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A genetic evaluation of some serum and milk production traits of dairy cattle Nash, Thomas Edward 1978

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A GENETIC EVALUATION  OF SOME SEBUM AND MILK PRODUCTION TRAITS OF DAISY CATTLE by THOMAS EDWARD NASH  B.Sc.  (Agr.),  University  o f B r i t i s h Columbia,  1576  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE GF MASTER OF SCIENCE in THE FACULTY  OF GRADUATE STUDIES  (Department o f Animal S c i e n c e )  8e a c c e p t t h i s t h e s i s a s c o n f o r m i n g to t h e required standard.  THE UNIVERSITY OF BRITISH COLUMBIA October, (c) Thomas  1978  Edward Nash, 1978  In p r e s e n t i n g t h i s  thesis  an advanced degree at the L i b r a r y s h a l l I  the requirements f o r  make i t  freely available  for  I agree  r e f e r e n c e and  f o r e x t e n s i v e copying o f  this  that  study. thesis  s c h o l a r l y purposes may be granted by the Head of my Department or  by h i s this  representatives. thesis  It  is understood that copying or p u b l i c a t i o n  f o r f i n a n c i a l gain s h a l l  written permission.  Department of  Animal S c i e n c e  The U n i v e r s i t y of B r i t i s h  2075 Wesbrook Place Vancouver, Canada V6T 1W5  Date  fi  fulfilment of  the U n i v e r s i t y of B r i t i s h Columbia,  f u r t h e r agree t h a t p e r m i s s i o n  for  of  in p a r t i a l  Columbia  not be allowed without my  ii AESTBACT  T h i s study  was i n i t i a t e d  certain  serum  cattle.  The p o p u l a t i o n  dairy  farms  c o n s t i t u e n t s and m i l k  in  the  under s t u d y upper  Serum s a m p l e s f r o m  analysis.  This represented  constituents  phosphate, glucose, albumin,  thyroxine  and  of d a i r y  on  commercial  region  cf British  were  groups with  used  i n the  approximately  significant  20  to  considered  between  herds  adjustments analyses lactation  s t u d i e d were lactation  having  a  under study.  important.  The  An  important  and s t a g e o f l a c t a t i o n  milk,  and  the  potentially Herd  a t time  appropriate  f o r serum  and  of  Seasonal seasons,  interaction  used  i n the  traits.  of bleeding  traits,  lactation  effects  and i n c l u d e d i n  covariables  t o t h e serum and p r o d u c t i o n  length  bicarbonate,  i n summer and w i n t e r  potentially  a l l traits.  were u n i q u e  start  sampling  creatinine,  for a l l traits.  and s e a s o n s was deemed  for  start  by  protein,  sampling. as  on t h e t r a i t s significant  as  covariables  lactation  be  estimated  o f serum  SGGT,  traits  inorganic  total  chloride,  first  were r e c o g n i z e d  influence  were e x p e c t e d  phosphatase,  f o r both  at time  calcium,  cholesterol,  Production  protein  i n progress  acid,  were  potassium,  amylase,  Several effects  effects,  alkaline  sodium,  f a t and m i l k  lactation  the  Valley  analysed  BOS, u r i c  bilirubin,  triglycerides,  were  located  animals  27 s i r e  traits  per group.  Serum  milk  545  the genetic aspects o f  production  was  Fraser  Columbia.  samples  to evaluate  were  while  Age a t were age a t  appropriate  i i i ccvariables All The  f o r the production  effects  traits  constants genetic  were e v a l u a t e d  under  with  study  traits  following  that  serum  squares  adjusted  by  data  her i n a b i l i t i e s  techniques.  the least  squares  subjected  amylase  (0,20),  o f 0.05 a n d 0.05, that  t ot h e  creatinine milk  Alkaline  correlated  highly  f o r both  phosphatase  production  traits  exhibited  reliable,  (0.32),  andt h e alkaline  {0.13),  deemed o f i n t e r e s t  and with  of  among  the traits  were  these  correlations  was  o f t h e magnitude o f t h e c o r r e l a t i o n and t h e  correlations  protein  significant  traits  potassium  existed  magnitude o f t h e a s s o c i a t e d genetic  a  respectively.  The e v a l u a t i o n  out i n light  have  creatinine  SGOT a n d EUN w e r e a l s o  evaluated.  The  to  t h e production  constituents:  Genetic correlations  relative  discerned  a l l  {0.30),  {0.08).  albumin  were  included  phosphatase  and  were  least  the resulting corrected  heritability  carried  by  analysis.  The  also  traits.  errors.  judged r e l i a b l e  were  as  fellows:  and negatively  with  both  milk f a t  first  correlated  f o r both  standard  and current  highly  lactations.  andn e g a t i v e l y  lactations.  Potassium  positive correlations  with  with  and  milk  a l l  albumin  production  alone. Genetic in  order  exhibited albumin, BON  was  correlations  t o elucidate a  positive  common  e x a m i n e d among s e r u m  underlying  correlation  anda negative negatively  were a l s o  with  c o r r e l a t i o n with  correlated  with  genotypes.  traits SGOT  amylase, p o t a s s i u * and alkaline  amylase  phosphatase.  and  positively  correlated with  with albumin.  alkaline  Creatinine  phosphatase,  and  negatively  w i t h amylase  positively  correlated  with albumin.  involving further, subset  some  serum  and  demonstrated  involvement i n other  traits.,  positively  alkaline  phosphatase  and  potassium.  tabulated the production  i t reduced the t o t a l which  correlated  amylase,  correlated  T h i s s t u d y e s t i m a t e d and  was  a  genetic  genetic  traits  number o f serum  Potassium  parameters  i n dairy  cattle,  constituents  component  was  or a  to  a  genetic  TABLE OF CONTENTS  Abstract List  *. v . . . . . . . . . . . . . . . . . . . « . . i i  of Tables  . . . . . . . . . . •.-.......»......... . . . . . v . . v i  A c k n o w l e d g e m e n t s ..., ............ •. .... •........ ...... . . v i i ;  Introduction Literature  ........................................v..  1 5  Review  H a t e r i a l s and Hethods  19  Lata C o l l e c t i o n and A n a l y s i s 19 S t a t i s t i c a l Models ......................,27 E s t i m a t i o n o f V a r i a n c e Components 29 V a r i a n c e o f V a r i a n c e Components . . . . 31 H e r i t a b i l i t y E s t i m a t e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Variance of H e r i t a b i l i t y Estimates ..................33 C o r r e l a t i o n E s t i m a t e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 V a r i a n c e o f C o r r e l a t i o n E s t i m a t e s . . . . « . . • • • . . . * . . . . . 34 35  R e s u l t s and D i s c u s s i o n  Conclusion Bibliography  Appendix  ........v.......••...••..............75  ................... ......  ......v.  79  .................. ...... • ...»••• ..»........•..• 8  L I S T OF TABLES  Table I  Analysis o f v a r i a n c e (ANOV) t a b l e and e x p e c t e d mean s q u a r e s (EMS) f o r t h e e x p e r i m e n t a l m o d e l s .  Table I I  Least sguares constants associated with summer and w i n t e r s e a s o n s and with t h e c o v a r i a b l e s .  Table I I I Least squares means and a s s o c i a t e d e r r o r s (SE) w i t h R for a l l correction 2  standard terms.  T a b l e IV  Variance components and h e r i t a b i l i t i e s a s s o c i a t e d SE f o r t h e i n d i v i d u a l t r a i t s .  Table  V  Phenotypic {below diagonal) (above d i a g o n a l ) c o r r e l a t i o n s a s s o c i a t e d SE. ,  Table  VI  Genetic genetic diagonal)  Table VII  with  and e n v i r o n m e n t a l among t r a i t s with  correlations {below diagonal) and components of covariance (above w i t h a s s o c i a t e d SE.  Genetic (above) and phenotypic (below) c o r r e l a t i o n s between s e l e c t e d serum c o n s t i t u e n t s and t h e p r o d u c t i o n t r a i t s . ,  Table VIII Genetic (above d i a g o n a l ) and p h e n o t y p i c diagonal) correlations among selected consitituents.  (below serum  vii ACKNOWLEDGEMENTS  The  author  acknowledge importance  the e f f o r t s  like of  i n the research  t h a n k s t o Dr. study  would  and  E . G .  to  those  Peterson  h i s discussion  people  who  f o r h i s help  during  a l l o f which proved  and  Duncan  Jeffries,  with  t h e computer  the  in  opportunity were  To  to  of  prime  thesis.  Many  initiating  research  invaluable.  and  Mabel  the  thesis Striker  many t h a n k s f o r t h e t e c h n i c a l a s s i s t a n c e  work r e g u i r e d i n t h i s  stimulating  discussion  manuscript.  Finally,  many t h a n k s  and  in  document.  this  and p r e p a r a t i o n o f t h i s  preparation,  encouragement  take  help  which  helped  the  to  study, in my  physical  the wife,  and  again,  for  writing of this Patricia,  for  preparation of t h i s  1 IMJOEUCTION  The  selection  of dairy c a t t l e  f o r the production o f s i l k ,  m i l k f a t and m i l k p r o t e i n h a s r e c e i v e d much a t t e n t i o n i n history. refined  Selection selection  conventional dramatic order  cattle.  progress  traits,  cost  the  researchers  t o such  a  are not l i k e l y  (Bobertson  interval,  thus  limiting  selection  procedures  production method the the of  traits  of  blood  for  replacement  These  response  c o u l d overcome t h e s e  a r e among  intermediary the animal.  synthesis process.  that  and a r e a v a i l a b l e  overcoming these  areas  of  dairy  females  to  an  age  s e l e c t i c n and  factors  effectively  t e s t e d and i n c r e a s e t h e g e n e r a t i o n  selection  traits  In  i n c l u d e t h e s e x - l i m i t e d nature of  per  unit  are  restrictions.  t h e more p r o m i s i n g  in l i f e  If  i t would  terms.  correlated  early  time.  restrictions,  r e p r e s e n t an improvement i n g e n e t i c and e c o n o m i c Selection  1966).  r e s e a r c h e r s must e x p l o r e  males.  number o f a n i m a l s  that  t o produce t h e  a r e m e a s u r e a b l e , and t h e i n i t i a l  replacement  have  degree  present i n the breeding  the cost of r a i s i n g  of r a i s i n g  limit  continue,  restrictions  the t r a i t s  and  i n t h e past  the r e s t r i c t i o n s  These  intense  procedures  improvements a c h i e v e d  that  where  been  m o d e l s and t e c h n i q u e s  selection  which e x p l o i t  the  has  recent  with t h e m i l k r e p r e s e n t s one  T r a i t s measurable i n  i n this  repect.  Bleed i s  fluid  between t h e mammary s y s t e m and t h e r e s t  As s u c h ,  i t c o n t a i n s i n p u t compounds t o t h e m i l k  process  as w e l l as some compounds s u p e r f l u o u s  to the  I n a d d i t i o n , b l o o d c o n t a i n s o t h e r compounds w h i c h  are  2 indicative makeup.  o f g e n e r a l body c o n d i t i o n The  latter  relationships production blood types. the  between  traits.  cell  has  been  studied  various  reaction,  alleles  and  accounted  relationships  milk  for  fat  studied  transferrin,  the  and  milk  were  red  and enzyme  these r e l a t i o n s h i p s  existed  genetic  examining traits  systems  serum  P i r c h n e r (1969) r e v i e w e d  strongest  by  polymorphic  Some p o l y m o r p h i c  antibody  or o f the animal's  and  noted  between c e r t a i n b l o o d  percentage,  yet  these  type  relationships  l e s s than t e n percent of the genetic  variance of  milk f a t percentage. Other  polymorphic  correlated there  polymorphic  other systems.  of  a  number  homozygosity  gene  response  effect,  reproductive  performance.  is  other  and  of f i n i t e  a  failing,  associated  with  that  a t one o r a few  selection  would  are loci.  cause  a  few g e n e r a t i o n s a t t h e l o c i  homozygosity  may  There  will  system  for are  in  result  a  example,  in  no  decrease  in  fitness  and  discrepancies  interpretation  classification variable  result  traits,  biological  continuously  polymorphic  of alleles  in  homozygosity in  system  this  highly  t o s e l e c t i o n . •• A l s o , due t o t h e p l e i o t r o p y o f  performance  statistical  to  difficulties  t o be a d d i t i v e ,  i n v o l v e d . . The a t t a i n m e n t o f further  p r o v e n t o be  By d e f i n i t i o n , t h e y a r e s y s t e m s  small  towards  not  In a d d i t i o n  intrinsic  A s s u m i n g gene a c t i o n trend  have  with milk production.  