� L � � r cs n� 0� �r Ler �  � 3 �, s c� ns c� L t�C � � � s l OLsu� u �

L � r L tc�� L � � uO�� 1zá6 .� z yg� 3 O    s L u� � � � �� x�u )� , � � NcO� �� 1áé6 .� áPz � 3 O    s L u� � �

� � � x�u )0� , � � C� L � s � L � a  tO3 s �  r T�c � � � � � �L D�� 1gj 6 .� gj y� 3 O    s L u� � � � �� x�u )� , � � 3 I cO� ��

1P6 .� Pz � 3 O    s L u� � � � �� x�u .� OTr c�� g(g)(� � L tcs � � �  �� � LOvL � � r cs n �0 �  s u� s l OLsu�

� s LutOtr , L � r L �� n�c t � O3 ns ct�L t � �  � � Lus � T�L �� cs � 1� ��  �� g(g)0� s � r n�L � s �  � t c� c�

 r T�c � � unr I u� � � � s l OLsu� , � ns c� OLsu(�

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�y � �

� �

� � � � � � � r mr � � OutcO� r � OvL � �  � � Lus � 3 r L � O�   � s l OLs � ns c� � s L tOL L t u� 1 � � 0�RttnqGGN�s ut� t(N�s (s cTGN� s ut�t G)(�

�� � LOvL � � r cs n �� �   � cT� � L � ur � t ccOts cOs � r L � ts t�   � � � gj á� 3 O    s L u� � � � �� x�u �

s l OL �u 0� � ut�� �L � s � ur � � s L � L tc�� OvL � L � � t c3 OL �� s u� n�Du u� � s 3 s � � un�o �0 �  � � � r �   � u �

ns uO� Os L �� L � u Tr L � s �  r T�c � � s L � � �u O� áj � 3 O    s L u� � � � �� x�u � 1gP0áh6 )0� , �  � � OLs � � LO� s 0�

� s L �  � 3 �, s c� � Lus � � � T�L �� s � s l OLs � 1f á� 3 O    s L u� � � N � tOl s u0� f g6 )(�

�� t L � L � O� � �  � � Lus � T�L �� cs � L � � r cs n�� u� � � c � O L t 0� us � c � ts � s � L � � un� o�0 �

c N   C� L � s � r L �� � Ou3 OLr � OvL � � � �   c � � s c� �  � f j 6 � � u� �  � �o s � áj j j (� � u � �� ut��

uOtr �� OvL0� L �  � � er �  � � Lus � u � R�� c � r � O� s � � s LuO� c��   3 L t � � n L � O L � s �

ncOL � On�   3 L t � � �  � � s N ct�� , � � 3 �L � �� �  � 3 c� �� s 0� � �  �� � �n �� O� �� � � �  �u � mn  s t�� Os L u�

, � us � c � ts � s � � �  �� � s Ls 3 D�0 �  s u� s l OLsu� Ls � R�L � � C�� s � � � ut�c � � L tcs � � �  �u � un � O u� � s L �

3 � , s c� La3 cs � � � �L O3 �   u� L �  � N � tOl s � T� L �� cs � 1 OTr c�� g(á)(�

� � � � � � � r mt � � N � tOl s � T�L �� cs � L � � un� o�� ns c� un � O � 1üg(j j j .� � � � � 0� áj gj )(�

Page 26: ddd.uab.cat · 2014-12-17 · iv RESUMEN! # # Parapoderimplementar #programasdemejoragenéticaenelsectorovinode carne,# es imprescindible definir y evaluar los caracteres biológicos

� � � � � � � � � � � � �

�A� �

� �

� � � � r mr � � Lus � T�L �� cs � L �  �� � LOvL � � r cs n � � 13 O   u� � � �L O3 �   u)� L � áj j P� 1 � � 0�RttnqGGN�s ut� t(N�s (s cTGN�s ut�t G)(�

� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �

�   3 � LO� � á(f yj � gá(Pzy� ááj � áA(hhé�

� r utcO� � f f f � g(PPé� Af � f (j Az �

� I  TO� � � gáA� á(Aj j � f á� A(f ág�

� r  T�c O� � g(zéy� yAy� z f j � éhz �

� ROnc áAé yA f gh zAy

� OL �3 �c � � � gj z � g(yz j � j � gá(f AP�

� u  s l LO� � gf P� zéj � áz � z f á�

� un�o � gP(égh A(j áj á(áAy áA(áPj

� uts LO� � éh� áf h� z � f Ay�

OL  �L � O� � ggh� Pgh� A� g(f hg�

c� L � O� é(égy gP(gPP g(áAé gz (hgj

c � O� � h(PPz � Aáj � z (géh� Pzá�

� s  �L � � � g(j PP� f (PPA� zgA� gá(gj h�

� r LTcD� � g(áf A� éj g� AA� f (f hf �

� c  � L � � � z (ééh� A(égA� gj � g(zAh�

� t�   O� � h(géy� A(zzé� Pyé� P(áyá�

ts LO� � Aé� f hj � gf � f hz �

Otr � LO� � zh� éég� gé� hPé�

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�

� u  s l �e r O� � f Aá� zhz � f A� ézg�

� r 3 �L O� � h(hhá� á(Ahz � hPh� A(géz �

� r � O� � yz j � g(yf h� j � g(yáP�

� s t�   � gj á(áPf � hP(zhP� gg(PzP� gyf (f gP�

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� � � � � � � � � � � � �

�é � �

� �

�� � OutcO� r � OvL � � �  �� T�L �� cD�� s l OL �� L � � un�o � � �� �c � � � 3 �u � �  � hA6 � � �  s u�

3 r LO� OnOsu� 1� LOts 0� áj j é)(� �  � T�L �� s � s l OLs � ut�� nc u L t � L � nc�� tO� �3 L t � ts � s �  �

t ccOts cOs � un�o s  0� � ut�� �L � s � � �u tO    �� , � vL0� � �u tO    � í �� � �L � R� 0� � mtc 3 � � r c�0 � � c�T vL � , �

� L � �   r � D� � 1 OTr c�� g(f )(� � s � c u�   � � �� �C s x0� � s L � á(f éh(j áj � �L O3 �   u0� � s 3 s �  �� ncs l OL � O��

un�o s  �� � s L � 3 �, s c� � Lus 0� u Tr O� �� � � � �� c u� , � � Or � �� � � �   0� er � ur n c�L �  � 3 O    vL � � �

N � tOl s u0� , � � � � cr  � , � � �c �T s x� 0� er � u � �n csmO3 �L � �   � 3 O    vL � � � � � � x�u � � � T�L �� s � s l OLs (�

� L �  �� t��  �� g(á� u � 3 r utc��  � La3 cs � � � s l OLsu� u TaL � tOns � 1� s c� cs u0� u 3 L t�   u� , �

s l C� u)0� n ct L � O L t u� � �  �u � � s 3 r LO� �� u� �r tvLs 3 �u � � � � un�o �( � � L �    �� u � nr � �

�n c � O�c �  � La3 cs � ts t�   � � � �L O3 �   u� , � t�3 � OI L �  s u� tOnsu� ncs � r � tOl s u� � � 3 � , s c�

c   l �L � O�( � �

� � � � � �

� A� � � � ó� � � � �� � � � � �

� � � r � � � � �� � � � � � � �

� � � � � � � � � � � r � � � �

� � � � r � � �

� A � r � �

� � � � � �

� � � � � � � � � � � �� Ar � �

� � A � � �

� � � � � �

a � � �� � � � � � �

� � � � � �

� A� � � C� � � � � � �

� gA� � � � r � �

� � � � �

� � � � � � � r mn� � OutcO� r � OvL � �  � � Lus � s l OLs � ns c� � s 3 r LO� �� u� �r tvLs 3 �u � L � � un�o � � 1� � � � 0�áj gj )(�

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� h� �� �

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� � � � � � � � � � � � � �

� P� �� �

� �  s u� � �u O� áf � 3 O    s L u� � � � �� x�u � � � s l OLs � mOut L t u� L � � un�o �0 � uO� u � tO L � L �

� r L t� �  � � �n tOtr � � ncs � r � tOl �0 � �   c � � s c� �  � há6 � us L � s l C�u � 3 �� c u� � utOL �� �u �

m�  r uOl �3 L t � ��  �� ncs � r � � OvL � � � � s c� cs u� 1gy� 3 O    s L u)0�  � gy6 � us L � s l C�u � � � s c� os �

1á0h� 3 O    s L u)0� r L � gj 6 � us L � � s c� c�u � � � c ns uO� OvL � , � r L � gy6 � � s c� cs u(� �u � c ncs � r � ts c�u �

er � n ct L � L � �� c�x �u � � �c LO� �u � u � r � O� �L � Nr L � �3 L t�   3 L t � L � � mtc 3 �� r c� 0� ��u tO    �� , �

vL0� � �u tO    � �  �� � �L � R�0 � � L � �   r � D�� , � � c�T vL(� � L � � r � L ts � ��  �u � s l C�u � � � s c� os 0�  �� 3 �, s c�

n� ct � � �  �u � 3 Ou3 � u� u � � s L � L tc�L � L �  �u � � s u� � �u tO    �u (� � � n u�c � � � er � � �t �   r o�� L s � u �

L � r L tc�� � L tcs � � �  �u � ncOL � On�   u� � s 3 r LO� �� u� �r tvLs 3 �u � � s L � 3 �, s c� � Lus � � � T�L �� s �

s l OLs0� ur � � s L tcO� r � OvL � ��  �� ncs � r � � OvL � � � � �c L � L � � un�o �� u� O3 ns ct� L t � 1� � � � 0� áj gj )(�

� � � � � � � � � � � � � � � � � � �

s u� s l OLsu� us L �  �� un � O � � s L �  � 3 �, s c� La3 cs � � � c�x �u � �� LOl  � 3 r L � O�   0� OL �  r us �

s � l O�L � s �  �u � L r 3 cs u�u � c�x �u � mtOLTr O� �u � R�u t��  �� �� tr �   O� �� � 1 � � 0� áj j é)(� �� � OutcO� r � OvL �

� �  �u � � OutOLt�u � c�x �u � L � � un�o � 0� u� � s 3 s � u � 3 r utc�� L �  � � t��  �� g(f (� �

� �u � �  � Pj 6 � � �  s u� N � tOl s u� u � �T cr n�L � L � A� c�x �u � 3 �, s cOt�c O�u 0�  �u � � r �   u� us L0�

ns c� s c� L � � � O3 ns ct�L � O� q� � cOL �0 � � �u �� � c�T s L u� 0� � �L � R T�0 � � Tr c o�� , � � �u t    �L �( � � L �

� r � L ts � �� � �t �   r o�� , � ts 3 �L � s � L � � r L t��  � c � r � O� s � � Lus � L � � s 3 n� c�� OvL � � s L �  �u � s tc�u �

� s 3 r LO� � � u� �r tvLs 3 �u � 1á0hg6 )0�  � � Lus � � �  �u � c�x �u � �r tv � ts L �u � � �t �   �L �u � c ur  t��

OLN cOs c� �   � � �  �� 3 �, s cD�� � � c�x �u � un�o s  �u 0� �r Ler � u � s � u cl �� er �  �� � Ons     u�� 1éj (j j j �

s l C� u)� nc � s 3 OL �� L � c  �� OvL � ��  �� � Ouer t� � 1gy(j j j � � �� x� u)� , � ��  �� � c�L u�� 1g(éj j �

� � � x�u )(� �

� �

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� � � � � � � � � � � � � �

� gj � �� �

� � � � r mn� � utO3 �� OvL � �  � � Lus � s l OLs � � un�o s  � u TaL � c�x �( �

� �x � � � Lus � 1 utO3 �� s )� � ntOtr � �

� cOL � � f (z j j (j j j � � � cL � í �  � L � �

� �u �� � c�T s L u� � á(gj j (j j j � � � cL �

� �L � R T� � g(Aj j (j j j � � � cL � í �   � R �

� Tr c o� � g(yj j (j j j � � � cL �

� � ut    �L � � g(áyj (j j j � � � cL � í �   � R �

� uu�N � Pj j (j j j � � R � í �   � R�x s u�

� Rr cc� � hj j (j j j � � R � í �   � R�x s u�

�� R� � yj j (j j j � � R � í �   � R�x s u�

� Tcr n�� OvL � L tc NOLs � í � NOLs � yj j (j j j � � � cL �

� � l �c c� � z éj (j j j � � � cL �

� C�  �� � � f j j (j j j � � � cL �

� s L t uOL � � gj j (j j j � � � cL � í �  � L � �

� l OLsu� nc � s � u� hj (j j j � � � cL � í �   � R�x s u�

� Ons     u� � éj (j j j � � � cL �

� � L �c O� � z j (j j j � � R � í � � � cL �

� COL Tc� � z j (j j j � � � cL �

� Ouer t� � gy(j j j � � � cL �

� c�L u� � g(éj j � � � cL �

� � � � � � � � � � � � � � � � � � � � � �

� mOut L � � ON c L t u� uOut 3 �u � � � ncs � r � � OvL � � L tcs � � �  � � T�L �� cD� � s l OL �( � � uts u�

� n L � L � �  � s � C tOl s � � � ncs � r � � OvL � , �  � � � Ouns LO� O  O� �� � T s Tc�NO� � 0� ncOL � On�   3 L t (� � u�

O3 ns ct�L t � t L c� L � � r L t�� r L �� u cO � � � N�� ts c u� � �u O� s u� n� c��  � u � ts c� s l OLs0� er � us L �  s u�

er � L s c3 �   3 L t � � t c3 OL �L �  � tOns � � � uOut 3 �� ncs � r � tOl s � 1� �x � 0� gPPé). L tc �    s u

� ut� � �L q�

� � � ts c u� tI � LO� s uq� c ncs � r � � OvL0� �   O3 L t�� OvL0� u�L O� �� 0� � �u � T L I tO� �0 � t� (� � s L �

N� � ts c u� er � OLN  r , L � L �  s u� n� c�3 tcs u� ncs � r � tOl s u� , � ns c� t�L ts � L �  �� c L t�� O  O� �� (� �

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� gg � �� �

� � � ts c u� utcr � tr c�   u� , � � s Lv3 O� s uq� t�3 �o s u� �  � c � �o s 0� tOns � � � NOL � �0 � 3 �L s � � �

s � c�� , � � � n� � Ot�� OvL0� uOut 3 �u � � � � s 3 c� O�   Ox�� OvL0� OL l cuOs L u0� t� (� � uts u� N�� ts c u�

ur   L � u c� 3 L s u� � s LuO� c�� s u� ��  �� Rs c�� � � 3 n x�c �  �� ncs � r � � OvL � , � uOL � 3 � �c Ts �

OLN  r , L � L �  s u� � s ut u� � � ncs � r � � OvL0� L �  � ncs � r � ts � NOL �   � , � L �  � � c L t�� O  O� �� � 1� r m�� I 0�

gPPA)(�

� r Ler � nr � L � � � cu � � s 3 � OL �� Os L u� , � � ON c L t u� Tc�� s u� � � ur n cnsuO� OvL0�  s u�

� ON c L t u� uOut 3 �u � � � ncs � r � � OvL � er � ns � 3 s u� L � s L tc�c � us Lq� mt LuOl s � tc�� O� Os L �   0�

u 3 Oí mt LuOl s 0� u 3 OíOLt LuOl s � � OLt LuOl s (�

� � � � � � � � � � � � � � � �

� �  � uOut 3 �� � � ncs � r � � OvL � mt LuOl s � tc�� O� Os L �   � 1 OTr c�� g(z )� 3 n   �� n�u ts c s �

� s L � r � O� s � s � L � Tc�L � u� � c� �� s u0� Ls c3 �   3 L t � uOL � ut�� r  �� OvL � LO� �   O3 L t�� OvL �

� s 3 n   3 L t�c O�( � s u� �L O3 �   u� l �c D�L � � � n us � , � � s L � O� OvL � � s cns c�   � � n L � O L � s � � �  ��

� Ouns LO� O  O� � � � � � �   O3 L ts 0� � �� s � ns c�  �� I ns � �� �  � �o s � , �  � u� � s L � O� Os L u� �  O3 �t s  vTO� �u �

1� r m� � I 0� gPPA)(� �u � OLut�   � � Os L u� �� s utr 3 � c�L � �� u c� cr � O3 L t�c O�u � s � OL �  r us � OL mOut L t u0�

� � uO� uOL � OL l cuOs L u� L � OLNc� utcr � tr c�u � , � � s L � 3 � L Cs � tc�� O� Os L �   � , � cr tOL �c Os (� �� OLt LuO� �� �

c ncs � r � tOl �� ur   � u c� 3 r , � � �C �0 � � s L � r L � n�c ts � ns c� �o s � , � ncs  ONO� O� �� � � �C �( � �� c L t�� O  O� �� �

� � ut � tOns � � � mn  s t�� Os L u� � n L � � �  � � s ut � � �  �� 3 �L s � � � s � c�� ns c� s l C�� , � � �  ��

I ns � � � �  � �o s � L � er � u � ncs � r � L �  s u� n� cts u(� � L � ut � tOns � � � uOut 3 �u � � � ncs � r � � OvL0�  � �

r tO  Ox�� OvL � � � 3 a  tOn   u� 3 �� Rs u� � r c�L t �  � u� � r � cO� Os L u� O3 nsuO� O  Ot��  � � O� L tONO� �� OvL � � �

n� t cLO� �� u.�  s � 3 Ou3 s � ur � � � � s L �  � u� 3 �t cLO� �� u� � r �L � s � u � nc u L t�L � 3 a  tOn   u�

n� cts u� Ls � � s L tcs  �� s u� ��  �� l x(� � uts � OL � O� � � Oc � t� 3 L t � us � c �  � � ns uO� O  O� � � � � � c �  Ox�c �

l �  r � � Os L u� T L I tO� �u 0� , �� er �  s u� ccs c u� L �  �� O� L tONO� �� OvL � � �  s u� ncs T LOts c u� u uT� L

 � � utO3 �� OvL � � � n� c�3 tcs u� T L I tO� s u� 1� �L � �   � Ó0� gPéj �) � , � l �   s c u� 3 Cs c�L t u� 1� uc�  � , �

�     c0� áj j j .� � �L � �   � Ó0� gPéj � )0� � c � L � s � OL � s LuOut L � O�u � L �  �� l �   r �� OvL � � � u 3 L t�   u�

ns c�  �� OL � ctO� r 3 � c � L �  �� OLNs c3 �� OvL � T L �   vTO� �� 1� 3 0� gPPá)� , � �N � t�L � s � OL �  r us �  �

ncs Tc us � T L I tO� s � � �  �u � � OutOLt�u � c�x �u � 1� � L s u� t� �   (0� áj j g.� � uc�  � , � �     c0� áj j j .� � �L �

�   � Ó0� gPéj � )(� �

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� gá � �� �

� mOut L � l � cO� �   u� �  � uOut 3 �� mt LuOl s � tc�� O� Os L �   � u TaL �  �� 3 s � �   O� �� � � � n�u ts c s �

1� r m� � I 0� gPPA.� � �x � 0� gPPé)q�

� � �u ts c s � ut�L t q�  � T�L �� s � n�u t�� L �  � 3 Ou3 s � tI c3 OLs � 3 r LO� On�   � � � s cOT L � , �

ncsmO3 O� � � u(�

� � c� Lut c3 Ot�L � O�q �  � T�L �� s � �n cs l � R�� c Tr  �c 3 L t � n�u ts u� � � s tcs u� tI c3 OLsu�

�   Ts � �   C�� s u0� � s L � ut�L � O� u� Nr c�� �  �  r T�c � � � s cOT L (� �

� � c� LuRr 3 �L � O�q � � s LuOut � L �  � �n cs l � R�3 O L ts � ut�� Os L �   � uOut 3 �t O� s � � �

n� uts u�   C� L s u(� � � l �  O3 Ot� � s � ns c� � s L � O� Os L �L t u� us � Os  �� s c�   u(� �

� � s 3 �� Ou3 sq� n�u ts c s � OtOL c�L t � uOL �  r T�c � NOCs � � � c uO� L � O�( � � s � mOut � L �

� un�o �( � �

� � � � � � � r (c � � mn  s t�� OvL � mt LuOl �( �

� � � � � � � � �

� s u� uOut 3 �u � � � mn  s t�� OvL � u 3 Oí mt LuOl su� 1 OTr c�� g(y)� us L � uOut 3 �u � L � n�u ts c s �

� s L � ut�� r  �� OvL � � r c�L t �  �� L s � R � , � �� l � u� OL �  r us � � r c�L t �  ��  �� t�� OvL0� t�3 � OI L �

� L s 3 OL �� s u� uOut 3 �u � tc�� O� Os L �   u� 3 Cs c�� s u� , �� er � � r L t�L � � s L � � O ct� � n  �L ONO� �� OvL � �

OLt LuONO� �� OvL � c ncs � r � tOl � (� � s L � 3 r , � � n L � O L t u� � �  �� � Ouns LO� O  O� �� � � � n�u ts u� , � u �

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� gf � �� �

ur n   3 L t� � � s L � �   O3 L t�� OvL � � s 3 n   3 L t�c O�0 � L s c3 �   3 L t � L �  �� I ns � �� � �

c ncs � r � � OvL(� �u � OLNc� utcr � tr c�u � � OLut�   �� Os L u� ut�L � 3 Cs c�� �u � L � c  � � OvL � � s L �  �

uOut 3 �� mt LuOl s � tc�� O� Os L �   0� � s L � c�� Os L �   Ox�� OvL � �  � 3 �L Cs � , � u�L O� �� � c  �t Ol �3 L t �

NO� O L t (� � r � � � �c � �   t�� c L t�� O  O� �� � uO�  � c L � O3 O L ts �  �� s c�   � u� �� � r �� s (� � u�  � uOut 3 ��

er � 3 �u � u � r tO  Ox�� �� tr �   3 L t � L � � un�o �� 1 r O    �r 3 t� , � � �C �0 � áj j g)(�

� � � � � � � r md� � mn  s t�� OvL � u 3 Oí mt LuOl �( �

� � � p � � � � �

� s u� uOut 3 �u � � � mn  s t�� OvL � u 3 OíOLt LuOl su� us L � uOut 3 �u � � � n� uts c s � tc� � O� Os L �  �

ut� L t 0� T L c�   3 L t � � s L � r � O� s 0� � s L � ut�� r  �� OvL � �� NOL � � � T ut�� OvL0� � r c�L t �  ��

 � � t�� OvL � , � ns c�  �u � L s � R u(� �� n  �L ONO� �� OvL � � �  �� �   O3 L t�� OvL � u � � s s c� OL �� � s L �  ��

c ncs � r � � OvL � , � u � � �L � DL � O� u� c ncs � r � tOl s u� 3 s � c�� s u� uO�  �� �  O3 L t�� OvL � u� � s cc � t�� , � u �

�n s , ��  � ncs � us � c ncs � r � tOl s � � s L � tc�t �3 O L ts u� Rs c3 s L �   u� L � ncO3 �l c�( � � �3 � OI L � u �

nr � L � nc u L t�c � � s 3 � OL �� Os L u� � � s � C tOl s u� � s L � � ON c L t u� �   t cL �t Ol �u � � � 3 �L Cs � , �

t � L s  s TD� 0� � s 3 s � n� uts c s � mt LuOl s � � � l O L tc u� , � s l C�u � � � c ns uO� OvL � � s L � �   O3 L t�� OvL �

L � � s cc�   � � � �L O3 �   u� n�c �� �� �u ts 0� � cD� � s � mnsuO� Os L u� T�L �� c�u (� � tc�� l �c O�L t � �  �

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uOut 3 �� u�  � n�u ts c s � � � �   t�� � LuO� �� � , � � s ct�� � r c�� OvL0� � s 3 � OL �� s � � s L � �   O3 L t�� OvL � s �

NOL �  Ox�� OvL � L � � s cc�   (�

� � � � � �

� uts u� uOut 3 � u� � � ncs � r � � OvL � u � � �c �� t cOx�L � ns c� u c� � � ut�� r  �� OvL � � s 3 n   t� �

1 OTr c�� g(A)(� � s L � ut � tOns � � � uOut 3 �� � � ncs � r � � OvL � u � s � tO L L � � r L s u� c ur  t�� s u� t�L ts �

ncs � r � tOl s u0� � s 3 s � � s Lv3 O� s u0� n cs � n�c � � � s Lu Tr Oc� c L � O3 O L ts u� vntO3 su� u� L � u�c Os �

� s L t�c � � s L � T L s tOnsu� u   � ts u� � � �   ts � LOl  � ncs � r � tOl s 0� c �   Ox�c � r L �� 3 r , � � r L ��

n  �L ONO� �� OvL � � �  �� �   O3 L t�� OvL0� ur � � s ut � , � � OutcO� r � OvL0� , � t L c� r L � � r L � c L � O3 O L ts �

 � � s c�   0� � Ou3 OLr , L � s �  � � s ut � � � 3 �L s � � � s � c�� ns c� s l C�( � � s c3 �   3 L t 0� uts u� uOut 3 � u�

� � ncs � r � � OvL � us L � � � t�3 �o s � 3 � O�L s � s � Tc�L � (� � � 3 �u 0� R�, � er � t L c� r L �� �t L � OvL �

u� LOt�c O�� � s L tOLr �0 � � � O� s � ��  �� � s L � L tc�� OvL � � � �L O3 �   u� , � r L � � r L � � s L tcs  � � � n�c ts u�

n� c�� l Ot� c� ncs �   3 �u � � � 3 �t cLO� �� u� � u� s Ls � O� �u 0� er � �r Ler � u � nc u L t�L � L � � �C s �

ns c� L t�C 0� nr � L � t L c� c n c� r uOs L u� L �  s u� ncs Tc�3 �u � � � 3 Cs c�� T L I tO� �( �

� � � � � � � r (e � � mn  s t�� OvL � OLt LuOl �( �

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� � LOl  � 3 r L � O�   � u � utO3 � � r L �� ncs � r � � OvL � � � h(áz j (j f g� ts L  �� �u � 3 I tcO� �u � � �

� � cL � L � � � L �   0� � s L � � us � c u�   L � ncOL � On�   3 L t � tc u� n�Du uq� � ROL �� � s L �  � áy0z6 0�

� r utc�   O�� � s L �  � h0j 6 � , � � r l � � �  �L � �� � s L �  � y0h6 � �  � ts t�   � � �  �� ncs � r � � OvL � 3 r L � O�   � � �

� � cL � � � s l OLs � 1 � � 0� RttnqGGN�s ut� t(N�s (s cTGN�s ut�t G)(�

� � LOl  � r cs n s 0� L � � s L � s c� �L � O�� � s L �  � � � OutcO� r � OvL � � �  s u� � Lusu� s l OLsu0�  ��

ncs � r � � OvL � u � � s L � L tc� � L � � OL � s � n�Du uq� � OLs � � LO� s 0� � un�o �0 � c�L � O�0 � � c  �L � �� � � t�   O�0 �

er � �� �c � �L � 3 �u � �  � hj 6 � �  � ts t�   (� � ut�� �L � us � c �  � c uts �  � � OLs � � LO� s � , � � un�o �0 � er �

ncs � r � L � L tc � �3 � s u� �   c � � s c� �  � yé6 � �  � ts t�   � � � � �c L � s l OL �� 1� ��  � � g(z .� � � 0�

RttnqGGN�s ut� t(N�s (s cTGN�s ut�t G)(� �u � c  �� Os L u� ncs � r � � OvLG� Lus � n�c �� � �c L � , �   � R � L �

� r cs n� � us L �  �u � 3 �u �   l �� �u � L �  � � s L t mts � 3 r L � O�  � 1� �x � 0� áj j á)0� Tc�� O�u � �   � � u�c cs    s �

t � L s  vTO� s 0� un � O�  3 L t � � �  s u� n�Du u� � �  �� � LOvL � � r cs n �( �

s u� c ur  t�� s u� � �  �u � L � r ut�u � T�L �� c�u � �  � áj j P� c �   Ox� � �u � ns c�  � � � � � �

c l  �L � er �  �� ncs � r � � OvL � � � � �c L � � � s l OLs � L � � un�o �� u� � � �n csmO3 �� �3 L t � gág(j j j �

� 3 � �L r �   u0� ��  � u� er � R�, � er � �o �� Oc� r L �� � �L tO� �� � �� O� Os L �   � � � � �c L � � utOL �� �� n�c ��  �

�r ts � s Lur 3 s � � �  s u� ncs � r � ts c u(� �� Ns c3 �� L �  �� er � u � � s 3 c� O�   Ox��  �� � �c L � � � s l OLs � L �

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�� 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 � � �

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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 (�

�� 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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� � V� B B� � � �V u� � S� � I � � B� � � BI � �� � J � � � � S� � �� D� � � � � BV� � � � � SBV� � � � BV� E�I �� � BI � � IBV�

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V� � V� �E �I S�� BV� � BI I � VEB � � � S� V� �r �

s d �� I �e � � B2� � � � � EB� � � V� �

pd �� eBB� � SI � � �

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t d � � B� I � � � � �

xd � � I �� S� I � VS� � �V � � � � � � � � � �� � �

i d � � I �� S� I � VS� � �V � � � � � � � � �I � �

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Capítulo  3  

REVISIÓN  BIBLIOGRÁFICA  

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Capítulo  4  

CHARACTERIZATION  OF  THE    RIPOLLESA  BREED    

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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  

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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  

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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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CHARACTERIZATION  OF  THE  RIPOLLESA  BREED  

  72        

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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  

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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).    

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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).    

 

 

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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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  81        

 

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  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.  

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  83        

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  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.    

 

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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.  

 

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  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  

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  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.    