are  composed  systems  (polymorphism)  of  in  correlating  with a t r a i t  (the milk production t r a i t s ) . .  d o e s have  an  input  into  the  the  a  that  I f the  continuously  3 variable  trait,  which give factors  the  must  trait  could  systematic  i t  be  small  i t s continuous systematic  factors  offer  present  study  in  distribution  their  be  and  distribution.  area  that not  to  the  factors  These  other  or environmental i n nature.  an  used t r a i t s  in relation  for  farther  were c o n s i d e r e d  based  The  study.  to  p r i m a r i l y on  be  This  continuous  a  polymorphic  are  continuously  system. There variable roughly those  are  and  many c o m p o u n d s i n t h e  relevant  grouped  c o m p o n e n t s , and which  milk  synthesis.  by  mediate  are  study in  to i f  approach.  their  animals. on of  a  For  i t  i s  not  the  Certain  be  material portion and  form a  when cost  material  constituents  to  of  may  reason,  profile  this  that  present  which  milk  the  into  milk  of  milk  which  appear  more  traits, to  study  provided  were added  must  an  several  field  cost  of  number  of  based l a r g e l y  efficient a  be  premising  yet  a large was  in  blood  results in a  were i n d i c a t i v e of  profile  milk,  synthesis,  measurement  production  this  part  be  pathways i n v o l v e d  selecting  to apply  of  synthesized  p r o h i b i t i v e when a p p l i e d  this  To  These compounds can  relevance  Ease and  with  that  biochemical  intended  constituents  body f u n c t i o n s .  do  considerations  measure.  a  mammary c e l l s  some o f  human b i o m e d i c a l blood  forming  which  relationship  m e a s u r e m e n t may  synthesis.  constraints of  other  constituents considered  the  those  Besides the there  milk  i n t o those  extracted  but  to  blood  survey  wide range  of  constituents  4 which  were  judged  to  production  or  satisfactory  analytical  If  one  theory,  certain  those  and  traits  traits  genetic  t o be  is  used  g e n e t i c and  present  that  production t r a i t s measured c a n  be  milk  which  had  conventional  parameters  must  or  concurrent  phenotypic  be  reliably  I f one  i f one  selection  estimated  d e s i r e s to  desires  with  known.  to  relate  changes i n  correlations  use  between  blood these  estimated. study, be  a more c o m p l e t e  proposed  in  individually.  traits  and  to  available.  trait  simultaneously,  parameters w i l l  present  a  relevance  g e n e t i c v a r i a n c e s must be  must a l s o be  In t h i s  to  techniques  change i n p r o d u c t i o n  constituents,  particular performance  genetic  a number o f t r a i t s the  of  reproductive  i s to u t i l i z e  Heritabilities for  be  some and  estimates estimated and  ascertained.  and  evaluated  mentioned  i n an  attempt  comprehensive i n v e s t i g a t i o n .  important  some  of a l l the above  subset  relationships of  the  serum  between  It the  constituents  5 LITESATOBE BEVIBB  In a p a r t i c u l a r few  major e f f e c t s .  order  study  The e x p e r i m e n t  to facilitate  the analysis  there are other factors importance, , accurately  that  tested  These secondary  of these e f f e c t s .  effects  must a l s o  t h e main i n t e r e s t  and c o r r e l a t i o n s  statistically,,  Beyond  identifiable  factors affect  quantified  and  analysis.  It  significant  effects  optimal as  the  they  to  h a s been c i t e d  possible,  in  the  range  as a f a c t o r  by many r e s e a r c h e r s .  processes that  present  in  this  to  157 months, w i t h  actual  prime  analysis.  derived  effect*  many  I f they a  and  effect  place remove  readily can  be  i n the as  order t o get a  and  many  complete  constituents.  levels  o f serum  Age may b e due t o a number o f matures.  Within  s t u d y , a g e may n o t be a s i m p o r t a n t  A l l animals  and t h e r e f o r e the age r a n g e The  of  the  be  i n governing  were a more homogeneous  under random c o n d i t i o n s .  represented. ,  could  a p p e a r o r change a s t h e a n i m a l  since the animals  lactating  in  deserve  understanding o f the nature o f the blood  constituents  In general,  was i n a s i r e  sire  identify  out i n  be q u a n t i f i e d a s  blood composition.  interpreted,  i s  i n one o r a  i s s e t up and c a r r i e d  a s p o s s i b l e and t a k e n i n t o a c c o u n t  heritabilities  Age  interested  which a r e s i g n i f i c a n t b u t n o t  In t h i s p a r t i c u l a r study, so  one i s u s u a l l y  age r a n g e  group  than  sampled i n t h i s of  0-23  i n this  months  study  a c o n c e n t r a t i o n i n t h e lower  expected study  were  was  not  was 23 months  portion of this  6 range.  There  continued  still  growth,  was  the  present,  maturing  however,  processes  the and  effects age  of  related  stresses. Serum c a l c i u m l e v e l s have been r e p o r t e d t o be s i g n i f i c a n t l y affected  by  significant  linear  significant and on  age.,  age  Kitchenham  Tumbleson  decrease  effect  and  Tumbleson  Rowlands  Sodium, p o t a s s i u m by  age,  a l . 1S73b  nitrogen  et  (BON)  point The  a  significance reduced this  than  study  studied our  appeared  lower  study.  decrease  significant  limit  were  by  t o two  Furthermore,  reported  years  was  t o 12.75  account  Since of  and  age  Rowlands  i s ,  Kitchenham (1976)  This  age, range  most o f  the  range,  the  f o r the s i g n i f i c a n c e  Kitchenham  urea  urea with  years.  be  (Tumbleson  Rowlands  i n s o d i u m and  24 months  to  stabilizing  and  and  range.  and  that  but  decrease 1.25  age.  Blood  e t a l , 1973b  experimental  may  with  curvilinear  pattern,  Kitchenham  i n the 0 -  based  relationship.  not  similar up  (1975)  Kitchenhas  e t aJL. 197 5).  a  No  was  i n the present study  (Tumbleson  significant  although the range wider  a  chlorine  by age  age.  a  herds.  reported  Kitchenham  affected  that  indicated  with  This conclusion  i t a significant linear and  reported  Kitchenham e t a l .  been n o t e d t o  followed  a l . 1975).  was  has  over the range  and  significantly after  (1S76).  level  w h i l e K i t c h e n h a m e t a l . (1975) and  (1976) termed  affected  calcium  a number o f  phosphate  a l . {1973b)  r e p o r t e d by  Rowlands  e t a l . (1973b)  relationship,  et  was  values o b t a i n e d over Inorganic  in  et  noted i n  (1976)  noted  7 that the  the herd  decrease under  Serum  o f s o d i u m w i t h age  showed  proteins  have  been  noted  increase  relationship of  significant  albumin  in  it  may  by  age,  be  expected  situation  by  y e a r s o f age three  and  years  age  selective culling Bilirubin significantly concurred  with t h i s  Thyroxine the  normal  fieineke  four  years of  present,  age.  albumin  noted  an  age  no  but  study  were  e t a l . (1S75) i n d i c a t e d  the  with  age  present  but  portion  of t o t a l  (1971)  speculated  This have  an the  Total (r=0.71).  serum l i p i d s  further  to  again  study.  with c h o l e s t e r o l  thereafter,,  and  affected  clarified up t o  decrease  the three after  been c a u s e d  by  animals.  noted  by T u m b l e s o n and  with  age,  Mylrea  Hutcheson and  (1971) t o  Healy  41968)  statement.  secretion rate  productive  1959).  age.  cholesterol levels increased  was  increase  in  Hutcheson  of c i d e r  was  by  increased  t r i g l y c e r i d e s l e v e l s are a l s o  decreased  of  protein  (1974)  the  correlated  that  stating  in  Arave  used  a large  T u m b l e s o n and  affected  Little  concentration  that  make up  be  decreased with  through  measured  positively  Triglycerides  to  thereafter.  cholesterol  were  two  to total  age./  triglycerides.  d i f f e r e d frcm  lipids  frcm  compounds  c h o l e s t e r o l and increase  with  relationship  Lipid  range  that  while a l k a l i n e phosphatase no  have been s p e c i f i c  study.  Tumbleson e t a l . (1973a) s t a t e d w i t h age  may  age  declined range  with  increasing  i n dairy goats  age  (Flamboe  over and  8 Kitchenham with  age i n a s t u d y  Rowlands  (1976),  relationship situation was  e t a l . (1975) n o t e d  in  between  was  0.5  a  a number o f h e r d s .  within  glucose  and age.  to  5  study  w h i c h showed  years  study,  Further  reported  obscuring  data, although  the lower  most  age l i m i t  P e t e r s o n and H a l d e r n  was  relationship  the  present  a g a i n much  age i n t h e l e v e l o f i n o r g a n i c p h o s p h o r u s , BON, phosphatase,  transaminase  (SGOT)  total  protein  Kitchenham  and  serum albumin,  and g l u c o s e . e t a l . (1975)  and  decreases  oxaloacetic  and s i g n i f i c a n t i n close  Tumbleson  study»s  cholesterol,  glutamic  T h e s e were  The  lower.  (1978) r e p o r t e d s i g n i f i c a n t  alkaline  This  no change t h e r e a f t e r . of  no this  (1971).  a significant  o f age, w i t h  range o f s i g n i f i c a n c e encompasses  with  herd  glucose  K i t c h e n h a m and  a r e p o r t by Tumbleson and H u t c h e s o n  a w i t h i n herd  from  involving  an i n c r e a s e i n serum  increases i n  agreement  etal.  with  (1973a)  and  p o t e n t i a l l y important  for  (1973b).,  Seasonal  e f f e c t s were c o n s i d e r e d  the  population studied.  and  winter,  and t h e d i f f e r e n c e  measure  of  included  differences  the  major c a u s a l  been  seasonal  s a m p l e s were t a k e n i n sampling  effects.  i n climate, diet  i n both  p e r i o d was  These  effects  summer  taken  as a  may  have  and f e e d i n g management  as  factors,  Relatively have  Blood  few  shown  significantly  lower  of t h e c o n s t i t u e n t s p r e s e n t to  vary in  seasonally.  summer  than  in  blood  Sodium was  winter,  while  noted urea  serum t o be and  9 albumin et  concentrations  a l . 1974).  Boss  were  and  f o r albumin,  significantly  higher  in  et  to  higher  increased  protein  flanston  1972,  This  and  effect  Halliday  serum c h o l e s t e r o l ,  l e v e l s i n summer.  Information phosphatase bleeding, seasonal result  on  was Boussel  difference  was tempered  aniffials  (0-2  years)  study,  and  Stallcup  (1966)  that,  Two  by  some s t u d i e s  ( L i t t l e and  this  in  this  by  the  when h e r d s  to  have a  summer l e v e l s o f  general  provided  conflict  lowered  with  cholesterol  strong b a s i s  f o r the  effect.  effects In  f o r SGOT  and  (1966)  enzymes.  noted  a  significant  The i m p o r t a n c e o f t h i s  t h a t t h e range o f study  age  and B e i n e k e  seasonal e f f e c t s  f o r the  was n o t t h e same a s i n t h e  statement  from s e a s o n a l  alkaline  a summer^winter s y s t e m o f  made  by  Boussel  age d i f f e r e n c e s i n t h e i r s t u d y  (Flamboe  indicated significant  e t a l . 1971)  higher  generally  Stallcup both  be due t o  the year.  was i n  noted  for  be l e s s e n e d  noted  limited.  separated  papers  shown  by t h e f a c t  present  be c o m p l e t e l y  Increased  Prewitt  may  (Payne  summer.  should  This  found  the  seasonal  and  was  in  seasonal  also  Drea  was  grass  Neither study  argument o f s i g n i f i c a n t  this  summer.  (1976)  who  disputed  in  throughout  although  e t a l . (1975)  summer.  e t a l . 1S75,  f e e d i n g program  Boss  Arave  the  been  Manston  BUN.  consistent  has  s'usier (Payne  and f u r t h e r s t a t e d t h a t c a l c i u m  protein intake i n  dietary  affect  be  in  H a l l i d a y (1976)  relationship  a l , 1974)  higher  and  could not  effects. 1959, H i x n e r  e t a l , 1962)  with  to thyroxine  regards  10 l e v e l i n the  Herd portion  blood. ,  effects of  the  constituents Hanston herd  feeding,  have  affected  by  Metabolic  et  the  largest  with  with  the  single  many  a l . 1974,  blood  Rowlands  observation  c o m p r i s e d o f a number o f  management and  a l l blood  and that  smaller e f f e c t s ,  microenvironment  a l l  albumin,  (CMP)  were  associated  and  regarding  Specifically,  they  Dinkel the the  uric  literature contributors  not  to  the  it  in  The  study  were  potassium the  herd  same  effect.  herd  on  the  phosphatase.  i n the  expected in  herd.  constituents  at  of  Ccmpton by  sodium,  of  studies  the  total  chlorine, bicarbonate,  variation  present  significantly  arrived  acid, cholesterol,  was  be  present  significance  well reported  reviewed,  (1968)  the  affected  calcium,  alkaline  SGOT, t r i g l y c e r i d e s ,  a m y l a s e was  the  importance  of c r e a t i n i n e and  h e r d e f f e c t on  and  phosphate,  noted  involved  these r e s u l t s .  studies  inorganic  to  in  (1974) i n two  significantly  with  Ccmpton  Wilson  conclusion  constituents  concurred  BON,  bilirubin,  literature  that  the  variability  i n the  involved  Payne e t a l . (1973) and  (1974)  glucose,  constituents  herd.  Profile  common t o  The  Payne  consistent  been r e p o r t e d  indicated  Hewett  are  for the  farm.  Virtually study  was  account  associated  1974,  This  differences  w i t h the  and  variablity  (Hewett  1976}.  notably  and  were n o t e d t o  literature. h e r d s w o u l d be  most  of  the  protein, thyroxine From  the  important parameters  11 measured. due  to  Moreover,  some  interactions  statistical  between  variation.  Payne  et al.  lactational  group  interaction  lactational  Since  group  the  associated  with  (1974)  stage  of  different  i n blood c o n s t i t u e n t  be  included  the  0-120 d a y s ,  with  at  the  of  h e r d by  comparison  time  stresses  into  account  levels. ,  measure  situation  to  little  effect  of  sampling i s  and  nutritional  as  Many  of  were i n o r g a n i c  source of i n the  stage i n t h e i r  Bowlands e t aj..  significant  beyond  a  studies  of lactation  due t o s t a g e o f l a c t a t i o n  t h i s general s i t u a t i o n  et  taken  some  general levels  of  in  arise  sources  that  large  I n one o f t h e Compton s t u d i e s ,  constituent  both  was  production  variation  noted  other  indicated  lactation  i t should  analysis.  and  may  effect.  regimes,  literature  herds  complications  variability  in  the  t h i s range. phosphate  {1975)  range  in of  Exceptions to and  potassium,  o f which showed no s i g n i f i c a n t t r e n d w i t h r e g a r d s t o s t a g e lactation.  al.  (1975)  More  indicated  protein  and c a l c i u m  showed  a  lactation with  looked using  progressed.  with  at  results  following:  of  glucosej  at the start  of  i n the early  BON,  total and  Sodium d e c r e a s e d as  stages.  relationship../  lactation  lowlands  lactation  A l b u m i n showed t h e s t r o n g e s t  positive of  the  increase thereafter.  especially  this  effect  the  were l o w e s t  significant  lactation,  concurred  specifically,  monthly g r o u p s a s s t a g e s o f l a c t a t i o n . ,  Little  Hewett  on an e x t e n d e d  relationship  (1974)  blood  He n o t e d  {1974) also  profile, inorganic  12 p h o s p h a t e and p o t a s s i u m  were  not a f f e c t e d  sodium, t o t a l  and  BUN  protein  were  by l a c t a t i o n , and  significantly  that  affected.  T h e s e r e s u l t s were i n a g r e e m e n t  «ith Rowlands e t a l . (1975).  the  reported  other  hand,  Hewett  (1974)  a l b u m i n were n o t s i g n i f i c a n t l y be p o i n t e d  out t h a t Hewett  range o f l a c t a t i o n s t a g e Hewett effect  (1974) a l s o  namely  inorganic  iodine.  effect  on  thyroxine  turnover  The  effect  Body  body f l u i d s  It  considering  the  noted  that  genetic  a much  there  was  no  total  e t a l . (1962)  noted  secretion  as  measured  iodine a  by  variability has r e c e i v e d Studies  and  significant  the  chemical  method.  an a n i m a l ' s g e n o t y p e on t h e v a r i a b i l i t y  level  i s of prime importance and  biological of  in  the  nutrients  accepted  that  either  many o f t h e s e  directly,  was  i n blood  associated constituent  some a t t e n t i o n involving  the  CHP  functions. under  as i n p r o t e i n s y n t h e s i s ,  l e v e l was  i n the  in  absorbed  processes are  with a p o r t i o n  of  present  compounds p r e s e n t  t h r o u g h c o n t r o l o f enzyme p r o d u c t i o n .  control  larger  measures o f t h r y o i d  iodine,  are the r e s u l t of synthesis  control,  should  significant  g u t , and o r a r i s e a s b y - p r o d u c t s o f b o d i l y  indirectly genetic  bound  metabolites  i s generally  It  and  i n h i s studies.  Mixner  of  constituent  study.  from  (1974) was  protein  thyroid  calciui  a f f e c t e d by l a c t a t i o n .  o f s t a g e o f l a c t a t i o n on any o f t h r e e  activity,  blood  glucose,  On  Whether  or  this  of the observed  under  question  and  literature. reported  a  significant  13 heritabilxty  f o r many o f t h e c o n s t i t u e n t s t e s t e d .  correlations,  Kithchenham  significant  heritabilities  (0.77±0.38) , a l b u m i n calcium and  and  inorganic  The  were  standard  design  resulted  group  size  study  by  However, protein.  estimates  analysis  on b e e f  differences  for  s t u d y by a  of  the  heritabilities  potassium  (0.40±0.23),  (0. 10±0.13) and  i n o r g a n i c phosphate  heritabilities  were  rough  with the  made. the  agreement One  range  i s that  o f age  used  these  estimated former  the second in this  and  thus  the  same  reduced  standard  errors.  only  for  e t a l . (1974)  constituents  two were  estimated  halfsib  eliminating  for  glucose  (0.19±0.17),  albumin  (0.18±0.16).  f o r BON  estimated  using  After  total  regions i n v o l v e d i n the  calcium  and  study, but study  in  progeny  showed  significant  weeks o f a<?e,  with  errors  (1976)  reliable  Bowlands  number  associated  with  small  regressions  more were  that  standard  Rowlands  c a l v e s o f 9-11  study, s i g n i f i c a n t (0.18±0.16),  high  Dam-daughter  with  heritabilities  insignificant  associated  (1959) s t a t e d  in  estimates  An e a r l i e r  Sodium, g l u c o s e  approximately  and  these  (0.50*0.34),  with  mainly  BUN  i n s m a l l progeny groups of  Kitchenham  heritability  for  protein  (0.53±0.34).  due  reported  experimental  result  unreliable estimates.  total  halfsib  to the  Bobertson  will  error)  errors  were q u i t e l a r g e ,  f o u r animals each.  (tstandard  associated  heritabilities which  (1976)  potassium  phosphorus  heritabilities.  fiowlands  (0.47±0.33) ,  (0.46±0,33) and  Osing  Nensignificant  sodium. two  dealt  present study.  T h i s was  points should  in be  with animals o u t s i d e The  other point i s  14 that to  the study  minimize  involving  the beef  the standard  errors  c a l v e s was o f s u c h  a design as  c f e s t i m a t e and t h u s  were  more  reliable. Wilson which  and D i n k e l  were  largely  They n o t e d  and  glucose  the  (0.19),  and  BON (0.46)*  SGOT  daughter  groups,  Furtmayr  (1975)  present  Bettini  (0,45),  The  (0.10),  first  uric  bilirubin  They  acid  (0.12)*  This  study  estimated  cholesterol  alkaline  phosphatase  used  (1966) c o n c u r r e d  213 cows i n e i g h t  reliable  blood  If this  constituent  effect  postulate variability  a  for  heritabilities for  and SGOT, serum  cholesterol  Stufflebeam with  level  levels  base  widely  a  and L a s l e y  (1969) and T a y l o r  high h e r i t a b i l i t y . genetic  by m e a s u r i n g t h e e f f e c t then  and t h u s  i n blood c o n s t i t u e n t  is  h =0,50 f o r d a i r y c a t t l e i n  this relatively  i s significant, genetic  esimates.  heritabilities  insignificant  A r a v e e t a l , (1975) n o t e d lactation.  (0.21),  (0.47),  significant  but r e p o r t e d  significant  phosphate  Some s t u d i e s have a t t a c k e d t h e p r o b l e m o f a of  i n t h e CMP,  e t a l . (1975)  listed  a d e s i g n which s h o u l d y i e l d  genetic basis of  reported.  study.  (0.22), i n o r g a n i c  (0.19).  bilirubin, cholesterol  S i Jl«  phosphatase,  present  f o r calcium  (0.40), albumin  their  t o those  h e r i t a b i l i t i e s f o r i n o r g a n i c phosphate,  alkaline  in  heritabilities  glucose  addition  constituents  h e r i t a b i l i t i e s f o r a l a r g e number o f c o n s t i t u e n t s under  examination  (0.12)  in  insignificant  creatinine reported  (1S68) r e p o r t e d on a s e t o f  i t may  be  a heritable  levels.  Heyns  basis  of breed.  reasonable  to  p o r t i o n of the (1971a)  noted  15 significant  breed  potassium,  differences  calcium  differences  and  occurred  phosphate.  Runkel  effects  for  a group  involved part The  to  results  repeatability  constituents  although they noted  further  this  i s an  to  high  (0.194),  creatinine  study  upper  the  limit  of  animals  majority  o f cows i n t h e  same as t h e  total  in lactating, present study  in  selection  of  (0.205)  and  performance.  The  correlations  among  Kitchenham age  and  and  breed,  Compton the  Bowlands noted  stock  and  albumin  condition.  c o u l d be  The  they  between  of great value  prediction  researchers  constituents (1976),  alkaline  were i n t h i s c o n d i t o n .  production t r a i t s  breeding  as  phosphate  C o r r e l a t i o n s among l e v e l s o f b l o o d c o n s t i t u e n t s and b l o o d c o n s t i t u e n t s and  in  They r e p o r t e d  (0.613),  nonpregnant  ones  reliable.  inorganic  protein  up  insofar  of h e r i t a b i l i t y . for  set  were  interest  SGOT ( 0 . 2 6 4 ) , g l u c o s e  for  repeatablilities  the r e s u l t s  were  (0.514),  (0.246)  environmental  S i n c e t h a t s t u d y was  repeatabilities  CO. 5 4 6 ) ,  indicating  basically  repeatabilities,  moderate  phosphatase  breed  (1S78) have e s t i m a t e d  present study.  of  inorganic  effects,  Waldern  estimate  and  breed  significant  w i t h i n b r e e d i s low,  o f serum  i n the  sodium,  noted  phosphatase  phosphatase,  Nonsignificant  albumin,  e t a l . (1953)  overshadow b r e e d  P e t e r s o n and  alkaline  glucose.  for  differences for alkaline that repeatability  for  have  of  future  noted  have  some  studied.  using c o r r e l a t i o n s adjusted f o r  a significant  positive correlation  between  16 calcium  and a l b u m i n  correlation was  (r=0.33).  Payne e t a l . ( 1 9 7 4 ) , u s i n g  c o e f f i c i e n t s , supported  apparent  only  positive correlation  potassium,  with  season.  reported with  inorganic  earlier  phosphate  their  importance  used  in  the  w h i c h was  (Payne  taken  significant w i t h sodium inorganic  into  be  sodium  that  account.,  (0.34), g l u c o s e with  with  large  and  (0.39),  (0.42),  means  for  with  (0.31)  listed  (0.40),  (0.43),  with sodium  phosphate  these  e t a l . 1975), was  albumin  albumin  were  variability,  Rowlands e t a l . (1974)  calcium  (BUS  although  erratic  Kitchenham  (0.34), albumin  inorganic  herd  Thus t h e w i t h i n herd  e t a l . 1973,  with i n o r g a n i c phosphate  albumin  (0.25 t o 0 . 6 0 ) ) ,  by t h e f a c t  both  a  e t a l . (1973)  coefficients  with  c o r r e l a t i o n s o f glucose with  phosphate  Payne  and  potassium  calculations. to  study,  (0.49)  with  noted  and r=0.24 i n t h e  partial correlation  was l i m i t e d  judged  correlations not  In  phosphate  i t  between i n o r g a n i c p h o s p h a t e a n d  i n t h e w i n t e r season  an  although  They f u r t h e r  r=0.40  some s i g n i f i c a n t  inorganic  finding,  i n t h e summer p e r i o d .  significant  summer  this  simple  glucose  glucose  with  (0.30),  albumin  calcium  (G.60),  and  sodium  with  c a l c i u m (0.31) . Heyns  (1971b)  reported  correlations  c o n s t i t u e n t s i n v o l v e d i n the present study. were  in  group. BUN  addition A  (r=-0.180)  mentioned.  to those  negative was Heyns  also  Several  several of  these  c o n s t i t u e n t s s t u d i e d by t h e Ccmpton  correlation in  among  agreement presented  between with  a  albumin  and  study  previously  significant  positive  17 correlations  of  creatinine  phosphorus  (0.397),  and  phosphorus  with  phosphorus glucose  (0.179).  w i t h BUN  With r e g a r d s levels  glucose  of  with with  (0.192), Negative  with  appeared to  and  to  the  study of  correlations  of  this  aspect.  the  scope  of  be d e s i g n e d  constraints. of t h i s  were  environmental  reported  correlations  Si  the c o l l e c t i o n  in  Furthermore,  in  away  "disappointing  sodium. of herd  milk  frcm  the  guality  potassium  design  correlation  average  in  into  a  w i t h a minimum should  both  allow  g e n e t i c and  o r with  is  and  greater  population  thus  literature.  than mean,  twe  Payne  standard  reported  that  was a s s o c i a t e d w i t h  high  lew l e v e l s o f a l b u m i n , the relationship  was much t h e s t r o n g e s t . basis  the  production  t h a t an " a b n o r m a l " l e v e l o f a  and y i e l d "  Of t h e s e i n d i c a t i o n s ,  albumin  order  T o be most w o r t h w h i l e ,  the  using the c r i t e r i o n  serum c o n s t i t u e n t i s one which  of  to  components.  a l . (1973),  levels  between  correlations  of data  to estimate c o r r e l a t i o n s  have r e c e i v e d seme a t t e n t i o n  deviations  for  t h a t most o f t h e  C o r r e l a t i o n s between b l o o d c o n s t i t u e n t s and m i l k traits  with  (-0.345).  interest  t h e study.  phenotypic  (0.290)  were n o t s e t up p r i m a r i l y  Instead, after  with  phosphatase  b l o o d c o n s t i t u e n t s , i t was a p p a r e n t  should  subdivision  alkaline  estimation  albumin  phosphatase  (-0.209) and w i t h c r e a t i n i n e  t o be i n i t i a t e d  expand  (0.212),  alkaline  s t u d i e s reported i n the l i t e r a t u r e deal  BUN  with  calcium or lew  These were d e t e r m i n e d  disregarded  individual  levels on a cow  18 differences. e t a l . , (1915)  Kitchenham regression  coefficients  (b«=-0.082),  glucose  indicating et  that  and  degree  of  at several  Furtmayr  (1975) , i n h i s I n a u g u r a l  involving  glucose  SGOT  (b» = 0. CC95),  relationship  but  and  exists.  were u n a b l e  and t h e milk  Sink  t o r e p o r t any milk  168 H o l s t e i n - F r i e s i a n s .  (0,33). milk  A  negative (-0.20),  yield  positively  correlated  correlated  with  Bondarenko  e t a l . (1976)  c o r r e l a t i o n s between  with  glucose  milk  and a l b u m i n .  of correlations  production  Dissertation,  was p o s i t i v e l y c o r r e l a t e d  and  cholesterol  albumin  contained extensive l i s t s  some serum c o n s t i t u e n t s  with  yield  between serum c h o l e s t e r o l and any  between  milk y i e l d  milk  trait.  Two o t h e r s t u d i e s  study  partial  i n t e r r e l a t i o n s h i p s between serum  traits,  significant correlations production  and  s  production  significant  between  BON ( b = 0 . 033)  some  a l . (1973) l o o k e d  lipids  reported  reported  His findings  with c h o l e s t e r o l  correlation Hilk  bilirubin  (-0,27) indicated  traits. on  were  and  existed  and  cholesterol  significant,  f a t percentage  that  (0,20) and between  f a t percentage (0.28)  a  was  negatively (-0.25). positive  and t r i g l y c e r i d e s ,  19  l i l J i S I M MS. METHODS a) B a i l C o l l e c t i o n The  and  animals  Analysis:  sampled  Hclstein-Friesian dairy in  the eastern  35  as  collect  herds  possible  technigues,  cattle  the  study  were located  of B r i t i s h Columbia.  These  were c h o s e n t o g i v e a s r e p r e s e n t a t i v e a  with  regards  microenvironments  production  present  i n commercial d a i r y herds  Fraser Valley region  co-operating  sample  in  data  to  within  from  a  nutrition, the  management  a r e a and a b i l i t y  recognized  data  to  collection  scheme. Specifically  the  characteristics. with  Several  a  stored  forages  exclusively  overall  classification  grass s i l a g e  included  traditional  freestall  systems,  systems were  varied  found  feeds.  throughout  of  cf  were a s s o c i a t e d  the  were used. summer  year.  and  feed  Included  were h a y , c o r n  and,  on  barns,  in  per Hany most  loose  Housing  Bucket and p i p e l i n e  farms systems  housing  b e i n g most numerous,  the  silage,  different  f e e d s were u s e d .  with the l a t t e r  with loose  the  silage  stanchion  stanchion  in  cows  cows  Several farms f e d conserved  stored  these  accordingly.  in  following  o f 40, 51, 56, and 59  system  and g r a s s - l e g u m e  combinations  the  and f e e d i n g s y s t e m s  pasturing  with  had  s i z e r a n g e d from 21 t o 136 m i l k i n g  management  utilized  supplemented  many  used  a guadrimodal d i s t r i b u t i o n  herd. farms  Herd  herds  and  ailking  milking  systems  barns, while v a r i o u s p a r l o u r  systems  and f r e e s t a l l  housing,  20 Cow  samplings  facilitate were  w i t h i n each  the v a l i d  discussed  out i n o r d e r t o  e s t i m a t i o n of a l l s o u r c e s o f v a r i a t i o n  in  considerations that  h e r d were c a r r i e d  the  literature  were employed  review.  i n the  The  that  methods  sampling  within  and herd  follow.  Age  at  lactation  sampling  scheme.  aniaals  with  start  Sampling  was  r e l a t i v e l y few  representative  animals and  in their  less  of  first  subject  of  Firstly,  lactation  would be  to s p e c i a l treatment  lactation,  concentrated  emphasize the were  in  relationships i t could  become  more  be  was  between  blood  expected  nutrients in relation  that  lactating This  days  selected that would  o f randomness.  of  lactation  start  Samplings  lactation  stress.  constituents these  was  secondly,  animals  also a consideration. 120  the  than older animals  Thus, younger  first  in  distribution  heavily  measured as d a y s from  the  pronounced  Seasonal e f f e c t s  skewed  less  period of greatest production  traits,  of  this  age.  t h e s t r u c t u r e o f many h e r d s ,  date of b l o o d sampling,  were  young,  of advanced  a sample i n k e e p i n g w i t h t h e a s s u m p t i o n  Stage to  consideration  c o n c e n t r a t e d on  were o f known p r o d u c t i v e a b i l i t y . yield  one  animals  done f o r a number o f r e a s o n s . was  was  and  If  to  there  production  relationships  will  i n periods of s t r e s s or marginal intake to  requirements,  were e s t i m a t e d by s a m p l i n g  each  herd  on  21 two  separate  occasions.  o u t between J u l y carried  out  sampling  seasons  climate,  19  and  p e r i o d s a t which adjust  to  sampled  i n both  The interest  28,  any  and  the h e r d s  sire  would  in this of  study. this  of The  that  they  of  would  the  ( l e s s than  standard  10 a n i m a l s .  On  subdivide the animals size.  The  total  this  sampled  for  which  at of  c o u l d be  to  were n o t  in  the  prime the  part  the  by  qenetic  (1959)  per group)  will  thus u n r e l i a b l e  stated  sire  are  constituents.  groups was  The  result in estimates.  according  the present study  ascertained.  time  sib analysis, small sib  1030  Hithin  serum s a m p l e s were a n a l y s e d , 27 s i r e  more d a u g h t e r s  represented  sampled and  Hobertscn  number o f a n i m a l s sampled  were a n a l y s e d f o r t h e serum  in  the s u b j e c t of  arrived  from  into  possible  met.  when s i b g r o u p s  basis,  two  adequate  heritabilities,  (1959), are minimized  These  also  c o u l d be  produce.  of  was  the c o n s i d e r a t i o n s of  10 i n d i v i d u a l s  error  as  estimates  and  carried  sampling  same i n d i v i d u a l s  were  when e s t i m a t i n g h e r i t a b i l i t i e s  Robertson and  The  had  was  1977.  numbers o f a n i m a l s  unacceptably high standard e r r o r s The  have  v a r i a n c e was  sampling  parameters  size  They  i t s assumptions  the r e l i a b i l i t y  group  F e b r u a r y 5,  i n order that  component  winter  much d i f f e r e n c e  s e a s o n a l changes. seasons  the  management.  considering  that  and  19 and  r e p r e s e n t e d as  d e s i g n and  distribution  summer b l o o d s a m p l i n g  1976,  between J a n u a r y  nutrition  statistical  The  between  10  was  s e t op  of  desirable  o f which the  groups  average  to  to  700  animals of  sire  10  or  group  22 size  was  approximately  with complete data. and  acceptable  Blood vacutainer samples  This  standard  was tubes  animal  should  for  biomedical  to c l o t  and  Serum was  split  into  The SHA  were c a l c i u m  phosphatase (T P ) ,  (Amyl),  (HC03), and  The  C.  T4  by  and  collected in These  the  into  blood  within  entire  subsamples of  immediately  2 ml  a  sample or  freezer  C until  6  sent  more. and to a  analyses. measured on  (Ca) , i n o r g a n i c  a Technicon phosphate  (Choi), albumin  (Alk P ) ,  (Na),  bilirubin which  potassium  present  AutoAnalyzer  (P04) ,  (Alb) , BUN,  glucose  uric  (Bili),  and  total  were measured on  (K), c r e a t i n i n e chloride  (Cl),  acid,  ether  (Great), bicarbonate  thyroxine.  i n i c n f o r m i n serum  chloride  a method m o d i f i e d  Biomedical  animals  estimates  were c e n t r i f u g e d  pipetted four  f r a c t i o n of  potassium,  determined  then  triglycerides (Trig),  constituents  sodium,  *B.  the  and  anticoagulant.  Other c o n s t i t u e n t s ,  e q u i p m e n t , were sodium amylase  for  SGOT, c h o l e s t e r o l  alkaline protein  1  545  reliable  s i x months a t - 1 8 °  serum c o n s t i t u e n t s  12/60  (Glue),  approximately laboratory  in  jugular vein  no  T h e s e s u b s a m p l e s were p l a c e d stored  for a t o t a l of  result  from t h e  containing  were a l l o w e d  one  animals  errors,„  obtained  hours of c o l l e c t i o n , . froB  20  and  bicarbonate.  from t h a t  Laboratories,  were  of K e s s l e r  Burnaby, B.  C.  calcium,  Calcius and  was  Helfman  23  (1964)./ from  This  modification  magnesium.  measurement  The d e t e r m i n a t i o n was  were  lilligrams  Sodium and p o t a s s i u m methods an  using  virtually  Units  method  and b i c a r b o n a t e  outlined  simultaneous determined analyzer.  determination  Units  Protein  of  measured  uric  acid,  determined and  bilirubin  and  were t o t a l  creatinine.  Determination protein  A l b u m i n was  analysed  by t h e method  was n o t e d t h i s serum  measurement BUS  reaction  technique  constituents,  was  this  carbon  the was a being  dioxide  metabolism  protein, Total  reaction  of t o t a l  It  both  constituents  protein  were grams  which a c o l o r i m e t r i c  for  were meg/1.  