 

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Capítulo  5  

BAYESIAN  INFERENCE  OF  MATERNITIES  

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O� i � N� � M   � � E�S � � nS �E � � NC� � � S �E � go� � BCM   � MsF� � nOBI E� o� M1 BI Oo� MCBFE� MO� NC� RT�� CTFE� MOgo� � � � N� �

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,aS � BO� M   � � �� � BMB9� �

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N� M�B I � � � � O� M   � � �9 � C� � � N� � M   � BC� F �C � I MO� � C� � � BI � 9�E T� O� FETO� �� � � I � � EB�I � O�S FEBI � M� CS �

� C�1 I � � CNS � ( ) 2,1

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�I � � � �S e� � �C B�I � � � � NS FNI � I MO� 1 � C� � OM�M� � � �O � M   � � 9�E T� O� FC� 9BNTOEs� N� M�B I � � � � s � ��O � EE�O �

� M� � Ee� naccpg� NI � M   � � O�S � � � �M�� O� M� n� �� E� � f el ge�

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� � 9� ENF� � � BI � M   BO� S �I TO� CBFMe� � NTI � �C B� O� � NC� M   � � � E�M� FCBNCO� N� � � � nrl occc � MN� l occcg� �I � �

9�C B�I � � � � NS FNI � I MO� nc � MN� l occcg� 1 � C� � �O OTS � � � E�C � � � I NT   � MN� � N9� C� �� 9� Cs � 1 B� � �

F � C�S � MCB� � OF �� � e� � � MNM�E � N� � af occc � � � CYN9� �   �B I � � NI M� � � �C EN� n� � � � g� � s � E� O� 1 � C� � CTI �

� NC� M   � � �I �E sOBO� N� � � ��   � OBS TE�M� � � FNFTE�MBNI o� �� M� C� �� � TCI rBI � F � CBN� � N� � l occc � � s � E� Oe� �   � �

E� I M   � N� � M   � � � TCI rBI � F � CBN� � 1 �O � E�C � � � I NT   � MN� � I OTC� � � NI 9� C � I � � e� �   BO� 1 �O �

� � M� CS BI � � � � TCBI � FC� EBS BI �C s � �I �E sO� O� � s � 9BOT�E � BI OF� � MBNI � �I � � � s � M   � � S � M   N�� N� � � �� M� Cs �

�I � � � � 1 BO� nl ppag� NI � 2a � �I � � i e� � � �

�   � � FCN� �� BEBMs� N� � �O OB I S � I M� MN� M   � � CB   M� � �S � n � � g� 1 �O � � �E � TE� M� � � � NC� � ��   �

E� S � � � M� M   � � � I � � N� � M   � � � �C YN9� �   �B I � � NI M� � � �C EN� n� � � � g� FCN� � OOe� � N� TOBI � NI � M   � � �M   �

E� S � o� M   BO� F �C �S � M� C� 1 �O � � � � BI � � � �O � Trdi IIPARD = o� 1   � C� � rdI � 1 � O� M   � � I TS � � C� N� �

� � � � � BM� C� MBNI O� �   NNOBI � M   � � CB   M� � �S o� �I � � TI � 1 � O� M   � � MNM�E � I TS � � C� N� � � � � � �

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� py � �� �

BM� C�MBNI O� nBe� eo� af occcge� � � F �C MTC� O� � CNS � �I � � i F � � M� � � cef � � � � � � TI � � C� � EBI � � �O OB I S � I M� MN�

M   � � M1 N� � �I � B� �M� � � �S O� 1 � C� � � 9�E T�M� � � � s � M   � � � � � � NFMBNI � BS FE� S � I M� � � BI � M   � � � � � ��

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� 9� ET� M� � � BI � M� CS O� N� � O� I OBMB9BMso� M   BO� F �C �S � M� C� � � BI � � � � BI � � � �O � M   � � FCNFNCMBNI � N� � � �S �

�O OB I S � I MO� � NCC� � MEs� B� � I MB� B� � � �O � OT�   e� NM� � M   �M� � � � � � C� OMCB� MO� MN� �� B9� I � OBS TE�MBNI �

�I � � � I �E sOBOo� 1   � C� �O � O� I OBMB9BMs� �� � NTI MO� � NC� M   � � FCNFNCMBNI � N� � MCT� � FNOBMB9� O� 1 BM   BI � ��

OBS TE�MBNI � O� � I �C BNe� � �

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� pp� �� �

� � M� � � � � � � � � � � M� � � � � �

�   � � F � C� NCS �I � � � N� � M   � � S � M   N� NEN s � NTMEBI � � � �� N9� � � NC� BI � � CCBI � TI YI N1 I �

S � M� CI BMB� O� BI � E�S � O� 1 �O � � 9�E T�M� � � � s � OBS TE�MBNI e� �   � � �I �E sO� O� F � C� NCS � � � NI � M   � �

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S � M� CI BMs� 1   � I � M1 N� � �I � B� �M� � � 1 � O� 1 � C� � BI 9NE9� � e� � � OFBM� � M   BO� �9 � C� � � F � C� NCS �I � � o� ��

C� S �C Y�� E� � � � C� � � N� �   � M� CN � I � BMs� 1 �O � N� O� C9� � � �� CNOO� OBS TE�MBNI O� O� � I �C BNOe� I � � � � o�

1 BM   BI rO� � I �C BN� � 9� C� � � � � � � � 9�E T� O� � ET� MT�M� � � � � M1 � � I � f aef 4 � nO� � I �C BN� � � bt� � � v� cel g�

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M   � M� C� 1 � � � � � � � OMBS �M� O� N� M�B I � � � � CNS � � ��   � �I � EsOBO� O   N1 � � � �� 1 B� � � BI M� C9�E � N� � 9�E T� Oo�

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f ef e� NM� � M   �M� M   � � C� OTEMO� � CNS � O� � I �C BNO� � � a� �I � � � � b� 1 � C� � � B� � � C� I MB�M� � � � � F � I � BI � NI �

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O   � � F � BI � TOMCso� 1 BM   � EN1 rMNrS N� � C�M� � F � C� � I M� � O� N� � �� �I � NI � � � E�S � O� BI � � NS S � C� B�E �

� EN� YO� n� E� i �I � � C� � M� � Eeo� l pybge� �   � C� �O � M   � � ENOO� N� � OBC� � �I � � � �S � B� � I MB� B� �MBNI � � �I � � � �

EBI Y� � � MN� M1 N� � B� � � C� I M� S �B I � � �T O� O� nBe� eo� S TEMBFE� � S �MBI O� �I � � �� �I � NI � � � E�S � Oo�

C� OF � � MB9� Esgo� M   � s � � NM   � C� Es � NI � S BOOBI � F � � B C� � � � �M�� n� �F F � � M� �E eoaccugo� 1 BM   � O� 9� C�E �

  � CS � TE� � NI O� RT� I � � O� MN� � C� � � BI � FCN C�S O� n� �I � � E� � Yo� l puc �o � t� � �I NO� � M� �E eo� accl ge�

� � 9� C� E� �I �E sMB� �E � �F FCN��   � O� 1 � C� � � � 9� ENF� � � MN� �� � NTI M� � NC� OBC� � TI � � CM�B I Es � n��C � NON�

�I � � � S F � ES �I oaccbt� � �F F � � M� � Eeoaccug� �E M   NT   � M   � C� � BO� �� E�� Y� N� � OF � � B� B� � �I � � EN1 r� NOM�

S � M   N� NEN B� O� MN� BI � � C� S BOOBI � � �S O� � CNS � �9 �B E�� E� � F   � I NMsFB� � �I � � F � � B C� � � � �M�e � NM� �

M   � M� M   BO� YBI � � N� � F � � B C� � � � �M�� ENOO� � �I � NI Es � � � � � � � BI BMB9� Es � C� � N9� C� � � � s � �F FCNFCB� M� Es �

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BS FE� S � I M�MBNI e� � EM� CI �MB9� � EN1 r� NOM� �F FCN��   � O� OT�   � �O � M   � � I �B 9� � S N� � EBI � N� � S BOOBI �

F � � B C� � � � �M�� nBe� eo� � I � MB� � CNTFOt� � � OM� EE� � M� �E eo� l pyyg� S TOM� � � � � NI OB� � C� � � �O � �I �

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C� � T� BI � M   � � EBOM� N� � O� E� � MBNI � � �I � B� �M� O� �I � � M   TO� � I NMsFBI � � NOMO� � NC� � NI � BCS BI � M   � �

S � M� CI �E � B� � I MBMs� N� � M   � � S NOM� BI M� C� OMBI � E�S � O� BI � M� CS O� N� � M   � � FC� � B� M� � � � �S e� � I � � C�

� � NI NS B� � C� OMCB� MBNI Oo� � I NMsFBI � � � � NCMO� � NTE� � � � � � BC� � M� � � MN� M   NO� � E�S � O� �O OB I � � � MN�

� EBM� � � 1 � Oe�

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� � M� CS BI �MBNI � TOBI � F   � I NMsFB� � �I � � F � � B C� � � � �M� o� 1 � � � � 9� ENF� � � OF � � B� B� � F �C �S � M� CO� � NC�

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BI � NCS �MBNI � nBe� e� OsOM� S �MB� � �I � � F � CS �I � I M� � I 9BCNI S � I M�E � � � � � � MOge� MO� F � C� NCS �I � � � 1 �O �

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� i F � CBS � I M�E � � BFNEE� O�� � EN� Y� Y� FM� �M� M   � � � � � � � � � � � � � � � � � � � � � � N� � � � C� � ENI �e � �   � � 1 B� � �

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�O OB I S � I M� TI � � C� M   � � � B� � � C� I M� O� � I � CBNOe� � TC� �9 � C� � � � � � � � 9� ET� O� 1 � C� � E�C � C� M   �I �

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OMCT� MTC� � N� � M   � � � �M�� O� MO� 1 NTE� � �E ON�   �9 � �   �� � �� E�C � � BS F �� Me� �

�   � � FCNFNCMBNI � N� � E�S � O� 1 BM   � � NCC� � MEs� B� � I MB� B� � � � �S O� � E�O OB� B� � � � � F � I � BI � NI � M   � �

� 1 � � 1 BM   �   B   � OM� FCN� �� BEBMs� N� � � �S � �O OB I S � I Mo� �I � � BMO� MC� I � o� 1 � C� � OMC�B   M� NC1 �C � Es �

�   � C�� M� CBm� � � � s � FENMMBI � O� I OBMB9BMs� � �B I OM� M   � � FCN� �� BEBMs� N� � � �S � �O OB I S � I M� n� B TC� O�

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nO� � I �C BNO� � � l o� � � a� � �I � � � � bt� � B TC� O� f eao� f eb�� �I � � f ef � g� TF � MN� pbeu4 � nO� � I �C BN� � � b� t� � � � E� �

f ebge� M� BO� BS FNCM�I M� MN�   B   EB   M� M   � � C� S �C Y�� E� � F � C� NCS �I � � � N� � NTC� �I �E sMB� �E � �F FCN��   �

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� RT�E � MN� NC� E� C � C� M   �I � yc4 e� �   � � F � C� � I M� � � N� � �I � EsO� O� S �M�   BI � M   BO� yc4 � O� I OBMB9BMs� 1 �O �

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f ebge� I � � I s � � �O � o� M   � � C� OTEMO� � CNS � M   � O� � O� � I �C BNO� C� 9� �E � � � �� 9� Cs � �F F � �E BI � F � C� NCS �I � � �

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� � � M� � � f ef � TS � � C� N� � CB   M� n� E�� Y� � NETS I Og� �I � � 1 CNI � n1   BM� � � NETS I Og� � �S � �O OB I S � I MO�nE� � M� � � �i BOgo� �I � � M   � BC� OT� O� RT� I M� O� I OBMB9BMs� nCB   M� � � �i BOgo� � NC� �� � I � NI � � � E�S � O� 1 BM   � M1 N�M1 BI r� BCM   � � �I � B� � M� � � �S O� 1   � I � M1 N� n�t � O� � I �C BN� � � ago� M   C� � � n� t� O� � I �C BN� � � bg� � I � � �E E� � NTC�E� S � O� n� t� O� � I �C BN� � � ) g� 1 � C� � �� �I � NI � � e� NM� � M   �M� E�S � O� 1 � C� � � E�O OB� B� � � � � F � I � BI � NI �M   � � � 1 � � 1 BM   �   B   � OM� FCN� �� BEBMs� N� � � �S � �O OB I S � I M� n� � �i BOge�

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TI � � COM�I � � M   � � � � � � � M� M   �M� � ��   � 9�C B�� E� �   � � � NI � M   � � � NCC� � M� � �S � � OOB I S � I Me�

� 9� CM   � E� OOo� M   � � S N� � E� � NTE� � � �O BEs� � � � � i M� I � � � � MN� � NS F�C � � �E E� N� � M   � � �� �I � NI � � �

I � 1 � NCI O� �M� M   � � O�S � � MBS � o� EBY� � M   � � S N� � E� � s � � �F F � � M� � Ee� naccuge� � TC� OBS TE� MBNI O� O   N1 � � �

M   � M� TI � � CM�B I � S �M� CI BMB� O� � NTE� � � � � �F FCNFCB�M� Es � S N� � E� � � TI � � C� �� � �s � OB�I � � FFCN��   � � s �

M�Y BI � �� 9�I M� � � N� � OsOM� S �MB� � n� e eo� � � � N� � M   � � � �S � �I � � MsF � � N� � � BCM   g� �I � � � I � MB� � n� e e�

� C� � � BI � 9�E T� g� ONTC� � O� N� � BI � NCS �MBNI e� �   BO� S � M   N� � � NTE� �   � EF � FC� 9� I M� � �B I OM�

BI � NCS �MBNI � ENOO� �I � � � I   �I � � � M   � � �� � TC�� s � N� � �O � � CM�B I BI � S �M� CI BMs� BI � EB9� OMN� Y�

FNFTE�MBNI Oe� �   � � � TCC� I M� � � 9� ENFS � I M� S TOM� I NM� � � � 9B� 1 � � � �O � �� OM�MBOMB� �E � MNNE� � NC� M   � �

S � OOB9� � C� � N9� Cs � N� � TI YI N1 I � S �M� CI BMB� Oe� I � � � � o� M   � � F � C� � I M� � � N� � TI YI N1 I �

S � M� CI BMB� O� BI � M   � � � TCC� I M� � i M� I OB9� � O   � � F � BI � TOMCs� BO� S N� � C�M� � MN� EN1 e� � 9� CM   � E� OOo� M   BO�

�F FCN��   � FCN9B� � O� �� OM�MBOMB� �E � S � M   N� NEN s � � NC� BI � � CCBI � TI YI N1 I � S �M� CI BMB� O� �M� �� 9� Cs �

EN1 r� NOMo� � � BI � TO� � TE� �O � �� FC� EBS BI �C s � OM� F � � NC� B� � I MB� sBI � C� FE�� � S � I M� � �I � B� �M� O� MN� � � �

� NI � BCS � � � � s � S �C Y� C� � I NMsFBI o� �I � � � NC� � BO� �C � BI � E�S � O� 1 BM   � �� EN1 � FCN� �� BEBMs� N� �

� � � NS BI � �� C� FE�� � S � I M� � �I � B� �M� � B9� I � BMO� �O OB I � � � � �S � �I � o� �O � �� � NI O� RT� I � � o� BMO�

C� OTEMBI � � C� � � BI � 9�E T� e� � EM   NT   � �I s � C� OTEM� S TOM� � � � � NCCN� NC�M� � � � s � � I NMsFBI � M   � �

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1 NTE� � � BO� �C � � O� 9� C�E � � �I � B� �M� O� � CNS � O� E� � MBNI o� M   TO� C� � T� BI � TI I � � � OO�C s � � I NMsFBI �

� NOMO� � I � � M   � � � NI O� RT� I M� � � NI NS B� � 1 �O M� e

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S � M� CI BMB� O� BI � O   � � F � 1   � I � �� C� � T� � � � I TS � � C� N� � NCF   �I � � � E�S � O� 1 � C� � BI 9NE9� � e� �   BO�

S TOM� � � � 9B� 1 � � � �O � �� TO� � TE� �I � � � NOMr� � � � � MB9� � �F FCN��   � MN� �� RTBC� �   B� � � I � BI � NCS �MBNI �

� CNS � M   � � F   � I NMsFB� � � �M�o � FCN9B� BI � C� E� 9�I M� � �M�� MN� M   � � OMN� Y� C� � � � COe� � EM   NT   �

S � M� CI BMB� O� S TOM� � � � � NI � BCS � � � � s � � I NMsFBI o� M   BO� FC� EBS BI �C s � O� C� � I BI � � s � S Bi � � � S N� � E�

�I �E sOBO� S TOM� �E EN1 � MN� � N� TO� � I NMsFBI � � � � NCMO� NI � M   � � S NOM� BI M� C� OMBI � E�S � Ot� � e eo�

� I NMsFBI � � NC� C� FE� � � S � I M� � �I � B� �M� O� � NTE� � � � � C� OMCB� M� � � MN� M   � � BI � � CC� � � N� � OFCBI � � CNS �

� EBM� � � 1 � Oe� �

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Capítulo  6  

REAL-­‐TIME  ULTRASOUNDS  IN    LIGHT  LAMBS  

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R � � -­‐ I L� R � ET-­‐ � � E � � � � T� � R � -­‐ �

ETLA� I � T� AE�

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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 �

n � �s 1s � kSq� le� Sñ� a � m1u� � � � B� ei � �� l � i � �� c   ei l � � �l l � s 1s � y � i 1e� � kñq� le� Pq� a � m1u� � � � B)�

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�s � � � � mm�d � 2� ñqq4B� � s l1� 1y �l � � l   � � s � � � � � ei � �� � di �l � � l eemc� le� � u�md�l � � � �i � �c c� � eb yec1l1es �

�s � � � es � eib �l 1es2� � u� s � 1s � �m1u� � 1s � 1u1� d�mc)� � � �mwl1b � � dml i �c eds � � l � �   s 1pd� c�   �u � � � � � s �

cd� � � cc� dmmg� dc� � � l e� yi � � 1� l � � e� g� � eb yec1l1es � 1s � m1u� cle� a� k � i i 1s � � l � � m)2� S889B)� �   � g�

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yi e� d� l � �s � � yi eu1� 1s �   1   mg� u�md�� m� � � �l �) � � ml   ed   � ceb � � cld� 1� c�   �u � � � u�md�l � � � l   � �

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� l � � m)2� ñqq90� � �   1s � � l � �m)2� ñqq90� � 1yemm� � l � � m)2� ñqq80� �   � i 1�d ml � � l � �m)2� ñqq8B2� � es l i �� 1� l ei g�

i � cdmlc� n � i � � i � yei l � � � 1s � m1   l � m�b � c)�

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i � 1es � k� �c � mm�c � � l � �m)2� ñqqH� B)� l � 1c� �� i dcl1� � b � � lwlgy � � � i � � � � n 1l   � � es u� v� yi e� 1m� 2� n   1l � �

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l   � � hy �c � d�mMwlgy � � m�b � � �   �i �� l � i 1c� � � � g� m1   l � � � � �l � cm�d   l � i � kSP)S� a � 6� q)S� a � e� � � �i � �c c�

n � 1   lB2� l   1c� � � 1s � es � � e� � l   � � b �1 s � l gy � c� e� � m�b � c� yi e� d� � � � 1s � l   � � � � 1l � i i �s� �s � i � 1es )�

� �b � c� n � i � � n � �s � � � �l � �s � � � � e� � �y yi ev1b �l � mg� Pq� � � �s � �   edc� � � 1s � eei c� kzñ� m�b � cfb ñB0�

l   � g� n � i � � � � � � n 1l   � y � mm� l � � � � es � � s l i �l � � � 1� l � kSR)S4D � � � 0� SS)H� � fa � e� � � B� �s � �   � g� � � �

� � � � � � 0� �s 1b �mc�   � � � � i � � � �� � � cc� l e� � i � c   � n �l � i )� �   � g� n � i � � cm�d   l � i � � � n   � s � i � ��   1s � ñ4�

le� ñ9� a � e� � m1u� � � � � kñ-­‐ )8� a � 6� q)S� a B� �l � �s � �y yi ev1b �l � � � � � e� � P� b e)� �

� � � � � � � � � � � � � n � � � � � � � � � � � � � � � � � � � � � � � � � o� � 1u� � � � � �s � � dml i � ceds � �

1b � � c� n � i � � i � � ei � � � � 49� le� Hñ�   � � � � ei � � cm�d   l � i � �s � � � g� l   � � c�b � � � vy � i 1� s � � � 2� n � mmw

l i � 1s � � � l � �   s 1� 1�s )� � � s l1� �m� � �l �� � emm� � l 1es � yi e� � � di � c� n � i � � �y ym1� � � l e� �mm� � i � � � c� �s � �

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� SSP� �� �

c� v� c)� � s � � vy � i 1� s � � � � �c c1cl �s l � i � cl i �1 s � � � l   � � m�b � � � g� a� � y1s � �� � s lm� � yi � ccdi � � n 1l   � l   � �

m� � l �   � s � � ds � � i � l   � � L�n � l e� yi � u� s l � � ei n �i � � b eu� b � s l2� �s � � ym�� 1s � l   � � i 1   l � � i b � �i eds � �

m� b � Mc� � �� a� l e� cley � � �� an �i � � b eu� b � s l )� � ml i �c eds � � 1b � � c� n � i � � e� l �1 s � � � dc1s � �� � � l �

S9q� � mdc� � pd1yb � s l � k� es e� 1l � 2� s � )2� � el   � mm2� � � 2� � � � B� �s � � �� � -­‐ q� m1s � �i � yi e� � � e� � ñ� l e� R�

o� � pd1yy � � � n 1l   � �� � es u� v� l i �s c� d� � i � k� es e� 1l � 2� s � )B)� � i � u1edc� cld� 1� c� � u1� � s � � � � l   �l �

l   � � dc� � e� � m1s � �i � ei � � es u� v� l i �s c� d� � i c� � 1� � s el � 1s � md� s � � � b � �c di � b � s l c� e� � � �i � �c c� l i �1 l c�

�s � � � es u� v� l i �s c� d� � i c� � u� s � yi e� d� � � � � � l l � i � � � � 1s 1l1es � e� � m�l � i �m� b dc� m� � � ei � � i c�

k� ei cm� g� � l � �m)2� ñqSñB)� � mm� i � � mwl1b � � dml i �c eds � � 1b � � c� n � i � � l �a � s � � l � l   � � i 1   l � c1� � � e� � l   � �

m� b � � 1s � ln e� � 1� � � i � s l � �s �l eb 1� �m� me� �l 1es c2� �l � l   � � l   ei �� 1� � k� � 0� � � l n � � s � l   � � Sñl   � �s � �

SPl   � i 1� cB� �s � � mdb � �i � k� � M� � � l n � � s � l   � � Scl � �s � � ñs � � mdb � �i � u� i l � � i �� B� m� u� mc)� �   1c�

�mmen � � � l e� � eb y �i � � dml i �c eds � � y � i � ei b �s � � � 1s � � 1� � � i � s l � �s �l eb 1� �m� me� �l 1es c� 1s � ei � � i � l e�

1� � s l1� g� �s � eyl1b db � me� �l 1es � le� � u�md�l � � � �i � � cc� �s � � � �l s � cc� l i �1 l c� 1s � m1   l � m�b � c)� � el   �

l i � s cu� i c�m� k� � B� � s � � mes 1ld� 1s �m� k� � B� u1� n c� n � i � � � emm� � l � � � �l � � ��   � �s �l eb 1� �m� me� �l 1es )�

ei � � cy � � 1� 1� �mmg2� � � � 1b � � c� n � i � � l �a � s � y � i y � s � 1� dm�i � l e� l   � � u� i l � � i �m� � emdb s � k� 1 di � �

-­‐ )SB2� � eu� i 1s � �mm� l   � � � � � � � � � � � � � � � � � � � � b dc� m� 2� n   1m� � � � � 1b � � c� n � i � � l �a � s � y �i �mm� m� le� l   � �

u� i l � � i �m� � emdb s � k� 1 di � � -­‐ )ñB2� 1s � l   � � b 1� � m� � e� � l   � � l el �m� m� s l   � e� � l   � � l i �s cu� i c� �

�y ey   gc1c)�

� � � I L� � B  O� � i � s cu� i c�m� i � �mwl1b � � dml i �c eds � � 1b � � )�

� ei � � � � 1b � � c2� �s � � s � �   e � s 1� � cdy � i � m�� � cl �s � e� � � k� es e� 1l � 2� s � )B� n �c � s � � � � � � l e�

� s cdi � � yi ey � i � � 1l � e� � l   � � l i �s c� d� � i � l e� l   � � � di u�l di � � e� � l   � � �s 1b �mMc� s �l di �m� � e� g� c   �y � 0�

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� SS4 � �� �

�� edcl1� � � m� n �c � dc� � � l e� yi eu1� � � � � l l � i � � es l �� l � � � l n � � s � l   � � yi e� � � �s � � l   � � cdy � i � m�� �

cl �s � e� � )� �   � � i 1   l � ym�� � b � s l � � ei � c� �s s 1s � n �c � � � l � i b 1s � � � � g� y �my �l 1s � l   � � i 1   l � c1� � � e� �

l   � � m�b � 0� �� l � i � l   �l 2� n eem� � 1� i � c� n � i � � c� y �i �l � � � n 1l   � �� � eb � )� � � � l �� m� � e1m� n �c � �y ym1� � �

l e� l   � � ca1s � �c � � edym1s � b � � 1db )� � i � ccdi � � es � l   � � l i �s c� d� � i �   � �� � n �c � a� yl � l e� �� b 1s1b db �

le� � ue1� � � eb yi � cc1es � e� � � el   � � �l � �s � � b dc� m� )� � ml i �c eds � � 1b � � c� n � i � � i � � ei � � � � � ei � m�l � i �

u1� n 1s � �s � � � s �mgc1c)�

� � � I L� � B  N� � es 1ld� 1s �m� i � �mwl1b � � dml i �c eds � � 1b � � )�

� ei � � ��   � � � � 1b � � � �l � � el   � � � � �s � � � � � me� �l 1es c2� ca1s � l   1� as � cc� k� � B2� cd� � dl � s � edc�

� � � aw� � l � l   1� as � cc� k� � � B2� �s � � � � y l   � k� � � B2� n 1� l   � k� � � B� �s � � �i � � � k� � � B� e� � � � � � � � � � � � � �

� � � � � � n � i � � e� l �1 s � � � n 1l   � l   � � b � � � � S)4ñp� ce� ln �i � � k� 2� ñqq8B� es � l   � � � �c 1c� e� � l   � �

� � m1y � i � �s � � i dm� � e� � l   � � � � l � S9q� � mdc� � pd1yb � s l � k� es e� 1l � 2� s � )B)� b � � c� n � i � � s el � yi � w

l i � �l � � � s ei � � es u� i l � � 0� � � � � �s � � � � � � n � i � � e� l �1 s � � � n 1l   � l   � � cl i �1   l � eyl1es � e� � l   � �

b � � � � ce� ln �i � � n   � i � �c � � � � � n �c � � �m� dm�l � � � n 1l   � l   � � � i � �   �s � � c� m� � l 1es � leem)� � el � � l   �l �

� � � � � s � � � � � � n � i � � l   � � b �v 1b db �   � 1   l � �s � � n 1� l   � e� � l   � � me1s � b dc� m� 2� n   � i � �c � � � � �s � �

� � � � n � i � � � u�md�l � � � eu� i � l   � � b �v 1b db � b dc� m� � � � y l   )� � s � l   � � el   � i �   �s � 2� � � �

b � �c di � b � s l c� n � i � � l �a � s � 1s � l   � � b 1� � ye1s l � � � l n � � s � l   � � Sñl   � �s � � SPl   � i 1� c� k� � � me� �l 1esB�

ei � � � l n � � s � l   � � Scl � �s � � ñs � � mdb � �i � u� i l � � i �� � k� � � me� �l 1esB)� � ei � � ��   � � � � 1b � � � �l � � el   � � � �

�s � � � � � me� �l 1es c2� � � 2� � � � � �s � � � � � � e� � � � � � � � � � � � � � � � � � � � n � i � � � mce� e� l �1 s � � � dc1s � l   � � c�b � �

ce� ln �i � � k b � � � � S)4ñp0� � 2� ñqq8B)�

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� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � i � � � l n � � s � 49� �s � � Hñ�   � �� l � i �

dml i �c eds � � b � �c di � b � s l2� m�b � c� n � i � � � �c l � s � � � �s � � cm�d   l � i � � � 1s � �� � eb b � i � 1�m�

cm�d   l � i   edc� � k� �i s c� � 1d� � � 2� � �m� � c� � � � es l � d12� � �i � � mes � 2� � y �1 sB)� s � � � � 2� m�b � c� n � i � �

� m� � l i 1� �mmg� clds s � � 2�   ds � � g�   1s � � m� c2� cld� a� 1s le� l   � � l   i e�l 2� ca1s s � � � � g� � 1cl1s � �s � �

� u1c� � i �l � � )� � �i � �c c� c� n � i � �   ds � � g� l   � � � �   1mm� c� l � s � es 2� 1� � s l1� 1� � 2� n � 1   � � � �s � � �   1mm� � �

�l � 4� ( � � � ei � ñ4�   )� � � l � i � l   �l 2� �   1mm� � � � � i � � cc� c� n � i � � n � 1   � � � �s � �   �mu� � )� � d� � dl � s � edc�

� � � aw� � l � l   1� as � cc� �s � � � � � 2� � � � � �s � � � � � � n � i � � l �a � s � es � l   � � i 1   lw   � m� � � �i � �c c� �� l � i �

i 1� � 1s � 1s � l   � � c�b � � �s �l eb 1� �m� me� �l 1es c� n   � i � � dml i �c eds � c� n � i � � yi � u1edcmg� i � 1cl � i � � �

k1)� )2� � � � �s � � � � B)� ei � � cy � � 1� 1� �mmg2� � � � 2� � � � � � s � � � � � � n � i � � b � �c di � � � n 1l   � �� i dm� i 2� n   1m� �

� � � � n �c � b � �c di � � � n 1l   � �� ym�s 1b � l � i � �� l � i � l i �� 1s � l   � � edl � i � y � i 1b � l � i � e� � l   � � � � � � � � � � � � � �

� � � � � � es � �� � l �l � � y � y � i )� � ei � � � �   � 1s � 1u1� d�m2� �mm� b � �c di � b � s l c� n � i � � i � y � �l � � � l   i � � � l 1b � c�

�s � � � u� i � � � )� �

� � � � � � � � � � � � L � � � � � � � � � � � � L � � � � � � � � � � � � � � � � � � � � � � � � � � �

� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � i � � � m1�� 1m1lg� e� � � � � � � � � � dml i �c eds � � b � �c di � b � s l c� 1s �

m1   l � m�b � c� n �c � � u�md�l � � � � g� l n e� 1s � � y � s � � s l � cl � l 1cl1� �m� �y yi e��   � c)� �   � � 1s l � s c1lg� e� � l   � �

i � m� l1es c   1y � � � l n � � s � dml i �c eds � � �s � � � � i � � cc� b � � cdi � b � s l c� n �c � b � �c di � � � � g� l   � �

� � �i ces � yi e� d� lwb eb � s l � � ei i � m�l 1es � � e� � � 1� 1� s l � k� B)� � �m� dm�l 1es c� n � i � � y � i � ei b � � �

� � l n � � s � dml i �c eds � � �s � � � 1i � � l � � �i � �c c� b � �c di � b � s l c� � ei � � � � 2� � � � 2� � � � � �s � � � � � � n 1l   �

l   � � � ei i � m�l 1es � yi e� � � di � � e� � � � � � u)8)S� k� � � � s cl )2� s � )2� � �i g2� � � B)� � � �u �1 m�� m� 2� 1s � � y � s � � s l �

� e� � � 1� 1� s l c� n � i � � � eb ydl � � � es � l   � � � �c 1c� e� � l   � � � 1� � � i � s l � �s �l eb 1� �m� me� �l 1es c� k1)� )2� � � � �s � �

� � B� �s � � dml i �c eds � � u1� n c� k1)� )2� � � � �s � � � � B)�

� i � � 1� l 1u� � �� 1m1lg� e� � dml i �c eds � � � �l �� es � � 1i � � l � � �i � �c c� b � �c di � b � s l c� n �c �

� u� md�l � � � � g� ds 1u�i 1�l � � m1s � �i � i � i � cc1es � � s �mgc1c)� � �m� dm�l 1es c� n � i � � �y ym1� � � � ei � � ��   �

� eb � 1s �l 1es � e� � dml i �c eds � � b � �c di � b � s l � k1s � � y � s � � s l � u�i 1�� m� B� �s � � � 1i � � l � � �i � �c c�

b � �c di � b � s l � k� � y � s � � s l � u�i 1�� m� B� n 1l   � l   � � � � i � cc1es � yi e� � � di � � e� � � � � � u)8)S� k� � � � s cl )2�

s � )B)�

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� SS-­‐ � �� �

� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � i � � ml   ed   � � ei i � m�l 1es � �s � �

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� �

dml i �c eds � � b � �c di � b � s l c� �s � � �mmen � � ei � cl i �1   l � ei n �i � � � eb y �i 1ces � n 1l   � yi � u1edc�

cld� 1� c� 1s � m�b � c2� � � b ei � � � � l �1 m� � � �s �mgc1c� n �c � 1b ym� b � s l � � )� � emmen 1s � �   � 1m� kS8-­‐ -­‐ B� �s � �

�   � i 1�d ml � � l � � m)� kñqq8B2� l   � � � i i ei � e� � yi � � 1� l 1es � � ei � dml i �c eds � � b � �c di � b � s l c� �c � n � mm� �c � 1l c�

� 1� � � i � s l � � eb yes � s l c� n � i � � � cl1b �l � � )� � i i ei c� n � i � � � u�md�l � � � � ei � � edi � dml i �c eds � �

u�i 1�� m� c� k1)� )2� � � � 2� � � � 2� � � � � �s � � � � � B� �l � l   � � l n e� �s �l eb 1� �m� me� �l 1es c0� � � � n �c � s el �

� u� md� l � � � � � � �d c� � l   1c� dml i �c eds � � b � �c di � b � s l � � edm� � s el � � � � l �a � s � � i eb � l   � � � �i � �c c� c)�

�   � � l el �m� b � �c di � b � s l � � i i ei � � pd�mc� l   � � b � �s � cpd�i � � yi � � 1� l 1es � � i i ei � k� � � � B2� n   1�   � n � c�

� � m� dm�l � � � � g� � u� i � 1s � l   � � cpd�i � � � � 1� � � i � s � � � � � l n � � s � � �i � �c c� k� � B� � s � � dml i �c eds � � k� � B�

b � �c di � b � s l c� k � i i 1s � � l � �m)2� S884B2� �

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n   � i � � � � n � c� l   � � l el � m� s db � � i � e� � 1s � 1u1� d�mc� � es l i 1� dl1s � � �l �) � � e� �   �i �� l � i 1c� � l   � �

� 1� � � i � s l � yel � s l1�m� cedi � � c� e� � i � c1� d�m� u�i 1� � 1m1lg2� � � � � � �s � � � � � � � eb yec� � � 1s le� l   i � � �

1s � � y � s � � s l � � eb yes � s l c� �c � yi eyec� � � � g� � � s �   ��i � � l � �m)� kS889B� �s � � � eb �i � �s � � �i � edv�

kñqqRB2� 1)� )2� SB� � i i ei � 1s � � � s l i �m� l � s � � s � g� k� � � B2� ñB� � i i ei � � d� � l e� i � i � cc1es � k� � B2� �s � � PB� � i i ei �

� d� � l e� � 1cldi � �s � � c� k� � B)� ei � � cy � � 1� 1� �mmg2� � � � � n �c � � �m� dm�l � � � �c � l   � � cpd�i � � � 1� � � i � s � � �

� � l n � � s � �u � i � � � � �i � �c c� k CSB� �s � � dml i �c eds � � k USB� u�md� c� � ei � � ��   � b � �c di � b � s l2� �

( )2USCSECT = �

�s � � �   �i � � l � i 1c� � � l   � � � 1�c � � � l n � � s � � el   � i � � ei � 1s � cgcl � b c2� n   � i � �c � � � � b dcl � � � � u1� n � � �

�c � l   � � � � y �i ldi � � � i eb � �� ds 1lg� cmey� � �� l � i � cl �s � �i � 1c1s � � �l �� � g� � eb � 1s 1s � l   � � cl �s � �i � �

� � u1�l 1es � e� � l   � � � �i � �c c� k CSSD B� �s � � dml i �c eds � � k USSD B� b � � cdi � b � s l c� �s � � l   � �

� ei i � m�l 1es � � e� � � 1� 1� s l � � � l n � � s � � � �s � � � � b � �c di � b � s l c� k� B2�

( )[ ]2USCS SDrSDER = �

�   � � m� cl � � eb yes � s l � k� � B� n �c � � �m� dm�l � � � �c �

( ) 221 USSDrED = 2�

�s � � �� � eds l � � � � ei � l   � � � i i ei � l   �l � � �s s el � � � � � vym�1 s � � � � g� l   � � i � i � cc1es2� i � yi � c� s l1s � l   � �

� 1cy � i c1es � e� � l   � � ye1s lc� �i eds � � l   � � i � i � cc1es � m1s � � k� � s �   ��i � � l � �m)2� S889B)� � c� s el � � � � g�

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�   � i 1�d ml � � l � �m)� kñqq8B2� l   � � l i d� s � cc� e� � l   � � b � � cdi � b � s l � 1c� �   �i �� l � i 1o� � � �g � � �� ; � � 2�

n   � i � �c � l   � � yi � � 1c1es � 1c� � u�md�l � � � � g� l   � � � � )� �

� � y �i l � � i eb � l   � � yi � u1edc� cl �l 1cl1� �m� y �i �b � l � i c2� n � � �mce� � �m� dm�l � � � l   � � cl �s � � i � �

� i i ei � e� � yi � � 1� l 1es � k� � � B� � ei � � � � 2� � � � 2� � � � � �s � � � � � � l i �1 l c2�

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� u� i � � � u�md� c� � ei � � � � 2� � � � � �s � � � � � � e� � � � � � � � � � � � � � � � � � � � n � i � � c1b 1m�i � l e� l   ec� �

i � yei l � � � 1s � m1   l � m� b � c� � g� � � i s �s � � o� � l � �m)� kS88HB� �s � � � 1yemm� � l � �m)� kñqq8B� �l � l   � � � � �

me� � l 1es � k� �� m� � -­‐ )S� l e� -­‐ )PB2� �ml   ed   � l   � c� � �d l   ei c� i � yei l � � � i � �l � i � u�i 1�� 1m1l1� c2� � u� s �

n   � s � � e� dc1s � es � m1   l � i � � �i � �c c� c)� � � �i � 1s � l   � � � � � 2� hy �c � d�mMwlgy � � m� b � c� dc� � � 1s � l   1c�

cld� g� n � i � � m� �s � i � l   �s � c1b 1m�i � 1s � 1u1� d�mc� � i eb � el   � i � � y �s 1c   � c   � � y � � i � � � c� cd�   � �c � � � c��

� i � es � c� � k� � m� �� � l � �m)2� S88R0� � 1yemm� � l � �m)2� ñqq8B2� �   di i �� � � s c1s �� k� 1yemm� � l � �m)2� ñqSqB� �s � �

�s �   � e2� � i 1s e� �s � � m� � � � � � i �s � � � F � � i 1s e� k� � i s �s � � o� � l � �m)2� S88H0� � 1yemm� � l � �m)2� ñqq8B)�

� � �s � � �i � �c c� c� �i � � e� � cy � � 1�m� 1s l � i � cl � � ei � l   1c� a1s � � e� � m�b � � yi e� d� l1es2� yi eu1� 1s �

�� � 1l1es �m� � � es eb 1� � u�md� � l e� m�b � � yi e� d� l1es � 1s � � � 1l � i i �s � �s � �i � �c � n   � i � � m� �s � i �

�s 1b �mc� �i � � y i � � � i i � � � k� m� es ce� � l � �m)2� ñqqS0� � es l � 1� � di s emc� � l � �m)2� ñqq-­‐ � �s � � � � 1v� 1i �� � l � �m)2�

ñqq-­‐ B)�

� 1i � � l � � �i � �c c� b � �c di � b � s l c� � 1� � s el � yi eu1� � � c1 s 1� 1� �s l � � � y �i ldi � c� k� � 7� q)qRB�

n   � s � � eb y �i 1s � l   � � l n e� �s �l eb 1� �m� me� �l 1es � k1)� )2� l   ei �� 1� � u� i cdc� mdb � �i 0� � �� m� � -­‐ )PB2� �s � �

l   � � c�b � � y � l l � i s � i � u� �m� � � � ei � dml i �c eds � � b � �c di � b � s l c� k� �� m� c� -­‐ )S� � s � � -­‐ )ñB)� �   � s �

� eb y �i 1s � � 1i � � l � � �i � �c c� �s � � dml i �c eds � � �u � i � � c2� c1 s 1� 1� �s l � � � y �i ldi � c� k� � z � q)qRB� n � i � �

� 1c� �i � � � � � ei � �mm� l i �1 l c� n 1l   � l   � � esmg� � v� � yl1es � e� � � � � )� l � 1c� 1b yei l �s l � l e�   1   m1  l � l   �l � i � �mw

l1b � � dml i �c eds � � ds � � iw� cl1b �l � � � � � � � k� � z � q)qRB� 1s � � mm� �s �l eb 1� �m� me� �l 1es c� �s � � yi e� � �

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� l � �m)� kñqq8B� i � m�l � � � l   1c� ds � � i � cl1b �l 1es � le� l   � � yi � ccdi � � e� � l   � � y i e� � )� �   � � i � b �i a�� m� �

ds � � i � cl1b � l1es � e� � � � � � cd � cl � � � �� m1b 1l � � � �y ym1� �� 1m1lg� e� � i � �mwl1b � � dml i �c eds � c� 1s �

y � c� d�mwlgy � � m1   l � m�b � c� � ei � � u�md�l 1es � e� � � �l s � cc� � � i � � 2� l   1c� � � 1s � �� � �c 1� � � i 1l � i 1es � � ei �

pd�m1lg� �s � � yi 1� � � 1s � � � 1l � i i �s � �s � � eds l i 1� c� k� �r d� e� � l � �m)2� ñqqqB)�

� � � � � B  O� � �s 2� cl �s � �i � � � � u1�l 1es � k� � B2� b �v 1b db � �s � � b 1s1b db � e� � l i �s cu� i c�m�dml i �c eds � � b � �c di � b � s l c� �l � � el   � l   ei �� 1� � �s � � mdb � �i � m� u� mc� ks � á� Sñ40� �u � i � � � � � � á�ñ-­‐ )98� 6� S)ñ4� a B)�

� i � 1l S� � �s � � � � 1s 1b db � �v 1b db �

�   ei �� 1� � m� u� m� � � � � �

� � � � � 2� b b � S)-­‐ S� q)ñ8� q)8S� ñ)4-­‐ �

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� db � � i � m� u� m� � � � � �

� � � � � 2� b b � S)R9� q)ñ9� S)qH� ñ)ñ-­‐ �

� � � � � � 2� b b � S)HP� q)-­‐ q� q)H9� R)4R�

� � � � � � 2� � b � ñ)P8� q)ñ-­‐ � S)9-­‐ � P)ñP�

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S� a1s � l   1� as � cc� k� � B2� cd� � dl � s � edc� � � � aw� � l � l   1� as � cc� k� � � B2� � � y l   � k� � � B2� n 1� l   � k� � � B� � s � � � i � � � k� � � B� e� �� � � � � � � � � � � � � � � � � � � u� md� l � � � � � l n � � s � Sñl   � � s � � SPl   � l   ei � � 1� � u� i l � � i � � � kl   ei � � 1� � m� u� mB� ei � � � l n � � s � Scl � � s � �ñs � � mdb � � i � u� i l � � i � � � kmdb � � i � m� u� mB)�

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� � � � � B  N� � �s 2� cl �s � �i � � � � u1�l 1es � k� � B2� b �v 1b db � �s � � b 1s1b db � e� � mes 1ld� 1s �m�dml i �c eds � � b � �c di � b � s l c� �l � � el   � l   ei �� 1� � �s � � mdb � �i � m� u� mc� ks áSñ40� �u � i � � � � � � á� ñ-­‐ )98�6� S)ñ4� a B)�

� � i � 1l S� � �s � � � � 1s 1b db � �v 1b db �

�   ei �� 1� � m� u� m� � � � � �

� � � � � 2� b b � S)S9� q)ñq� q)-­‐ 4� S)-­‐ -­‐ �

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� db � � i � m� u� m� � � � � �

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� � � � � B  D� � �s 2� cl � s � �i � � � � u1�l 1es � k� � B2� b �v 1b db � �s � � b 1s1b db � e� � l i �s cu� i c�m� � �i � � cc�b � �c di � b � s l c� �l � � el   � l   ei � � 1� � �s � � mdb � �i � m� u� mc� ks áSñ40� �u � i � � � � � � á� ñ-­‐ )98� 6� S)ñ4� a B)�

� i � 1l S� � �s � � � � 1s 1b db � �v 1b db �

�   ei �� 1� � m� u� m� � � � � �

� � � � � � 2� b b � ñ)Pq� q)-­‐ 9� q)9P� -­‐ )qq�

� � � � � � 2� � b � ñ)4H� q)ñH� S)Hq� P)Sq�

� � � � � � 2� � b � � 4)Rq� q)P8� ñ)-­‐ q� R)4q�

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� db � � i � m� u� m� � � � � �

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� � � � � � 2� � b � � 4)-­‐ q� q)4ñ� P)-­‐ q� R)-­‐ q�

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� � � � � � � � � � � � � � � � � � � � � � � � i � � � �i ces � yi e� d� lwb eb � s l � � ei i � m�l 1es � � e� � � 1� 1� s l c�

� � l n � � s � i � �mwl1b � � dml i �c eds � � �s � � � 1i � � l � � �i � �c c� b � � cdi � b � s l c� �i � � c   en s � 1s � � �� m� � -­‐ )4)�

� 1l   � l   � � esmg� � v� � yl1es � e� � � � � 2� �mm� l i �1 l c� yi eu1� � � �   1   � �s � � c1 s 1� 1� �s l � k� � z � q)qRB�

� ei i � m�l 1es c� n 1l   � � cl1b �l � c� i �s 1s � � � l n � � s � q)-­‐ SS� k� � � 2� mdb � �i � m� u� mB� �s � � q)994� k� � � 2�

l   ei �� 1� � m� u� mB)� � el � � l   �l � � ei i � m�l 1es c� n � i � � c1b 1m�i � k� � 7� q)qRB� � ei � � el   � � � � � s � � � � � me� �l 1es c�

k� �� m� � -­‐ )4B2� � s � �   1   � i � � ei � l i �s cu� i c�m� dml i �c eds � � u1� n c� l   �s � � ei � mes 1ld� 1s �m� u1� n c0�

� 1� � � i � s � � c� �b es � dml i �c eds � � u1� n c� n � i � � c1 s 1� 1� � s l � k� � z � q)qRB2� n 1l   � l   � � esmg� � v� � yl1es � e� �

� � � � �l � l   � � l   ei �� 1� � m� u� m� k� � z � q)SB)� �   � c� � i � cdmlc� cd � cl � l   �l � mes 1ld� 1s �m� u1� n c� � edm� � � � �

e� � m1b 1l � � � 1s l � i � cl � 1s � m1   l � m�b � c2� n   � i � �c � � el   � l   ei �� 1� � �s � � mdb � �i � �s �l eb 1� � m� me� �l 1es c�

b dcl � � � � u1� n � � � �c � �ml � i s � l 1u� � eyl1es c� n 1l   � c1b 1m�i � y � i � ei b �s � � c)� � �

� � � � � B  G� � ei i � m�l 1es � � e� � � 1� 1� s l � � � l n � � s � � 1i � � l � � �i � �c c� �s � � dml i �c eds � � b � �c di � b � s l c� 1s �m1   l � m� b � c)�

� � ml i �c eds � � u1� n �

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� � � � � � db � � i � m� u� m4� q)-­‐ qñWWW� q)PS9WW�

� � yl   � e� � l   � � � � � � � � � � � � � � � � � � � � b dc� m� � � �

� � � � � �   ei �� 1� � m� u� m� q)994WWW� q)-­‐ 8PWWW�

� � � � � � db � � i � m� u� m� q)9RHWWW� q)-­‐ SSWWW�

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� i � �� e� � l   � � � � � � � � � � � � � � � � � � � � b dc� m� � � �

� � � � � �   ei �� 1� � m� u� m� q)HS8WWW� �

� � � � � � db � � i � m� u� m� q)HH-­‐ WWW� �

S� � i y � s � 1� dm� i � l e� l   � � mes 1ld� 1s � m� � v1c� e� � l   � � m� b � )� ñ� � i � mm� m� le� l   � � mes 1ld� 1s � m� � v1c� e� � l   � � m� b � )� P� � l n � � s �Sñl   � � s � � SPl   � l   ei � � 1� � u� i l � � i � � )� 4� � l n � � s � Scl � � s � � ñs � � mdb � � i � u� i l � � i � � )�

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� ei i � m�l 1es � � e� � � 1� 1� s l c� � ei � � � � � n � i � � men � i � l   � s � l   ec� � i � yei l � � � � ei �   � �u 1� i �

�s 1b �mc� kPñ� le� -­‐ P� a 0� � � � � c� � l � �m)2� ñqq90� � �   1s � � l � �m)2� ñqq90� �   � i 1�d ml � � l � �m)2� ñqq80�

� b � s   � 1c� i � � l � �m)2� ñqSq�2 � B)� � 1yemm� � l � �m)� kñqq82� ñqSqB� �mce� yi eu1� � � �   1   � i � � ei i � m�l 1es �

� e� � � 1� 1� s l c� � ei � m1   l � m�b � c� � i eb � �   di i �� � � s c1s �� �s � � � �c �� � i � es � c�� c   � � y � � i � � � c2�

n   � i � �c � edi � � cl1b � l � c� � ei � � � � � � v� � � � � � � l   � � � ei i � m�l 1es � i � yei l � � � � g� � � 1v� 1i �� � l � �m)� kñqq-­‐ B�

�s � � �   � i 1� dml � � l � �m)� kñqq8B)� � ml   ed   � � � � � � ei i � m�l 1es � � e� � � 1� 1� s l c� e� l �1 s � � � 1s � l   1c� cld� g�

n � i � � �mm� c1 s 1� 1� �s l � k� � z � q)qRB2� l   � g� n � i � � men � i � l   �s � � ei � l   � � i � b �1 s 1s � l i �1 l c)� �   1c� � �s � � � �

� d� � l e� i �s � eb � � i i ei � 1s � dml i �c eds � � b � �c di � b � s l c� 1s l i e� d� � � � � g� l   � � b 1s1b �m� yi � ccdi � �

�y ym1� � � es � l   � � l i �s c� d� � i �   � �� � � di 1s � dml i �c eds � � � emm� � l 1es 0� l   � � yi � � 1c1es � e� � l   � �

dml i �c eds � � � pd1yb � s l � � edm� � �mce�   �u � � �� cd� cl �s l1�m� 1b y �� l � 1s � l � i b c� e� � i �s � eb � � i i ei �

n   � s � b � �c di1s � l   1s � l1ccd� c� m1a� � l   � � cd� � dl �s � edc� � �l � m�g � i � 1s � m1   l � m�b � c)� ei � eu� i 2� l   � �

� i � cc1s � yi e� � � di � c2� s �b � mg� l   � � ca1s s1s � �s � � l   � � � dl l1s 2� � � s � m� �� � l e� �ml � i �l 1es c� e� � l   � �

� � � aw� � l � l 1ccd� � n   1�   � � �s � � � � i � cyes c1� m� � � ei � l   � � men � i � � ei i � m�l 1es � � e� � � 1� 1� s l c� e� c� i u� � �

k� � 1v� 1i �� � l � � m)2� ñqq-­‐ B)� �

� ei i � m�l 1es c� � ei � � � � � �s � � � � � � � m� �i mg� � v� � � � � � � l   � � es � c� yi eu1� � � � � g� � 1yemm� � l � �m)�

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c   en � � � � ei i � m�l 1es c� cb � mm� i � l   �s � yi � u1edc� � cl1b � l � c� yi eu1� � � � � g� � � i s �s � � o� � l � �m)� kS88H0�

q)99B2� � 1mu�� � l � �m)� kñqq-­‐ 0� q)8RB� �s � � � �   1s � � l � �m)� kñqq90� q)9ñB)� �   1c� � edm� � � � � m1sa� � � l e� l   � �

yi e� � � � � � �d c� � �mm� l   � c� � cld� 1� c� dc� � � �� y i e� � � e� � i � �l � i � � i � pd� s � g� kR� le� 9� oB2� i � cdml1s �

1s � i � � l � i � 1b � � � pd�m1lg)� � � u� i l   � m� cc2� n � � � eds � �   1   � i � � ei i � m�l 1es c� � � l n � � s � dml i �c eds � �

�s � � � � i � � cc� u�md� c� 1s � geds � i � m�b � c� l   �s � � � � � c� � l � �m)� kñqq90� q)HRB2� � b � s   � 1c� i � � l � �m)�

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�s � � men � iwl   � swes � � � c� c   en s � 1s � � 1 di � c� -­‐ )P� le� -­‐ )-­‐ )� �   � � i � m�l 1es c   1y � � � l n � � s � � �i � �c c� �s � �

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� � y � s � 1s � es � l   � � � s �mgo� � � l i �1 l )� s � � � � 2� � � y �i ldi � c� � i eb � l   � �   gyel   � l 1� �m� 1� � s l1lg� m1s � �

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ds � � iw� cl1b �l 1es � n   � s � b es1lei1s � � g� dml i �c eds � c2� l   ei �� 1� � � � � 2� � � � � �s � � � � � � y i eu1� � � �

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ds � � iw� �s � � eu� iw� cl1b �l 1es c� � ei � cb �mm� �s � � m�i � � u�md� c� e� � � �i � �c c� l i �1 l c2� i � cy � � l 1u� mg� kc� � �

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i � yei l � � � � g� � 1yemm� � l � �m)� kñqq8B� �s � 2� �c � cd � cl � � � � g� l   � b 2� l   1c� � edm� � � � � � d� � l e� yi � ccdi � �

es � l   � � yi e� � � �s � � b dcl � � � � b ei � � 1b yei l �s l � 1s � � �l l � i � m�b � c)� s � � � � 2� ds � � i � cl1b �l 1es � 1s �

� � l l � i � 1s � 1u1� d�mc� n �c � yi � u1edcmg� i � yei l � � � � g� c� u� i �m� �d l   ei c� cd�   � �c � � � � � c� � l � �m)� kñqq9B�

�s � � � 1yemm� � l � �m)� kñqSqB)� �   � c� � i � cdmlc� � i � � � � n 1l   � l   � � men � i � � ei i � m�l 1es � � e� � � 1� 1� s l c�

i � yei l � � � � ei � � � � � k� �� m� � -­‐ )4B� �s � � b dcl � � � � m1sa� � � l e� yi eyei l1es �mmg� i � �l � i � � i i ei c� e� �

b � �c di � b � s l � l   �l � � �s � � � � �s l1� 1y �l � � � � ei � l i �1 l c� �l � cb �mm� � � yl   c� k� eds � �s � � � � � a� i 2� S884B)�

� � b �1 s 1s � l i �1 l c� yi eu1� � � � m�i � i � i � i � cc1es � � e� � � 1� 1� s l c2� �ml   ed   � cd � cl � � � �� cm1   l � � 1�c �

�   � i �� l � i 1o� � � � g� c1 s 1� 1� � s l � k� � z � q)qRB� 1s l � i � � y l � l � i b c� 1s � �mm� � �c � c)� �   � � � ñ� � u1� � s � � � � l   �l �

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b � �c di � b � s l c� � ei � � � � 2� � � � � �s � � � � � � k� 1 di � c� -­‐ )4� le� -­‐ )-­‐ B)� �   � c� � � cl1b �l � c� n � i � � cm1   lmg�

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s � � s � i �m2� edi � � i i ei � � cl1b �l � c� � ei � � � � � �s � � � � � � k� �� m� c� -­‐ )R� � s � � -­‐ )-­‐ B� n � i � � men � i �

l   � s � l   ec� � i � yei l � � � � g� �   � i 1�d ml � � l � �m)� kñqq8B� � ei �   � �u 1� i � k4H� a � e� � m1u� � � e� g� n � 1   lB� �s � �

em� � i � kSPq)8� � B� m� b � c2� � u� s � n   � s � dc1s � �� y i e� � � n 1l   � c1b 1m�i � � i � pd� s � g� �s � � n 1l   edl �

c   � �i 1s � l   � � n eem)� �   � s � � � � � n �c � � � � eb yec� � 2� l   � � � � � � i i ei � n �c � m1sa� � � l e� l   � �

ds � vym�1 s � � � u�i 1�s � � 2� i � yi � c� s l1s � l   � � yei l1es � e� � � � � � l   �l � � edm� � s el � � � � � m1b 1s �l � � � � g�

m1s � �i � � ei i � � l 1es c� �s � � s � � � c� l e� � � � b 1s1b 1o� � � k�   � 1m2� S8-­‐ -­‐ B)� � ei � l i �s cu� i c�m� dml i �c eds � �

1b � � c2� n � � � eds � � l   �l � � � � � es l i 1� dl � � � l   � � b ecl � l e� � � � � k4P)RD � le� 89)qD � e� � l   � � l el �m�

� i i ei 0� � �� m� � -­‐ )RB2� � v� � yl � � � � � kl   ei �� 1� � m� u� m2� Sñ)RD0� mdb � �i � m� u� m2� P9)RDB)� � es 1ld� 1s �m�

dml i �c eds � � 1b � � c� c   en � � � �� c1b 1m�i � l i � s � � k� �� m� � -­‐ B2� n   � i � � � � � � � es l i 1� dl � � � l   � � b ecl � l e�

� � � � � ei � � � � � k7-­‐ RDB� �s � � � � � � � u1� � s � � � � � es l i �� 1� l ei g� i � cdmlc� n   � s � � eb y �i 1s � l   ei �� 1� �

k� � � á� HR)9DB� �s � � mdb � � i � k� � � á� S9)RDB� 1b � � c)� �   � c� � i � cdmlc� y � i l1�mmg� � 1c� i � � � n 1l   �

�   � i 1�d ml � � l � �m)� kñqq8B� n   e� � eds � � men � i � � � � u� md� c� kSSD � �s � � PqD � � ei � � � � � �s � � � � � 2�

i � cy � � l 1u� mgB)� �   � � � 1� � � i � s l � � � � u�md� c� e� l �1 s � � � � edm� � � � � � vym�1 s � � � � g� l   � � � �� l � l   �l �

�   � i 1�d ml � � l � � m)� kñqq8B� dc� � � �� y i � y �i �l 1es � l �� m� � l e� i � cl i �1 s � l   � � m�b � c� �s � � l   � � n eem� � m� � � � c�

n � i � � c   � � i � � )� s � l   � � cld� g� � es � d� l � � � � g� �   � i 1�d ml � � l � �m)� kñqq8B2� l   � � �s 1b �m� b eu� b � s l �

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� di 1s � l   � � � �y ldi � � e� � dml i �c eds � � 1b � � c� 1s � d� � c� l   1c� l gy � � e� � � i i ei 2� �ml   ed   � l   � i � � � edm� �

�mce� � � � el   � i � cedi � � c� cd�   � �c � l   � � i 1� � 1s � yi e� � cc)�

� s gn � g2� l   � c� � i � cdmlc� �s � � l   � � � � y �i ldi � c� n 1l   � �   � i 1�d ml � � l � � m)� kñqq8B� cd � cl � l   �l �

�y yi eyi1�l � � i � cl i �1 s � �s � � c   � �i 1s � e� � l   � � m�b � c� � edm� � � � � e� � cy � � 1�m� 1s l � i � cl � 1� � dml i �c eds � �

b � �c di � b � s l c� �i � � cgcl � b �l 1� �mmg� 1b ym� b � s l � � � � g� l   � � c   � � y � 1s � dcl i g� � ei � � �i � �c c�

� u� md� l1es � e� � m1   l � m� b � c)� � � �

�   � � � � � � i � u� �m� � � �c � �� i � m� u�s l � � es l i 1� dl1es � le� l   � � � � � � kq)q4D � le� Sq)ñD0� � �� m� c�

-­‐ )R� �s � � -­‐ )-­‐ B� n   � s � � e� dc1s � es � � � � � �s � � � � � � kS4)HD � le� 9q)qDB2� l   1c� � � 1s � m� cc� i � b �i a�� m� �

� ei � � � � � �s � � � � � � 1s � � � � u1� n c� kq)HD � le� SR)8DB� �s � � b e� � i �l � � 1s � � � � u1� n c� kS4)HD � le� Hñ)ñDB)�

�   1c� a1s � � e� � � i i ei � �   �i �� l � i 1c� � � l   � � � 1�c � � � l n � � s � � �i � �c c� �s � � dml i �c eds � � b � �c di � b � s l c�

�s � � � �s � � � � � �c 1mg� i � b eu� � � � g� �� � 1s � l   � � u�md� � e� � l   � � � 1�c � l e� l   � � dml i �c eds � �

b � �c di � b � s l )� � c� s el � � � � � � ei � 2� l   � � yi � ccdi � � �y ym1� � � es � l   � � l i �s c� d� � i �   � �� � � di 1s �

dml i �c eds � � � emm� � l 1es � � edm� � � � � y �i l1�mmg� i � cyes c1� m� � � ei � l   1c� a1s � � e� � � i i ei 2� l   1c�   gyel   � c1c�

� i � � 1s � n 1l   � l   � � m�i � cl � � es l i 1� dl1es � es � y � i y � s � 1� dm�i wlewl   � wl i � s c� d� � i � b � �c di � b � s l c�

k� � � � �s � � � � � B� l   �s � l   � � i � b �1 s 1s � es � c)�

� 1s �mmg2� � � � i � yi � c� s l � � � l   � � � � y �i ldi � � � i eb � l   � � ds 1lg� cmey� � n   � s � ymel l1s �

dml i �c eds � � b � �c di � b � s l c� � �1 s cl � � �i � �c c� b � �c di � b � s l c)� �   1c� � i i ei �   �� � �� b e� � i �l � wlew

men � � es l i 1� dl1es � le� � � � � kS)qD � le� ñR)qD0� � �� m� c� -­‐ )R� � s � � -­‐ )-­‐ B2� �s � � i � yei l � � � u�md� c� � ei �

� � � � � s � � � � � � � � mm� n 1l   1s � l   � � i �s � � e� � � cl1b �l � c� yi eu1� � � � � g� �   � i 1�d ml � � l � �m)� kñqq8B� 1s �

� d� � ema� �s � � � ei c� l � m�b � c)� � ml   ed   � l   1c� a1s � � e� � � i i ei � � �s � � � � � ei i � � l � � � � g� i � i � cc1es2� l  � �

� � � 1b ym1� c� l   �l � l   � � � 1�c � 1c� s el � � es cl �s l � �s � � u�i 1� c� � � y � s � 1s � es � l   � � b � s 1ld� � � e� � l   � �

b � �c di � b � s l c� k�   � i 1�d ml � � l � �m)2� ñqq8B)� �

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� � � � � � � eb yes � s l c� �

� i � 1l S� � � � ñ� � � � � P� � � 4 � � R� � � � � -­‐ �

�   ei �� 1� � m� u� m� � � � � �

� � � � � � � � q)qq9� q)qqR� q)qqñ� q)qqS� q)SRñ�

� � � � � � � q)qS9� q)qqR� q)qqñ� q)qSS� q)Sñ9�

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� � � � � � � q)8Pq� q)qPñ� q)S4R� q)HRP� q)8ñR�

� db � � i � m� u� m� � � � � �

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� � � � � � � q)qñP� q)qq9� q)qqR� q)qSq� q)ñS9�

� � � � � � � q)SP4� q)qqS� q)qqP� q)SPq� q)P-­‐ 8�

� � � � � � � q)HHP� q)SñP� q)SP4� q)RS-­‐ � S)q99�

S� d� � dl � s � edc� � � � aw� � l � l   1� as � cc� k� � � B2� � � y l   � k� � � B2� n 1� l   � k� � � B� � s � � � i � � � k� � � B� e� � � � � � � � � � � � � � � � � � � �� u� md� l � � � � � l n � � s � Sñl   � � s � � SPl   � l   ei � � 1� � u� i l � � i � � � kl   ei � � 1� � m� u� mB� ei � � � l n � � s � Scl � � s � � ñs � � mdb � � i �u� i l � � i � � � kmdb � � i � m� u� mB)� ñ � � � 5� b � � s � cpd� i � � y i � � 1� l 1es � � i i ei )� P� � � 5� � i i ei � e� � � � s l i � m� l � s � � s � g)� 4� � 5� � i i ei �� d� � l e� i � i � cc1es )� R� � 5� � i i ei � � d� � l e� � 1cldi � � s � � )� -­‐ � � � 5� cl � s � � i � � � i i ei � e� � y i � � 1� l 1es )�

� c� � � n   em� 2� l i d� s � cc� e� � dml i �c eds � � � �l � � n �c � � u�md�l � � � � g� � � � � �s � � � � 2� � u1� � s � 1s �

b e� � i �l � � � � y �i ldi � c� l   � l � � edm� � � � � � � c1mg� � ei i � � l � � � � g� � yymg1s � �y yi eyi1�l � � � ei i � � l 1s �

� � � l ei c� �s � � i � i � cc1es � yi e� � � di � c2� i � cy � � l 1u� mg)� � s � l   � � � es l i �i g2� yel � s l1�m� cedi � � c� e� �

u�i 1�� 1m1lg� k� ) )2� m�b � � i � cl i �1 s l2� n eem� c   � �i 1s B� b dcl � � � � a� yl � l e� l   � � b 1s1b db � n   � s �

�y ymg1s � dml i �c eds � � l � �   s 1pd� c� 1s � ei � � i � l e� b 1s1b 1o� � l   � � i � m� u�s l � 1b y �� l � e� � � � )� �   1c�

� es � mdc1es � � edm� � � � � e� � cy � � 1�m� i � m� u�s � � � 1s � m1   l � m�b � c� n   � i � � l   � 1i � cb �mm� i � � e� g� c1o� c�

� edm� � ei 1 1s �l � � yi eyei l1es �mmg� i � �l � i � � i i ei � e� � b � �c di � b � s l � � ei � l i � 1l c� � l � cb � mm� � � yl   c�

k� eds � �s � � � � �a � i 2� S884B)� �

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� � � � � B  B  � � � c1� d�m� � � � eb yec1l1es � e� � mes 1ld� 1s �m� dml i �c eds � � b � �c di � b � s l c� � eb y �i � � � l e�� � i � � cc� b � �c di � b � s l c� �l � � el   � l   ei �� 1� � �s � � mdb � � i � m� u� mc)�

� � � � � � � eb yes � s l c� �

� i � 1l S� � � � ñ� � � � � P� � � 4� � � R� � � � � -­‐ �

�   ei �� 1� � m� u� m� � � � � �

� � � � � � � � q)qSR� q)qSñ� q)qqñ� q)qqS� q)S9P�

� � � � � � � q)q8R� q)qS4� q)qq8� q)qHñ� q)S8P�

� db � � i � m� u� m� � � � � �

� � � � � � � q)qñS� q)qS4� q)qqR� q)qqñ� q)S44�

� � � � � � � q)SRS� q)Sq8� q)qS4� q)qñ9� q)PqP�

S� d� � dl � s � edc� � � � aw� � l � l   1� as � cc� k� � � B� � s � � � � y l   � k� � � B� e� � � � � � � � � � � � � � � � � � � � � u� md� l � � � � � l n � � s � Sñl   � � s � �SPl   � l   ei � � 1� � u� i l � � i � � � kl   ei � � 1� � m� u� mB� ei � � � l n � � s � Scl � � s � � ñs � � mdb � � i � u� i l � � i � � � kmdb � � i � m� u� mB)� ñ � � � 5�b � � s � cpd� i � � y i � � 1� l 1es � � i i ei )� P� � � 5� � i i ei � e� � � � s l i � m� l � s � � s � g)� 4� � 5� � i i ei � � d� � l e� i � i � cc1es )� R� � 5� � i i ei � � d� �l e� � 1cldi � � s � � )� -­‐ � � � 5� cl � s � � i � � � i i ei � e� � y i � � 1� l 1es )�

� � � � � � � � � � � � � � � � � � � � � � � � � � � i � �   � � � � � � 1c� � s � �ml � i s �l 1u� � cl �l 1cl1� � � eb b esmg� dc� � � l e�

u�m1� �l � � l   � � �� � di �� g� e� � dml i �c eds � � b � �c di � b � s l c� 1s � el   � i � cy � � 1� c� � dl � s el � 1s � m�b � c�

k� b � s   � 1c� i � � l � �m)2� ñqSq� B)� s � l   1c� cy � � 1� c2� l   � � ds 1pd� � i � cdmlc� �u �1 m�� m� � 1s � l   � � c� 1� s l1� 1� �

m1l � i �l di � � l e� � u�md�l � � l   � � �� � di � � g� e� � dml i � ceds � � b � �c di � b � s l c� � vym�1 s1s � l   � � � 1� � � i � s l �

yel � s l1�m� cedi � � c� e� � i � c1� d�m� u�i 1�� 1m1lg�   �u � � � e� dc� � � es �   � �u g� m�b � c� k�   � i 1�d ml � � l � �m)2�

ñqq8B)� � � � � n �c � �mce� � �m� dm�l � � � �c � �s � �ml � i s �l 1u� � cl �l 1cl1� � l e� � u�md�l � � l   � � y i � � 1c1es � e� � l   � �

dml i �c eds � � b � �c di � b � s l c)� � � � c� � ei � � � � � 1s � l   1c� cld� g� k� �� m� c� -­‐ )R� � s � � -­‐ )-­‐ B� n � i � � cm1   lmg�

i � �l � i � l   �s � l   ec� � i � yei l � � � � g� el   � i � �d l   ei c� k� �1 l � � l � �m)2� ñqqR0� � � � � c� � l � �m)2� ñqq90�

�   � i 1�d ml � � l � �m)2� ñqq80� � b � s   � 1c� i � � l � �m)2� ñqSq� B2� � ml   ed   � � m� �i mg� cb �mm� i � l   �s � l   � �

b � v1b db � u�md� � e� � q)ñR� � b � � cl �� m1c   � � � � g� � �1 l � � l � �m)� kñqqRB� � ei � � �l � b � �c di � b � s l c� 1s �

m� b � c)� �   � � � � � c� � ei � � � � � k� �� m� c� R� � s � � -­‐ B� n � i � � men � i � l   �s � l   � � u�md� c� i � yei l � � � � g�

�   � i 1�d ml � � l � �m)� kñqq8B)� s � l   � � � �c � � e� � � � � 2� l   � � i � cdml1s � � � � � kq)PR� �s � � q)PH� � b 2� � ei � � � � �s � �

� � � me� � l 1es c2� i � cy � � l 1u� mgB� � edm� � s el � � � � � eb y �i � � � c1s � � � l   � i � � �i � � s e� yi � u1edc� cld� 1� c� l   �l �

i � yei l � l   � � � � � � � ei � l   1c� b � �c di � b � s l )� �   � � � � � � � ei � � � � � 1s � l   � � yi � c� s l � cld� g� kq)8P� �s � � S)q8�

� b ñ� � ei � � � � �s � � � � � me� �l 1es c2� i � cy � � l 1u� mgB� n �c � men � i � l   �s � l   ec� � i � yei l � � � � g� � � 1l � � l � �m)�

kñqqRB� kS)8ñ� le� ñ)S9� � b ñB2� � � � � c� � l � �m)� kñqq9B� kS)RR� � b ñB� � s � � � b � s   � 1c� i � � l � �m)� kñqSq� B�

kS)9-­‐ � l e� ñ)ññ� � b ñB2� cd � cl1s � l   �l � edi � dml i �c eds � � � cl1b �l � c� �i � � b ei � � �� � di �l � � l   �s �

l   ec� � e� l � 1s � � � 1s � � � im1� i � i � yei l c)�

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Capítulo  7  

CARCASS  TRAITS  AND  MEAT  COMPOSITION  OF  LIGHT  LAMBS  

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� � � � � �

� � � � � � � � � � �

� CB   CD� B   � � � EDB� i B� E� � B   � � H   � � F � CD� 1HBGt f� � EDH1N � G� FG� � � G� D� � H� BEW�G � H� IC   B� �D � �

IEW� � �B � I�N � H�   �3 � � � � � D� � i B� DHC3� It � G� FEGB� � � CD� HE1B   � GD� � 1GEF� � m� I� EDHE� � B� �I df� . vv( s�

� EDB� C� � 1GDEIH� � B� �I df� . vv6s� � � Ci � CG�f � � � BEHf� � E� GC 1� Hf� � � I� �� �D � � � � � �3 � hf� . vv6wd� �N � H�

�G � � Bt FC� �I It � HI�1   B� G� � � �� B� G� W� �D CD � m�B � �� E1B� ( v � T � IC3� � W� C   Bw� EG� �� B� G� �� H   EGB�

� � BB� DCD � F � GCE� � m. v,2v� T � IC3� � W� C   Bw� CD� H� 3� G�I � �G � �H � E� � B   � � � � � CB� GG�D � �D � � � HCDd�

� EG� E3� Gf� HE� C� Bt rH� � ED� � GD� �� E1B� N � �B � � EN FEHCBCED� �D � � CBH� G� I�B CEDH   CF� WCB   �   1N �D �

  � �I B   �   �H � CD� G� �H � � � � 1GCD � B   � � I�H B� � � W� � � � �� � H� m� � GD�� � 1� �D � � � � D� � GEf� . vv; wf� �D � � CH�

� 1GG� DBIt � � � T� t � FECDB� � EG� B   � � H   � � F � � �G NCD � CD� 1HBGt � m� EGBCN � G� � B� �I df� . v( vwd� � N ED � EB   � G�

BtF � H� E� � N � � Bf� I�N � � N � �B �   �H � � � � D� H1 � HB� � � �H � � � G� I� 3�D B� HE1G� � � E� � EN � � ,2� � �B Bt � �� C� H�

m� d df� IED , �   � CD� FEIt1DH�B 1G�B � � � � � BBt � �� C� H� m� � � � Hw� �D � � H� 3� G�I � � EDS1 �B � � � ICDEI� C� � �� C��

m� � w� CHEN � GHwf� WCB   � � � D� � C� C�I � � � � � � BH� ED� �B B� D1�B CD � � �G � CDE � D� HCHf� � B   � GEH� I� GEHCHf� B   � �

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� �

�I I� G � DC� CBt � B   �D � G� � � N � �B � � GEN � EB   � G� � EN � HBC� � HF � � C� H� m� 1� � �� � B� �I df� . v( ( wd� �   � �

� 3C� � D� � � FGE3C� � � � � t � B   � H� � HB1� C� H� WE1I� � � � � E � � HF � � C�I � G� I� 3�D � � � BE� HBE� T� G� � � � GHf�

�I B   E1   � DEB� �I I� E� � CB� � �D � � � � � CG� � BIt � � i BG�F EI�B � � � BE� �I I� I�N � � FGE� 1� BCED� HtHB� NH� C3� D� B   � �

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�

� � IB   E1   � IC   B� I�N � � FGE� 1� BCED� G� FG� H� DBH� B   � � I�G � HB� H   �G � � E� � B   � � I�N � � N �G T� B� CD�

N � Dt � � � � CB� GG�D � �D � � E1DBGC� H� m� �M1� E� � B� �I df� ( kk) wf� ICBBI� � CH� TDEWD� �� E1B� � �G � �H H�

F � � 1IC�G CBC� H� �D � � N � �B � � EN FEHCBCED� �B � � �G It � � � H� E� � I�N � Hd� � EG� E3� Gf� B   CH� TCD� � E� � I�N � �

FGE� 1� BCED� CD3EI3� H� N 1IBCFI� � H   � � F � � G� � � Hf� CD� I1� CD � �1 BE�   B   EDE1H� BtF � H� H1�   � �H � B   � �

� 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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( 20 � �� �

WCB   � � � WEGI� WC� � � � CHBGC� 1BCEDf� Cd� df� B   � � N � �B ,BtF � � �� �1 D� f� �D � � �� � F �D CH   � �1 BE�   B   EDE1H�

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� �

B� �   DCL1� H� ED� IC3� � �D CN �I H� �D � � � CG� � B� N � �H 1G� N � DBH� ED� B   � � � �G � �H H� �� B� G� HI�1   B� Gs�

� 1GB   � GN EG� f� B   � � � �B Bt � �� C� � FGE� CI� � E� � B   � � � � � � � � � � � � � � � � � � � � N 1H� I� � W�H � �I HE� �   � G�� B� GCh� � d�

�   CH� HB1� t � W�H � � �G GC� � � E1B� BE� FGE3C� � � � � B�C I� � � TDEWI� � � � �� E1B� B   � � � ENN � G� C�I � E1BF1B�

� GEN � B   CH� BtF � � E� � � � � CB� GG�D � �D � FGE� 1� BC3� � HtHB� N � �D � � BE�   C   IC   B� � �D � C� �B � � B�G � BH� � EG�

� 1GB   � G� � G� � � CD � FGE G�N H� CD� H   � � F � � G� � � Hd� � �

� � � � � � � � � � � � � � � � � � �

� II� �D CN � I� FGEBE� EIH� W� G� � �F FGE3� � � � t � B   � � � DC3� GHCB�B � � 1BODEN �� � � � � �G � � IED��

m� � II�B � GG�f � � F �C Dw� � DCN � I� � �G � � �D � � � H� � � ENNCBB� � � mG� � � G� D� � � � � � � ( . 6vw� �D � � W� G� �

� � HC D� � � �D � � � ED� 1� B� � � �� � EG� CD � BE� B   � � 1C� � ICD� H� E� � � E1D� CI� � CG� � BC3� � ) 6y6vky� � ��

m� 1GEF� �D � � ENN1DCBC� Hf� ( k) 6w� ED� B   � � FGEB� � BCED� E� � �D CN �I H� 1H� � � � EG� H� C� DBC� C� � F1GFEH� Hd�

� � � � � � � � � � � � � � � � � � � � �

�   CH� HB1� t � � E� 1H� � � ED� BWE� N � �B ,BtF � � H   � � F � � G� � � Hf� � CFEII� H�� �D � � �� �1 D� f� �H � W� II�

�H � B   � CG� � CGHB, � D� G� BCED� � GEHHd� � N ED � B   � � �1 BE�   B   EDE1H� H   � � F � � G� � � H� CD� � F �C Df� B   � �

� CFEII� H�� CH� B   � � N EHB� �� 1D� �D B� � G� � � � CD� � �B �I EDC�� mCd� df� B   � � DEGB   � �H B� GD� G� CED� E� � � F �C Dwf�

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� � G� � HH� W� C   B� E� � ( 2� BE� ( 6� T d� B� CH� �� G1HBC� � N � �B ,BtF � � � G� � � � WCB   � � ED3� i � FGE� CI� f�

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� W� H� m� � GG� Bf� � E� CD� �D � � � � G� �� C� Gf� ( kk; wd� � � BEB�I � E� � ( . 0� � CFEII� H�� mD� j � z zwf� �� �1 D� � mD� j �

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� GEN � B   � � � �N �� � GC�� � �H BEH� � � IE� T� m� C 1� H� C� � C� IIHf� � �G � � IED� f� � F �C Dw� W� G� � CD3EI3� � � CD� B   CH�

G� H� �G �   d� �

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� � � � � � � � s� � EDBCD1E1H� �� � � HH� BE� � G� H   � W�B � G� W�H � �I HE� 1�G �D B� � � d� �   � � I�N � H� W� G� �

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B   � � uF �H � 1�I r,BtF � � I� N � H� FGE� 1� � � � CD� B   � � � � � CB� GG�D � �D � G� CEDd�

� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �

� WE� � C� � � G� DB� � � � � � � � � � � � � � � � � � 1IBG�H E1D� � CN � � H� W� G� � G� � EG� � � � F � G� I�N � � 0) � BE�

z . �   E1GH� � � � EG� � HI� 1   B� G� � t � B   � � H�N � � BG�C D� � � B� �   DC� C�D � 1HCD � � � B( ) v� � I1H� 1IBG�H E1D� �

� L1CFN � DB� WCB   � �� � 6v � � ED3� i � FGE� � � E� � . � BE� ; � � h� m� EDE� CB� f� D� df� � EB   � IIf� � � f� � � � wd�

� IBG�H E1D� � CN � � H� W� G� � B�T � D� �B � B   EG�� C� � m� � s� � � BW� � D� B   � � ( . B   � � D� � ( 2B   � GC� Hw� �D � �

I1N � �G � IE� �B CEDH� m � s� � � BW� � D� B   � � ( HB� �D � � . D� � I1N � �G � 3� GB� � G�� wd� � B� � ��   � IE� �B CEDf�

BG� DH3� GH� � m� � s� F� GF� D� C� 1I�G � BE� B   � � 3� GB� � G�I � � EI1NDf� � E3� GCD � �I I� E� � B   � � � � � GEN � B   � �

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Capítulo  8  

DISCUSIÓN  

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D� � D9� � � � 19� � Ieo91� r� � � � � e� � I1� 9r   �� IL1� � 91� �1 u� rI9rI� �� � �� � e� �   9   � 1u9� I   o9eI� INIu�� N��

� a� Nl � � IL1� � 1� uI� � � n� e� N� � � IL1� �� � � l �� �� � � � � 1I   �N � e� o�r �� r� o9eI� IL1í� D� � I� 1� 9e� �

1� � � e�r I�� N� 9� u� 1� IL1� � � I1� 9r   �� IL1� �� o�r uIr� � � � el e� � �   INI�r � e � Ir� � u9eí� N9e� � l �N � e� e� �

� � e� �r u�1 � � 9   9� � �1 � I� �u 9e� o9u� 1� I�N � e� �� e� N� � � IL1f� � e� o9r� � NN9� s l � � N� � l uINIi � � IL1� � � � N��

u� � 1I� � � � � � l Nur�e 91I� 9e� �� uI�   o9� r� �N � r� or� e� 1u�� l 1� �a �1 � � � I   o9ru�1 u� � o�r �� N�� I1� l eurI��

9aI1�í � o� r   IuI� 1� 9� N� � � a�Nl �� IL1� or� aI� � � N� e�� rI� I� I9� � � � u9� 9e� N9e� �1 I   �N � e� n�   � bI   Ii �1 � 9�

N� � NIeu� � � � � � �1 � I� �u 9e� �� e� N� � � IL1z� � 1� � 91e� � l � 1� I�í � u�   � I� 1� N�� I1u� 1eI� �� � � � � e� N� � � IL1� n�

o9r� N9� u�1 u9� N�� r� eol � eu�� �� N�� e� N� � � IL1� e� � a� r�1 � 9ouI   Ii �� �e f� � �

� eul � I9e� or� aI9e� D� 1� r� o9ru�� 9� D� r� � �� INI� �� � e� s l � � a�1 � � � � yíéM� �� yí( é� o�r ��

or9� l 1� I� �� � � � � N� � r� e� � el � � l u� 1� �� , � 9 �r uní� éj j qz� � � r1�1 � � e� � u� �Nfí� xyyáz� � �b �� � u� �Nfí�

xyy) ;í� � � � yíé( � n� yíxÉ� o� r�� or9� l 1� I� �� � � � �   t e� l N9� � � � � � � � � � � � � � � � , l e�I 1� � u� �Nfí� xyy) z�

� �b �� � u� �Nfí� xyy) ;í� � � � yíMy� o�r �� �   oNIul � � � � �   t e� l N9� � � � � � � � � � � � � � � � , � � r1�1 � � i � � u� �Nfí�

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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  � � I�1 u� � l Nur�e 91I� 9ef� � eu9� o� r   IuIrS� � or� � � � Ir� � N� r� 1� I   I� 1u9� � � � N�� � �1 � N� �1 u� e� � � N�

e� � rI� I� I9í� � N� o9r� � 1u�U � � � � � � �r 1� � � 9   � r� I�NIi �� N� � , � INe91í� éj j xí� éj j Éz� 9l Du91� �1 � �

� l rNI1 1u91í� éj j xz� � INe91� � u� � Nfí� éj j Mz� � rrI1 � � u� �Nfí� éj j á� z� �   NI1� � u� � Nfí� éj j qz� � D� o�r � �

� 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�

� 1� l 1�� o9� N�� IL1�   �n 9r� � � � �1 I   �N � e� � � � oI� � � � � � rS�� n� � � � r� � �4 9e� � 9   � r� I�N � e� , � INNI�   eí�

xyyé;f�

� l 1s l � � � N� r� Ieur9� � � � N9e� � �u 9e� o�r �� � N� � 91ur9N� � � � r� 1� I   I� 1u9e� r� or� e� 1u�� l 1��

u�r � ��   �e � o� r�� N�e � � boN9u�� I91� e� 9aI1�e í� uI� 1� � l 1�� r�1 � I   o9ru�1 � I�� o�r �� N9e� �1 �� � r9e�

n�� s l � � or9o9r� I91�� l 1� r� Ieur9� or� � Ie9� � � � u9� 9� N9� s l � � el � � � � � � 1� � N� r� � �4 9í� o� r   IuI� 1� 9�

u9   � r� N�e �   � � I� �e � n� � � � I91� e� �� � � l �� �e � � 1� � �e 9� 1� � � e�r I9f� � � �   �e � or9o9r� I91��

I1� 9r   �� IL1� u�1 u9� or9� l � uIa�� � 9   9� � 1� �NL I� �í � N�� � l �N � l 1�� a� i � � 1�NIi �� �� n� � � ol r�� � � 19e�

o� r   Iu� � or� � � � Ir� � N� a�N9r� � 1� uI� 9� � � � N9e� �1 I   �N � e� o�r �� N9e� � �r �� u� r� e� � � � I1u� r� eí� eI�   or� �

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�

N� e� � � � IeI91� e� r� � � r� 1u� e� �� N�� e� N� � � IL1� � � � 1l � a9e� I1� IaI� l 9e� � � � r� o9eI� IL1í� �e S� � 9   9�

e� N� � � I91�r � �� N9e�   � U9r� e� � �1 � I� �u 9e� o�r �� � 1ur�r � �N � � � 1ur9� � � � u� eu�U � � � � �   9rl � � 9e� � � � N��

r�i �� 9aI1�� � Io9NN� e�í � N9e� � l �N � e� �� ul �N   � 1u� � e� � e� N� � � I91�1 � u9   �1 � 9� � 1� � l � 1u�� N��

or9NI� I� I� �� � n� N��   9r� 9N9 S�í � � l � rI� 1� 9� � 91� N� e� � � r�� u� rSeuI� �e � � � N� � eu�1 � �r � r�� I�Nf�

� N� � es l �   �� � � � e� N� � � IL1� � � � N�� r�i �� � Io9NN� e�� D�� orI9rIi �� 9� � � e� � � � N� orI   � r�

  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 �

  9r� 9NL I� �e � � � � I1I� �e � � 1� � N� � eu�1 � �r � r�� I�Ní� eI� 1� 9� eI�   or� � � N� 9� U� uIa9� � I1�N � � � N� � es l �   � �

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� é( y � �� �

� � � e� N� � � IL1� I1� r�   � 1u�r � N�� or9� l � � IL1� � � � � �r 1� f� � 9e� r� el Nu�� 9e� 9� u� 1I� 9e� � 1� � N� or� e� 1u� �

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�

� � r� � u� r� e� � � � N� � � �1 �N � n� � � � N�� � �r 1� ;í� o9r� N9� � l �N � e� � ol � � � � � eu�� N� � � r� l 1�   9� � N9� � � �

� a� Nl � � IL1� � 1� uI� � � I1I� I�Ní� I1� 9ro9r�1 � 9� � eu9e� � �r �� u� r� e� or9� l � uIa9e� � 9   9� l 1� 9� U� uIa9�

� � � e� N� � � IL1�   �e � � 1� � N� or9 r�   �� � � �   � U9r�   I� 1u9� � 1� uI� 9� � � � N�� r�i �� 9aI1�� � Io9NN� e�f �

� � � � �   � 1� I91�r � s l � � N�� or� e� 1u� � or9ol � eu�� r� s l � rIr�� � eul � I9e� � l ul r9eí� I1� Nl n� 1� 9�

� eul � I9e� � � � eI   l N�� IL1í� � l �1 � 9� e� � u� 1 �� � N�r 9� � N� o�1 9r�   �� n� eIul �� IL1� � 1� N�� s l � � e� �

� 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�

� � � � � e� r� � � 1ur�� 9� � 1� N�e � u� 1� � 1� I� e� � � N�   � r� �� 9í� s l � � e91� N� e� s l � � � I1�N   � 1u� �

� � 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�

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Capítulo  9  

CONCLUSIONES  

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� � � � � � � �

� 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�

� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �

nu � �U � � �S � L� � � �� � U� � � � � � LS�U � OI � � � � � U� L� � � � � �� �U � � �� � � � � NI � U�

� �6 � U� �� Ua� �O L   � � � �� � � � � US� � S�U � � I � � S� U� � � � � � � L� �� � C� � �� � � � � S�� � 6�

� � � � S� � �c � � d� � � � � S U� U� US� � �S � � U� � � � � �� � � �� � � � � � �� � �� L� � 6� S� O � � � � O�L S a�

6� � d� � � � � S � � � � � S� � � �� � S�   � � � � � � � S� U� � �� u�

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BIBLIOGRAFÍA  

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