biuret  (1964). ,  in  other  both  related to  study  by a m o d i f i e d  Hochstrasser  units  this  with  (meg/1).  (1972). = B r i e f l y ,  with  o f measurement  in  flame  measurement  on a Beckmann c h l o r i d e  and compounds  were  spectrophotoletric  were measured i n t h e serum b y  by Kenny a n d Cheng  by c c l o r i m e t r y  hy  l i t h i u m , sodium  e l e m e n t s were m i l l i e g u i v a l e n t s p e r l i t r e Chloride  interference  (mg/dl).  were d e t e r m i n e d  standard.  any  c o l o r i m e t r i c and u n i t s o f  per d e c i l i t r e  a Beckmann p o t a s s i u m ,  internal lithium  removed  albumin,  by Skeggs  e t al.. (1971)  with bromcresol green  most  notably  and  (g/dl)4:  o f Doumas  from  was  colorimetric  per d e c i l i t r e  free  BUS,  protein  as described was  that  was  interference  with  bilirubin.  used. some  Units  of  were g / d l .  determination  et al..(1965).  This  utilized  method  the  method  reported  entailed the reaction  of  by Marsh  urea  with  24 diacetyl-monoxine color  development.  methods  The version based  to u t i l i z e  was c o l o r i m e t r i c  method  o f compounds t h a t  The a d v a n t a g e o f t h i s  was i n i t s a b i l i t y  Determination  in  i n the presence  of  of  uric  acid  were  mg/dl.  Total and  bilirubin  bilirubin  was of  differential with  units  by  reacted a  Determination  was e s t i m a t e d Gambino  and  was c o l o r i m e t r i c and  with  by t h e method o f J e n d r a s s i k  and  Schreiber  diazotized  caffeine-sodium  (1964).,  sulfanilic  benzoate  acid  reagent  Serum i n the  to  give  reaction,  o f mg/dl. was m e a s u r e d by t h e J a f f e Method w h i c h i n v o l v e d  cclcrimetric  determination of the  and  picrate.  alkaline  Enzymes  florganstern  resulted  of s t a b i l i z i n g  c o l o r i m e t r y between a s a m p l e and a b l a n k  Creatinine  phosphatase  which  I t was  y  Grof a s a d a p t e d  presence  agents.  (1965).  complex  s t a b l e c o l o u r development i n t h e presence  units  was an a u t o m a t e d  o f t h e method s e t o u t by Sobrinho-SimeT>s  intensifying  earlier  were m g / d l .  on r e d u c t i o n o f a p h o s p h o t u n g s t a t e  color  over  r e a c t i o n s o f low a c i d i t y .  and u n i t s  analysis  method  enhance t h e  reaction  ietween  U n i t s o f measurement were mg/dl.,  measured  in  the  serum  were  and a m y l a s e . , SGOT was d e t e r m i n e d et  SGOT, d i a l y z e d  a l . (1966). and c o u p l e d  salt.  T h i s method  control  and c o r r e l a t e d  creatinine  SGOT,  by the  method  O x a l o a c e t i c a c i d was g e n e r a t e d with  Fast  Ponceau  L,  e l i m i n a t e d t h e need f o r r u n n i n g well  alkaline  with  other determination  a  of from  diazcnium  a concurrent techniques.  25 This  was  a  colcrimetric  millilnternational Alkaline hydrolysis  for in  of  was  p-nitrophenyl This  c o r r e c t i o n with  phosphate as  with  by  use  and  a blank s t a n d a r d .  of  of enzysatic  subsequent  reported  interference  units  (mU/ml)•  measured  method,  a l . (1965), e l i m i n a t e d  color  by  Mcrganstern  by b i l i r u b i n  and t h e need  Units  o f measurement  were  mU/ml. The  by  per m i l l i l i t r e  phosphatase  development., et  Units  measurement,  measurement o f a m y l a s e  t h e method  method  was  reaction  outlined  by  Binderknecht  b a s e d on r e l e a s e  o f amylase  with  B e m a z o l b r i l l i a n t Blue.  l e v e l i n serum was  e t a l . (1971).  of soluble starch  an  insoluble  accomplished This  f r a g m e n t s due t o  starch  labelled  with  D e t e r m i n a t i o n was c o l o r i m e t r i c and  units  were Soraogyi u n i t s .  The  two l i p i d  cholesterol.  ;  Triglyceride  B u c c l o and D a v i d triglycerides hydrolysis  constituents  (1973) and  by  a  determination  of  spectrophotometer.  This  for  Units  Cholesterol e£ §1*  (1967).  involved  was This  involved  enzymatic  lipase  through  and  use  determination  and  was by t h e method o f  a  was s u b s e q u e n t l y c o u p l e d  glycerol  triglycerides.  determination  microbial  procedure  measured were t r i g l y c e r i d e s  of  hydrolysis protease.  to  an  The  enzymatic  absorbance  was s t a t e d  of  t o be  in  a  specific  o f measurement were m g / d l .  analysed  by  the  was a m o d i f i c a t i o n  method  of  o f an e a r l i e r  the r e a c t i o n of c h o l e s t e r o l with g l a c i a l  levine  method and  acetic  acid,  26 acetic  anhydride  cclcrimetry  and  and  sulfuric  glucose,  inorganic  determined  by  corrected  by  the  method  used  of  two  location  method  for  (DHI)  day  serum  Data  was  T4  Glucose  reduction described  from  the  measurement  formation by  of  stannous  by H u r s t  Slaunwhite  used i n t h i s  The  yields  first  of  (1964),  (1976)  thyroxine.  and  of P e r f o r m a n c e herds  study  measured  of milk,  milk and  Since the  during  the  particular  (BOP)  was  provincial  from  lactation  bleeding.  these  on t h e  obtained  E. C.  the  time o f  was  (1S74).  b a s e d on  fraction  from h e r d s  lactation  both  were  w i t h u n i t s o f mg/dl.  of the  program was  f e d e r a l Record  were c o l l e c t e d  by  was This  14  in  decilitre.  Victoria,  progress a t the  was  O n i t s were mg/dl.  by Cheung and  production data  in  protein  the  Mead  radioimmunoassay t e c h n i q u e s  sources.,  305  and  subsequent  This  determination  milk  included  the  and  m i c r o g r a m s per  Improvement  in  thyroxine.  measurement was  described  utilized  The from  determination.  acid  method  i n the  and Bcndar  a colorimetric technique  method units  of  phosphate  chloride-hydrazine.  The  measured  phosphate  a blank  phosphomclybdic  was  were  reaction involved a spectrophotometry  Inorganic  It  Determination  u n i t s were mg/dl.  O t h e r compounds which  determination  acid.  Dairy  central  Herd data  data  collected  fat  and  silk  the l a c t a t i o n  number o f h e r d s  program farm  obtained  was  visits.  in on  s m a l l , data Type  of  27 data  collected  herds  with  from  t h e BOP h e r d s .  but for  this  reduction  those  herds  vas i d e n t i c a l  data  protein  were a v a i l a b l e  i n the f i n a l the  numbers r e c o r d e d  not  to  at t h e time  a t the c e n t r a l data  several  with t h e s i r e  The  match  of  blood  locations,  reason  some  cow  sampling  deletion of  size,,  effects  effect  being  to  be a c c o u n t e d  fori n this  o f prime i m p o r t a n c e .  and c o v a r i a b l e s were n o t common t o b o t h  serum t r a i t s ,  appropriate  available  f o r 701 a n i m a l s ,  analyses.  inability  each dependent  model  variable  was  Since  production  analysed  by  e l i m i n a t e unique  effects,  a n a l y s e s were p e r f o r m e d  cn a d j u s t e d d a t a  to obtain estimates  various  genetic  obtained using least  parameters.  sguares  Adjustment  and then  the  to  the  DEI  Models:  were  effects  t o t h a t from  was  and m a i n t e n a n c e o f o p t i m a l s i b group  There analysis  was  present  b) S t a t i s t i c a l  and  Serum  o n l y 5 4 5 cows were used  outliers,  all  BOP  the e x c e p t i o n that milk  identification with  from  factors  constants  from  the  traits,  the  model  joint of were  appropriate  analyses. For  the  adjustments Yijk  milk  production  were b a s e d  was a s f o l l o w s :  = u > H i + S j • H S i j • b A i j k > cLijjk • € i j k  where u = overall  mean common t o a l l s a m p l e s .  Hi = t h e e f f e c t  o f t h e i - t h herd.  on  which  28 Sj = the e f f e c t HSij = the e f f e c t season.  o f t h e j - t h season of f r e s h e n i n g . o f t h e i - t h herd  Aijk = the covariable start. b = the coefficient Lijk  = the covariable  serum  age (mo.) t o a p p r o p r i a t e  lactation  with  Aijk.  length  associated  € i j k .= t h e u n e x p l a i n e d with i n d i v i d u a l  the  the j - t h  lactation  c = the c o e f f i c i e n t  For  to  associated  specific  with  (days). L i jk.  environmental deviations samples.  traits,  associated  t h e model on which a d j u s t m e n t s were  b a s e d was a s f o l l o w s :  Tijk  = u + Hi • S j • HSij  • bAijk  + eijk  + cLijk  where u = overall  mean common t o a l l s a m p l e s .  Hi = the e f f e c t  c f the i - t h herd.  Sj = t h e e f f e c t  c f the j - t h season o f b l e e d i n g .  HSij = the e f f e c t season, Aijk  of  = the covariable which b l e e d i n g  b = the coefficient lijk  specific to the j-th  age (mo.) t o s t a r t occurred, associated  with  of  lactation  = t h e unexplained with i n d i v i d u a l  associated  with  in  Aijk.  = the covariable days from lactation b l e e d i n g date (stage o f l a c t a t i o n ) .  c = the coefficient eijk  t h e i - t h herd  start  to  Lijk.  environmental deviations samples.  associated  29  The  effects  o f herd,  season  and h e r d  were t r e a t e d a s f i x e d  effects.  normally  and independent  zero.  distributed  Constants  appropriate analysis.  were e s t i m a t e d  terms The  in  raw  i l l  data  were  adjusted  recognizing  t h a t an i n s i g n i f i c a n t  Using estimated  regardless  significantly  the adjusted from  of  be  tested f o r  a l l  squares  f o r a l l appropriate  significance.  adjustment  nor introduce  data, the s i r e  to  with e x p e c t a t i o n s e q u a l t o  t h e above models b y use o f l e a s t  terms,  the data  interaction  e f f e c t s were assumed  and s i g n i f i c a n c e  adjustment  alter  x season  term  T h i s was done would  neither  bias.  component o f v a r i a n c e was  t h e f o l l o w i n g model  Yij = u + Si + € i j where u = overall  mean common  Si = the effect €ij  Sire normally  c)  of the i - t h sire.  = the unexplained environmental with i n d i v i d u a l samples.  was t r e a t e d a s a random e f f e c t distributed  Referring  t o Table  components  deviations associated  and was assumed  to  be  with expectation equal to zero.  Estimation of Variance  following  t o a l l samples.  Components; I,  this  of variance  present  study  utilized  (with d e f i n i t i o n s  the  as o u t l i n e d  30  Table  I . , A n a l y s i s o f v a r i a n c e (ANO?) t a b l e s q u a r e s (EMS) f o r t h e e x p e r i m e n t a l  and expected models.  mean  —i  i I  |  Sources o f  least  1  variation  Square  Cgrrgctj,gn  r  I I I  Covariables i Aqe (months) | Staqe o f l a c t a t i o n ( d a y s ) |  I  Herd  I  J  Season  I  I  H x S  I  |  Genetic  |  Sires  I  1  d.f.  2 1 1  EMS -1  ! I  0 e *  d.f.se  I  0*e + k 3 6 2 s e  d.f.hs  I  0*e •  k26*hs  6*e •  kie^s  d.f.h  !  Individual/Sires  ! ! 2  k4e*h  !  J  d.f. s i r e s I  I  d.f. e  i  Q2e i  shere 1  k1 =  (N.,;- ( g  For production  traits,  Ni.)*/N. . ) / d . f . s i r e s t h i s becomes l e n q t h o f  lactation.  31  by  Falconer  (1960):  6 e •= t h e component c f v a r i a n c e a s s o c i a t e d w i t h i n d i v i d u a l s within sire groups. This component of variance contains 3/4 o f t h e a d d i t i v e g e n e t i c v a r i a n c e {V ( A ) ) and a l l the environmental variance (V(E)). Honadditive genetic variances were assumed t o be n e g l i g i b l e i n t h i s system. 2  C s  = t h e component o f v a r i a n c e a s s o c i a t e d w i t h d i f f e r e n c e s among s i r e g r o u p s . T h i s was assumed t o l e a d to an estimate of 1/4 V (A). Variances a s s o c i a t e d with a d d i t i v e by a d d i t i v e i n t e r a c t i o n s were assumed t o be n e g l i g i b l e i n t h i s system.  2  The v a r i a n c e components o f V (A) ,  V(E)  calculated  d)  and  =  V(E)  = e e  V(P)  = V(A)  variance  this  study  were  V(P).  These  were  relationships:  2  -  2  Variance  this  in  4C s 30 s 2  • V(E)  of Variance  Variances  in  phenotypic  using the following  V(A)  formulae  the  interest  of variance  given study,  1) V a r ( V ( A ) )  Components:  by S e a r l e  components (1971a).  the appropriate  =  were  calculated  F o r t h e components  sampling  variances  by  the  estimated  were:  16Var(6 s) 2  where Var(6 S) 2  = 2fl*eN. .  (N. «-r1) (A-1) / ( N . . -rA) (N, . > S 2 )  2  2  2  + ( 4 6 e c S l J . . )/(N..2-S2) 2  2  + 26*S(N.. S2 • S2 2  2  - 2N. . S 3 ) / ( N . .  2  . - S2)  2  32 and  »••'..= Z N i . 52 =  iNi.z  53 = 2 N i . 3 A = the Ni. 2)  Var(V(E))  = the  number o f s i r e  groups  number o f s a m p l e s i n t h e i - t h s i r e  =  Var(62e)  + 9Var(62s)  *  26*e/ (N. .--A)  -  group  6cov<e2ee*s)  where Var(fi2e) cov(62ee s) 2  3)  = -N. . (A-1) Var ( 6 e ) / ( N . .2  Var«V(P)} =  Var(V{A) +  =  Var ( 6 2 s +  = Var ( 6 s ) 2  €  )  Heritability  V(E)) 6 e) 2  • Var(62e)  (1960).  estimates  were  Intraclass  and  environmental  hz * 4 t =  2cov(6 e62s) 2  derived  as  described  c o r r e l a t i o n s (t) between  g r o u p s were e m p l o y e d , making use sire  •  Estimates;  Heritability Falconer  -S2)  2  of the  factors  components o f v a r i a n c e .  V(A)/(V(A) • V(E))  =  462s/(62e *  halfsib  comprising In  general,  6 S) 2  by  the  33 *~) V a r i a n c e o f H e r i t a b i l i t y Variances  of  making use o f t h e intraclass  the  Estimates:  heritability  relationship  correlation  estimates  between  between  a modified  =  and  The  for halfsib  the  sampling  analysis  was  (1959).  16Var(t)  = 32f 1 + ( k 1 - 1 ) t ] 2 ( 1 - t ) a / k 1  9) C o r r e l a t i o n  <k1-1) (A-1)  Estimates;  Estimates  of  genetic,  c o r r e l a t i o n s f o r each p a i r tabulated  groups.  v e r s i o n o f t h a t r e p o r t e d by R o b e r t s o n  Var(hz)  calculated  heritability  halfsib  variance of the i n t r a c l a s s c o r r e l a t i o n  were  environmental  of traits  and  phenotypic  measured i n t h i s  study  were  using the f o l l o w i n g formula;  r(XaXb) =  cov(XaXb)/SXaSXb  where r(XaXb) = t h e a p p r o p r i a t e c o r r e l a t i o n Xb (a#b) QXa = t h e a p p r o p r i a t e trait 6Xb = t h e a p p r o p r i a t e trait  standard standard  between t r a i t s  deviation deviation  and where cov (XaXb) was t h e s o l u t i o n the f o l l o w i n g g e n e r a l f o r m u l a ; Var (Xa+Xb)  = e Xa a  • €*Xb • 2cov(XaXb)  of of  Xa  and  the a - t h the  b-th  o f a rearrangement o f  34 h)  Variance of Correlation  Estimates:  Variances of the c o r r e l a t i o n s estimated  Est.  by t h e a p p r o x i m a t i o n  calculated  i n t h i s study  g i v e n by S c h e i n b e r g  (1966).  V a r (rXaXb) = 2 u r (XaXb)/k 1 f ( s V a / d . f , s i r e s w v a/d.f.e)/4C*a 2  2  2  were  2  *  2  *• ( s v z b / d . f . s i r e s +  w*v b/d.f.e)/40*b 2  • s [ (VaVb + V a b ) / d . f . s i r e s • w v a b ) / d . f . e) ] / 2 c o v (Xaxb) 2  2  2  (vavb •  2  - (sVaVab/d.f. s i r e s + w vavab/d.f.e)/6 acov(XaXb) 2  2  - (sVbVab/d.f. s i r e s • w v b v a b / d . f . e) / O b c o v (XaXb) 2  •  2  (sV ab/d.f.sires 2  • w v ab/d.f.e)/26 a0 b} 2  2  2  2  where Va = s i r e  mean s g u a r e  associated  with t r a i t a  Vb = s i r e  mean s q u a r e  associated  with t r a i t  Vab  = sire  b  mean c o v a r i a n c e f o r t r a i t s a and b  va = r e s i d u a l  mean s g u a r e a s s o c i a t e d  with t r a i t  a  vb = r e s i d u a l  mean s q u a r e a s s o c i a t e d  with t r a i t  b  vab = r e s i d u a l  mean c o v a r i a n c e f o r t r a i t s  a and b  and u = 16, 1 and 1 f o r t h e genetic, environmental phenotypic c o r r e l a t i o n s r e s p e c t i v e l y  and  s = 1, 9 and 9 f o r the genetic, environmental phenotypic c o r r e l a t i o n s r e s p e c t i v e l y  and  w = 1, k1+3 and k1-3 f o r t h e g e n e t i c , e n v i r o n m e n t a l and phenotypic c o r r e l a t i o n s r e s p e c t i v e l y  35 BESPITS JND  DISCUSSION  Before s u b j e c t i n g parameters, systematic of  least  the data  to  squares  h e r d , s e a s o n , h e r d by s e a s o n account.,  interaction,  covariables,  was  records, this  was t h e age t o s t a r t  current at  of the l a c t a t i o n  second  covariable  milk p r o d u c t i o n bleeding  in  traits  the  and  case  of  one  of  lactation.  For both  was t h e age  sampling  took  lactation  at  s g u a r e s c o n s t a n t s a s s o c i a t e d w i t h h e r d and t h e  term  were  to  be drawn from  constants  these  with  c o v a r i a b l e s a l l o w e d us t o s t a t e trait the  was  covariable.  sguares the  h i g h e r o r how  i s  The  than w r i t t e n  most  the  two  the  time  The  i n which season  the l e v e l  actual  abridged  and  with table  the  a  two  of  a  respect to of  This table  and t h e c o v a r i a b l e s i n  were  sguares  the l e v e l  was c h a n g i n g  of  interaction  least  seasons  presented i n Table I I .  trends f o r the seasons  concise  However,  F o r t h i s r e a s o n , an  constants  of the  s i n c e no s p e c i f i c i n f e r e n c e s  effects.  associated  place.  i n the case  t h e serum c o n s t i t u e n t s .  interest  the  lactation  least  of l i t t l e  of  For f i r s t  of l a c t a t i o n  stage  effects  covariables  this covariable  i n w h i c h serum  was l e n g t h  The  start,  of f i r s t  genetic  identifiable,  and two  a l l traits.  p r o d u c t i o n a n d serum t r a i t s ,  start  The  to  for  the  were u n d e r t a k e n . ,  Age a t l a c t a t i o n  common  for  adjustments  environmental e f f e c t s  were t a k e n i n t o  analyses  form  least shows more  text.  noticeable of these l e a s t  squares  c o n s t a n t s were  Table I I .  Least squares w i n t e r seasons  c o n s t a n t s a s s o c i a t e d with and with the c o v a r i a b l e s .  —•  "-  ' * I Trait |  Sumner  Winter  Age  suamer  Length  and  — ————i Stage  |  *== = = = = M = ========== ========= ========================== ==-=====.f 1 Hilkl  |  -214.4  214. 4  41.98  28.56  na  |  | Fat 1 |  -6. 351  6. 351  1. 6C5  1. 250  na  |  I Protl |  -5. 454  5. 454  1.549  .9522  na  |  I HilkC |  -195.3  195.3  22. 36  34.26  na  |  I Fat C |  -5. 708  5. 708  . 8728  1.272  na  |  I Protc |  -4.556  4.556  . 6318  1. 127  na  |  1 Ca  |  . 0379  -.0379  -.0021  na  .0005  |  I P04  |  . 0139  -.0139  -.0104  na  .0011  |  ! Glue  j  -2. 090  .0194  na  . 0 140  |  I BUN  |  . 1333  -.1333  0178  na  .0098  |  ! Uric  |  . 1130  -.1130  -.0001  na  -.0004  |  I Chol  |  -6. 949  -. 1069  na  .0523  |  I T P  |  . 0406  -.0406  . 0100  na  -.0006  |  I Alb  |  -.2449  .2449  . 0004  na  .0003  |  1 Bili  |  -.0049  .0049  .0002  na  -.0002  |  1 Alk P |  -.6511  .6511  -. 1720  na  .0187  |  | SGOT  |  -.4055  . 4055  -.0543  na  .0536  |  I Creat |  -. 0693  . 0693  -.0004  na  -.0000  | |  2- 090  6. 949  1 Trig  |  . 0890  -.0890  0122  na  .0030  1 Na  |  -.8073  .8073  -.0154  na  .0016  I K  1  .0167  -.0167  -.0009  na  -.0000  |  1 CI  |  . 6487  -.6487  -.0071  na  .0050  |  I HC03  |  -.9059  . 9059  -.0044  na  -.0019  |  1 T4  |  2352  .2352  -.0087  na  .0047  |  I  |  -8. 706  8. 706  .5502  na  .0122  |  fiffiyl  J  37  those  associated  lactations. of  The  production  compared over  an  had  higher  present  and  examined  p o r t i o n of t o t a l effects  in this  the  alkaline  study,  phosphatase  relatively effects  variability however,  in  relative model.  Thus the  that  statistic  for  means and  the  study  partial B the 2  2  effect  of herd  was  from  for  the t r a i t s 0,20 8  for  This indicated  the  total levels,  a l a r g e p o r t i o n of  yielded  an i n d i c a t i o n i n the the  became a of  summarized  the  environmental  determining  importance  standard e r r o r s i n Table  accounted For  beyond whether  of determination are  total  F o r serum a l b u m i n  s i g n i f i c a n t , and  relative  production,  systematic, in  in  model  separate e f f e c t s  went  reflected  f o r the  ranged  2  {23-36  age.  above.  were d e t e r m i n i n g  of  was  the  phosphatase.  partial 8  effect  Coefficients  The  The  importance  of  this  effects  variability.  2  that,  show as s t r o n g a  variability  B  lactation  constants f o r  t o 0.704 f o r a l b u m i n .  of a l k a l i n e  these  (B )  listed  total  s m a l l importance utilized  was  lactation  much l o w e r  of determination  measures  lactations  which  Current  had  environmental  current  This indicated  23-157 months, d i d not  consequently  a measure o f t h e  total  of  the  first  in f i r s t  a more p r o n o u n c e d e f f e c t  range  and  for  for current lactation.  coefficient  the  first  constants f o r a l l three  considerably  those  age  The  for  sguares  the  higher a s s o c i a t e d constants.  relationship  by  in  t h e s m a l l e r r a n g e o f age  with  is  age  least  were  to  m o n t h s ) , age the  with  the  adjustment adjustment  discriminating  that with  of  the  adjustment. least  squares  III. ,  significant  for a l l traits in  this  38  Table  I I I . L e a s t square means and a s s o c i a t e d c o r r e c t i o n terms.  Trait  Milkl  |  Mean  SE  standard  errors  (SE)  with  R*  fora l l  Staqe  Total |  T |  Herd  Season  HxS  Aqe  lenqth  |  5994kq  45. 2  I  .256*  .029*  .085*  . 012*  .068*  na  .494  |  1 |  222.kq  1. 79  I  .224*  .017*  .086*  .012*  .087*  na  . 465  |  195. kq  1. 52  I  .262*  .0 20*  .064  .017*  .054*  na  .464  |  MilkC |  7160kq  66. 2  I  .184*  .009*  .050*  .108*  . 127*  na  .553  |  Fat C |  262. kq  2. 59  I  .166*  .005*  .056*  .120*  .127*  na  . 505  |  ProtC  |  230.kq  2. 1 1  I  . 198*  . 005*  .053*  .095*  .142*  na  .533  |  Ca  |  9. 54mq/dl  .021  I  .230*  .003  .206*  .008*  na  .002  .471  |  P04  |  4. 98mq/dl  . 039  I  .114*  . 000  .1 18*  .072*  na  .004  .354  |  Glue  I  5a.1mq/dl  . 316  I  .188*  .036*  .298*  . 002  na  .006*  .561  |  BUN  |  114. 1mq/dl  . 125  I  .333*  . 001  .304*  .010*  na  .014*  .671  |  Uric  t  1. 07mq/dl  .011  I  .165*  . 104*  . 1 19*  , 00 0  na  .005*  . 438  |  Chol  |  206.mq/dl  1.78  I  .210*  .018*  .121*  .003  na  .004  .375  |  T P  |  7.62q/dl  . 022  I  . 138*  .003  . 086*  .154*  na  .003  .481  |  Alb  |  3.66g/dl  .013  I  . 302*  . 189*  .120*  .000  na  .001  .704  |  Bili  |  .170my/dl  . 00 4  I  .177*  .002  .063  .004  na  .010*  .317  |  P |  UO. 7 mil/ml  . 960  I  .089*  . 001  .060  .038*  na  .002  . 208  |  |  136. mU/ml  1. 18  I  .289*  .000  . 1 33*  .002  na  .008*  . 447  |  Creat |  .9 57mq/dl  .005  I  .128*  .158*  .189*  .004*  na  .000  . 552  |  Trig  1  9. 68mq/dl  . 121  I  .243*  .001  .136*  .009*  na  .002  . 402  |  Na  l  141. meq/1  . 127  I  .272*  . 035*  .197*  .010*  na  .000  . 542  |  K  1  4.24raeq/l  .016  I  .142*  . 001  . 124*  .003  na  .000  .310  |  CI  |  97.2neq/l  . 108  I  .237*  .029*  .240*  .003  na  .006*  .570  |  HC03  |  23.3meq/l  .080  I  .276*  .092*  .223*  .002  na  .001  .622  |  T4  |  5. 25yug/dl  . 058  |  . 131*  .016*  .216*  .017*  na  .024*  .470  |  A my 1  j  193. SOD. 0  3. 58  I  .159*  .008*  .073  .024*  na  .000  .306  |  Fat  Protl  Mk SGOT  |  * S i g n i f i c a n t a t P<0.05  -  1  39 study,  accounting  v a r i a b i l i t y . „.  This  management and agreed  with  microenvironment  Hanston  important  S,9%  between reflected  earlier  Rowlands and most  for  papers 1976)  and  the  33,3%  effects  of  of  nutrition,  a s s o c i a t e d w i t h each (flewett  1974,  which s t a t e d in  determining  factors  in  the  herd,  and  Payne e t a l .  that herd  factor  total  the  1974,  was  the  single  level  of  serum  constituents. The  other  significant discussed  o r as  in detail  Season was accounted  important  for  as  the  herd  significant only  a  f o r a l l milk production  small  proportion  of  schemes, and  s i n c e a l l herds  t h i s adjustment  was  a  d i d not  significant  exception  of  season  significant  a  are  traits  but  variability.  various  warranted cow's  i n view  was  much l a r g e r  lactation  traits.  constants  were  of  environment.  p r e s e n t on  a year-round  dairy  pasture  many  farms.  basis.  Also,  e x p e r i e n c e a p a r a l l e l s e a s o n a l change i n interaction  T h i s was  found  first  still  s t o r e d f e e d s on  expected.  for  They  i n t h e s e s e a s o n a l c h a n g e s were t h e c h a n g e f r o m  farms u t i l i z e d  feed,  the  has l o n g been r e c o g n i z e d by  to s t o r e d f e e d , a management s y s t e m Other  universally  effect.  t h e s e a s o n a l c h a n g e s i n many a s p e c t s o f t h e Included  as  at t h i s p o i n t .  Adjustment f o r season recording  model were n o t  for a l l  lactation  between  herd  season  was  production  traits  with  the  protein yield.  I t was  noted  that  f o r a l l production t r a i t s  p r o p o r t i o n of the This indicated  similar  and  {Table  variability  that,  while the  I I ) , t h e r e was  but  accounted  i n the c u r r e n t least  sguares  relatively  more  no residual  error  resulted  involved  in less  lower p a r t i a l  A  v a r i a b i l i t y accounted f o r  factors  rationale  for  a  means of  for  this  factors  production  age  was  i n determining  situation  first  this an  higher yet  for  partial R  indicated  This thus  l e n g t h have  by  the  a  also  records.  significance  variability  indication  For  length  for  first  of of  first  lactation.  first  lactation  were  age  and  least  of  all  l e n g t h as sguares  of  milk  This resulted  lower o v e r a l l  respect  residual  least  strength  first  v a r i a b i l i t y around  in  lower p a r t i a l  of  the  traits,  to l a c t a t i o n in  traits.  lactation  length,  yet was  R, 2  regression.  s g u a r e s c o n s t a n t s were  lactation  for  lactation  lactation  regression of  the  did  constants  compared t o c u r r e n t  Again, t h i s r e s u l t e d  similar residual  greater  a lower p a r t i a l B  2  in for  traits. , serum  interpretation straightforward  large.  current  with  the  slope,  lactation,  and  variability,  although the  was the  for the  lactation  a larger  regression  effects  and  production  were h i g h e r f o r c u r r e n t  z  that  exhibited  For  the  existed  were  age  season  lactation  reinforced  R e g a r d i n g age,  on  and  standardizing  seasonal e f f e c t s .  This  by  traits. ,  traits.,  similar  traits,  lactation  2  been u t i l i z e d as  t h e s e two  current  R.  Adjustment  The  in the  of as  for  constituents, the the  significant  the  adjustment production and,  in  significance  terms traits.  was  not  Although  general, accounted  for  and as herd the  41  largest  portion  varied  of the v a r i a b i l i t y ,  greatly  with  regards  the other adjustment  to  significance  and  factors relative  importance. „ Herd  effect  changes i n the nutrition,  I t was  animal  will  functions. directly  l e v e l s o f serum t r a i t s  metabolism  feeding,  herds.  a n i i r a l and  on  that  altered  by  and  constituents,  to  differences  indirectly  which are  interrelated.  chronicle  the  Specifically,  was  expected  in  seasonal  by  parallel  mentioned,  changes  the  effects in  pasture  these  i n the  the  animal  were  expected  and  stored  seasonal  by s e a s o n  covariables associated  w i t h t h e serum  to  environment. feed  altered  metabolism  and  expected.  changes  over a l l herds, a herd  serum  constituents  inputs to  particular  both  other  i n p u t s t o the animal's  i f  among  i n i n p u t s to the  q u a n t i t a t i v e c h a n g e i n some serum t r a i t s c o u l d be previously  in  which  affecting  Seasonal  t h e change from  the n u t r i t i o n a l  differences  t o have i t s e f f e c t  m e t a b o l i c pathway i n v o l v i n g or  of  various metabolic a c t i v i t i e s  due  This alteration  the  a reflection  microenvironment  d i f f e r e n c e s i n the environment  on t h e  greatly  about  management,  expected be  brought  was  interaction  a As  were  not  could  be  age  at  expected. The lactation  start  and  s a m p l i n g , were b o t h time.  As  occur  and  stage  of  the affect  stage  associated  animal the  lactation  of  lactation  of  proceeded,  at  with changes i n  matures,  levels  traits,  m e t a b o l i c and some  serum  h o r m o n a l and  time  of  blood  metabolism hormonal  changes  constituents. nutritional  over  As  balance  42 stress  were  both  involved  in  the  altering  of  some  serum  constituents. Calcium a  number  of  adjustment effects  body  were  (1976),  inorganic  al.  to  has  been  season  insignificant BON  stage of  levels  in  was  of  Prewitt et However,  to  this  (Little  have  a nonsignificant fact that was  not  this  been p r e v i o u s l y r e p o r t e d age  (Kitchenham et Tumbleson  In  and  of  and  reported  as  by  influence  significant  effects 1975*  Hutcheson  an  disruptive.  Seasonal  al.  of  (Hewett  herd, season, herd  lactation.  significance  reports  effect  seriously by  (Boss  significant  study  not  present  and  Stage  a  stage of  and  with  (1975)  other  and  lactation.  al, as  or  effect  The  age  agreement  significance.  reported  significantly  BON  such  the  disagreement 1976  in  s e a s o n , but  significantly affected  a l . 1974).  Rowlands  by  no  l e v e l s , so  was  the  and  p h o s p a t e have b e e n r e p o r t e d  affected  variously  serum g l u c o s e has  have b e e n i n  and  be  found  interaction  (Payne e t  and  inorganic  lactational  G l u c o s e was  on  and  (Rowlands e t a l , „ 1975)  insignificant  interaction  K i t c h e n h a m e t •••al.  1976)  calcium  season  in  trends for  (1973b)  lactation  on  same  This  a l , 1974)  1974)  by  the  interrelated  significant,  (Payne e t  effect  showed  Herd, herd  C a l c i u m and  Halliday  p h o s p h a t e , which are  systems,  factors.  Tumbleson e t  and  and  on  glucose  Kitchenham  1971)  but  was  interaction,  age  study. affected  view of  Hanston  by  the  1972,  expected  herd,  effect Hanston to  of et  1971),  i t  was  be  pointed  out  e a r l i e r , management on  diet  on  serum  a l . 1975,  and  significant. many f a r m s  now  includes herd  stored  feeds e x c l u s i v e l y .  seasonal  effect  on  major c h a n g e s i n d i e t . significance  the  exception The  was  lipids  was  there  adjusted  The  g e n e r a l agreement w i t h  by  a l l factors  that  were  the  with  was  cholesterol  little  similarity  for  herd,  season  between  adjusted  interaction  f o r herd,  interaction  above s i t u a t i o n f o r c h o l e s t e r o l  previously  age was n o t e d  cholesterol  and  and  published  studies  by P e t e r s o n and Ha I d e m  by Tumbleson and H u t c h e s c n (1971) t o have a on  for  f o r the s i g n i f i c a n c e o f t h e adjustment terms.  effects.  exception  support  by  term.  w h i l e t r i g l y c e r i d e s were  age  within  largely affected  indicated  measured  although  these c o n s t i t u e n t s  and  i n turn  the  o f age.  triglycerides,  effects  This  s i g n i f i c a n t l y affected  serum  Cholesterol  would l e s s e n  any c o n s t i t u e n t s  of the i n t e r a c t i o n  Uric acid  This  level.  This  age  r a n g e o f age p r e s e n t , w h i c h c o u l d  explain  with  the  the  (1978) and  significant  e f f e c t was  was i n  effect  d e p e n d e n t on t h e finding  in  this  protein  and  study. , Measures o f p r o t e i n albumin.  Albumin  interaction, age  while t o t a l  effects.  to  lactation, albumin  and  protein  (1973b) who  be  received  was  total  close  reported  total  protein  (1975)  affected and  by h e r d ,  age and  f u r t h e r adjustment f o r  in  significantly  Rowlands e t a l . total  were  s i g n i f i c a n t l y affected  This  Tumbleson e t a l . albumin  was  i n t h e serum  by  Little  agreement protein age.  with  but In  (1974)  not early found  b o t h t o be s i g n i f i c a n t l y a f f e c t e d  by  44 stage of l a c t a t i o n , study.  Hewett  closely  a result  (1974),  approximating  not forthcoming  using  that  a range  in  the  r e l a t i o n s h i p of stage o f l a c t a t i o n Bilirubin Tumbleson  was a f f e c t e d  and  Hutcheson  by  differed  with t h e p r e s e n t study  affected  by a g e .  Alkaline was  in  1973a).  phosphatase  agreement  with  comprised  This part  significant  indicated  of  while  the  been c i t e d All  climatic  as an e f f e c t  levels.  supported t h i s  Sodium, electrolytes  and  in  and  the are  on  age.  and  This el; a l , .  stage  of  effects  which  terms  were  The r e s u l t s f o r  p a s t s t u d i e s i n t h a t age  lactation  1978)•  were  utilized  The a v a i l a b l e l i t e r a t u r e  chloride  in  on t h i s  ( P e t e r s o n and H a l d e r n  and  bicarbonate  which a r e i m p o r t a n t  instrumental  interaction  effect  of  also  1968).  blood  A l l four  were n o t .  (1968)  was  (Tumbleson  interaction  present finding  potassium,  and  eguilibrium.  seasonal  bilirubin  by h e r d  from  no  lactation.  and H e a l y  ( P e t e r s o n and Waldern  creatinine  and H y l r e a and H e a l y  functions  differed  stage  constituent  and  found  of  nutritional  changes  f a c t o r s except  adjusting  season  Mylrea  studies  that  herd  SGOT i n t h e p r e s e n t s t u d y  1978  stage  i n stating  previous  more  albumin.  and  was a f f e c t e d  present  stage  study,  SGOT was a d j u s t e d f o r h e r d , i n t e r a c t i o n  lactation.  has  with  and  the  of lactation  present  herd  (1971)  from  in  constituents factors potassium.  with  i n many b o d i l y  maintaining  were the  are  affected exception  Additionally,  osmotic by h e r d , of the  sodium  was  45 significantly lactation. by  affected The  by  absence  previous  age  and  chloride  by  stage  o f age e f f e c t s was g e n e r a l l y  studies  of  supported  (Tumbleson e t a l . 1973b  and  K i t c h e n h a m e t a l . 1976). Thyroxine agreed Mixner  was s i g n i f i c a n t l y  with  results  of  Flamboe  by a l l f a c t o r s .  and  Beineke  This  11959)  and  e t a l . (1962).  Amylase was s i g n i f i c a n t l y  Once  significant  eliminated random  from  components  present  heritabilities,  additive  study,  Since  this  of  study  of the genetic v a r i a b i l i t y  that  the  environmental  effects exclusive of the additive  were and  of  the  environmental  efficiency.  variance  nonadditive genetic v a r i a b i l i t y ,  study  and  environmental  present  effects  Because  genetic  and a g e .  only genetic  high r e l a t i v e  estimates genetic,  that  remained.  the  separable with  and  portion  estimating  environmental  variability  yielded  correlations.  by h e r d , s e a s o n  t h e d a t a , i t was presumed  were  separation  affected  systematic,  environmental  design o f t h i s  this  affected  This  components, and  phenct y p i c  estimated  only  the  and had no means o f i t  was  parameters  assumed  in  included a l l  effect.  T a b l e I ? r e p r e s e n t s a summary o f t h e g e n e t i c , e n v i r o n m e n t a l and each  p h e n o t y p i c v a r i a n c e s and t h e h e r i t a b i l i t i e s trait.  It  was  apparent  v a r i a n c e and some h e r i t a b i l i t i e s This  was  instigated  in  cases  that  associated  with  some g e n e t i c components o f  were a r b i t r a r i l y  set  to  zero.  where t h e g e n e t i c v a r i a n c e was  Table  IV.  V a r i a n c e components and h e r i t a b l i t i e s with a s s o c i a t e d the i n d i v i d u a l t r a i t s .  Trait  |  Milkl  | 15. 25  68728±64289  1 | 15. 25  Fat  Protl  k  | 13. 56  V (A) ±SE  V (E)±SE  SE f o r  V(P)±SE  h*iSE  571378±69953  640106±43127  .10741.0976  153.9±113.9  852.2±116.2  1006.168.37  .15301.1078  106.0±83.77  576.2i85.33  682.3±49.09  .15541.1172  903688±187688 1362593193041  .33701.1382  MilkC | 17. 35 459265+217238 C | 17. 35  587.5±297.6  1495.±265.9  2082.1138.8  .28211.1270  ProtC | 15. 52  534.7±249.6  809.2±210.9  1344.±99. 14  .39791.1560  Ca  | 18. 10  0.0  . 1950±.0167  .19501.1190  0.0  P04  | 17. 48  ,0086±.0409  ,6078±.0550  .6164±.0384  .01391.0664  G luc  | 18. 13  0.0  42.78±3.661  42.78±2.618  0.0  BON  | 18. 14  .3057±.4792  6.312±.5952  6.618±.4060  .0462±.0718  Uric  | 18. 17  .0002±.0034  .05481.0047  .0550±.0034  .0045±.0616  Chol  | 18. 06  0. 0  1352.±116. 1  1352.±82.92  0.0  T P  | 18. 13  .0015t.0133  .2119±.0184  •2134±.0131  .00701.0624  Alb  | 18. 13  .0064±.0063  .0689±.007T  .0753±.0046  .08521.0810  Bili  | 18. 03  0. 0  .0068±.0006  .0068±.0004  0.0  P | 18. 06  116.4±56.74  272.4±49.94  388.8±25.65  .29941.1283  | 17. 93  30.53±43.73  554.9153.46  585.4136.14  .05221.0739  C r e a t | 18. 28  .0035±.0016  .00731.0014  .C108±.0007  .32481.1328  Trig  J  18. 17  0.0  6.3391.5417  6.3391.3876  0.0  Na  | 17. 83  0. 0  6.9501.6025  6.9501.4290  0.0  K  | 17. 83  .0144*.0108  .09721.0113  .11161.0070  . 12881. 0921  CI  | 17. 87  .0837±.3218  4.8121.4310  4.8961.3020  .01711.0657  HC03  | 17. 87  0. 0  2.7201.2354  2.7201.1677  0.0  T4  | 18. 44  0.0  1.4721.1244  1.4721.0893  0.0  Amyl  | 17. 82  1082. ±6 30. 1  4265.1601.6  5347.1341.9  .20241.1085  Fat  Alk SGOT  47 negative, resulting theoretical value  was  was  very  in  lower  taken  limit  negative of  was  to one  heritability.  heritability  as a s t a t i s t i c a l  close  heritability  a  zero.  estimate  Taken  t h a t was  Since  i s zero, a  the  negative  of a h e r i t a b i l i t y t h a t  another  way,  not s i g n i f i c a n t l y  a  negative  different  from  estimates  from  zero. Comparison this  and  components  p r e v i o u s s t u d i e s was  traits.  The  present  lactations. first  of the  It  lactations  study  was  noted  i f the  lactation.  Any  on  lactations  current  regards comply  to the  reliable  f e e d i n g and  Estimated Butcher with study.  Variance  somewhat due was  estimated  lower  than  current  were sampled d u r i n g t h e i r  first  subject  to  based cn f i r s t  considerations  different  less  with  measures  genetic  theory.  modern  biased  or  lactation  r e c o r d s h a v e been c o n s i d e r e d  are  by  more  selection  or  management.  of  variance  a l l lactations from  these  agreed  reported  by  reasonably  well  current lactations  estimates.  t o a number o f f a c t o r s . , lower  and  indeed  components a s s o c i a t e d w i t h than  production  were  r e g u i r e d by  components  e t a l . (1967) on  those  were  they  first  between e s t i m a t e s  lactation  as  both  milk  that current lactations  comparisons  first  insofar  preferential  used  animals  assumptions  variance  p o s s i b l e for the  degree t o which the  with the  Historically,  of  Since  first  i n this  lactation  This discrepancy  first  present  lactation  were  could  be  production  c u r r e n t l e v e l p r o d u c t i o n , i t appeared t h a t  the  HQ  v a r i a n c e and indicating  t h e mean o f t h e s e a  nonnormal  c o u l d h a v e been t h a t lactations sires  lactations. confounded  to  range  to  tended  the  first  to  This  indicated  that  extent  or  sire  that  for current  age  h e r i t a b i l i t i e s f o r the  1957).  while  younger  their  first  and  age  were  adjustment  was  production t r a i t s f a l l  into  the  (Butcher et a l .  1967,  H e r i t a b i l i t y estimates f o r the c u r r e n t  production  t r a i t s were c o n s i d e r a b l y h i g h e r t h a n t h o s e  lactation.  T h i s may  statistical  have a r i s e n  analysis  used  mentioned, s i r e e f f e c t current and  lactation  in  with  were  completed  of b l o o d sampling. the  inflation  lactation. lactation  In important  of  in age  to  this were  two  As  somewhat  that  adjustment  first  y e a r , was may  environmental  T h i s would t e n d t o r e d u c e  the  first  of  the  previously  the a d d i t i v e Also,  not  for  confounded  a number o f y e a r s  S i n c e age,  lactation  aspects  study.  inflated.  over  model, an i n a d e q u a t e  resulting  due  the result  t h e h e r i t a b i l i t i e s were  records year  and  in  lactation.  generally reported in the l i t e r a t u r e  Johnson  in  effect  the  cause  daughters  lactation,  predominantly  correlated,  possible  have  daughters  some  were  Another  have  somewhat i n a d e q u a t e The  distribution. ,  older sires  subsequent  would  distributions  in  variances lactation  preceeding  the  accounted  for  have o c c u r r e d w i t h a  variance  for  first  heritability for  first  traits.  discussing  the  p o i n t must be  heritabilities considered.  For  of  serum  traits  production t r a i t s ,  an the  49 adjustment  model i n v o l v e d e f f e c t s  with  minor e x c e p t i o n , s i g n i f i c a n t . ,  net  one  a l l of t h e e f f e c t s  This  became  important  insignificant  insignificant sums  removed  This  degrees  since  The  had  wasted  when h e r d  34  that  or  degrees  trait  may  Heritability agreement  Rowlands  from  and  this  the  thus  degrees  have been  error  of  freedom terms  then the  associated  not  errors  to  be  Firstly,  an  reduce  the  term  still  the  sire  in  the  became  were  genetic  especially  insignificant,  f o r both  heritability  terms.  If  estimated  underestimated.  study  were i n c l o s e  literature,  albumin  ( h = 0 . 0 8 5 ) , an 2  et a l .  (1975) the  i n view  of e s t i m a t i o n .  of  and  and Rowlands e t a J L .  previous  research  h e r i t a b i l i t y e s t i m a t e d f o r albumin  unsupported  was  estimate i n  i f not magnitude, w i t h Kitchenham  i m p r e s s i o n from low  were  o f o v e r e s t i m a t i n g the  were i n v o l v e d  in this  Bettini  overall  was  line  e s t i m a t e s f o r serum c o n s t i t u e n t s  the s i g n i f i c a n t , study  traits,  terms  underestimating  interaction  o f freedom  heritable  (1976), The  certain  causes. ,  the combined e f f e c t  agreement of s i g n i f i c a n c e ,  that  two  with e s t i m a t e s p r e s e n t i n t h e  significantly  (1974),  i f  were,  significant.  d i d not s i g n i f i c a n t l y  t h e s e t e r m s were i n s i g n i f i c a n t for  model were  However, t h e i n s i g n i f i c a n t  of freedom  environmental variance  important  that  from  term  sguares.  These  variance.  in  that  F o r many serum  adjustment  arose  adjustment of  analysis.  covariables  then t h e e n v i r o n m e n t a l v a r i a n c e would tend  overestimated.  error  i n the  and  was in  p r e v i o u s e s t i m a t e s and  50 Alkaline  phosphatase (0.299).  heritability previously  reported  noted  but  estimate  was  i n  Peterson  that  insignificant,  estimated  This  ( B e t t i n i e t a l . 1975, further  was  the  was  have  and  that  a  higher  agreement  of  than  those  significance  adjustment  this  moderate  1978).  Haldern  interaction  indicating  to  I t  was  term  estimate  may  was be  an  underestimate. The  heritability  (0.325).  high  repeatability  Peterson for this  present estimate Potassium studies  estimate  were  relative  Amylase, corroboration and  small  with from  present estimate adjustment indeed  Two  These  lowlands  (1976)  and  magnitude the  moderate  standard  of  previous  present  one  estimates  but  higher i n d i c a t i n g  standard t h e lower  estimates. heritability  error,  was a r e l i a b l e  one.  o f 0.202, had no  I n view o f  i t s magnitude  i t i s proposed Given  that the  that the interaction  f o r amylase, t h i s  heritability  may  underestimate.  constituents  exhibited  and  The  was i n s i g n i f i c a n t  be an  with  previous studies.  relative  range.  i n agreement  of the higher a  that the  was i n a c o r r e s p o n d i n g  correspondingly  reliability  a  a result  higher than  also  reported  (0.129),  Kitchenham  were s u b s t a n t i a l l y  (197 8)  Waldern  of h e r i t a b i l i t y  Rowlands e t a l . (1974).  was m o d e r a t e t o  c o n s t i t u e n t o f 0.514 i n d i c a t i n g  was h e r i t a b l e  by  errors  and  for creatinine  heritabilities  estimates  were  utilized  in  this  below t h e c r i t i c a l 0.046  and  0.052  study,  BUN and SGOT,  significance f o r BON  and  level. SGOT  51 respectively.  Kitchenham  heritability  and  heritability  for  BON, b u t  removed f r o m  (1974)  this  both  estimate  noted  that the standard  components value  was  done  allow  a  errors of  constituents,  that  c o u l d prove  may  be  animals  study.  heritability  low  completely  SGOT h a s been  I t was f u r t h e r for  will exist.  spite  these  two  w i t h i n which a nonzero  i n a selection  in  a  heritability  Since, f o r  t h i s range o f confidence reached  interesting  high  significant.  1975).  range o f confidence  effective  a  ( B e t t i n i £t a,l. 1975) and a s  (Furtmayr  f o r the population estimate  two  on  i n this  heritable  heritable  being  a non s i g n i f i c a n t  t h e age r a n g e p r e s e n t  significantly  reported reported  estimates  reported  reported as s i g n i f i c a n t l y not  (1976)  B e t t i n i e t a l . (1975)  f o r BON,  Rowlands e t a l .  a n d Rowlands  into  these values  program, BON and  of  their  SGOT  insignificant  heritabilities. The  remaining  heritable.  was  that  present i n d i c a t i n g  failure These  trait.  to f u l l y include  there  fixation The  ;  identify  for  future  were  not  significantly  possible  a l l nongenetic  c a u s e c o u l d have  sources  e f f e c t s which in  marginally  additional  One  genetic variance  the  of  context  significant  environmental  been  variation.  were n o t random b u t of  the  p o s s i b l e c a u s e s h o u l d be c o n s i d e r e d  traits of  studies,  no a d d i t i v e  causes.  o f t h e genotype r e s p o n s i b l e f o r the  environmental  The s e c o n d  those  was  other  were n o t a b l e t o be measured study.  traits  T h i s may have been due t o two s e p a r a t e  possibility  particular  serum  present  seriously  heritability. effects  should  In be  52 quantified heritable  to  determine  in this  Cholesterol previous (Arave  study was  studies  e t al.,1975,  In the  close  to  zero  error,  the  low  Glucose  to  have  a  and  Furtmayr  estimate  would n o t  in a situation  Rowlands  (1976) r e p o r t e d no  derived  frcm  the  present  the be  and  with  this  of c h o l e s t e r o l  was  large relative  standard  in dispute.  heritability  study  some  component  concur  e t a l . , (1975)  such  in  1969,  s i m i l a r t o t h a t of  Bettini  significant  Lasley  heritability  considering  component.  qenetic  (1975) d i d not  study,  marqinally  reported  large  Stufflebeam  be  genetic  constituent  and,  indicated  to  serum  present  was  found  have a s i g n i f i c a n t  Rowlands e t a l . ( 1 9 7 4 ) , (1976)  traits  one  T a y l o r e t a l . 1966). findinq.  if  and  while  Furtmayr  Kitchenham  significnace.  indicated  cholesterol.  The  and  estimate  the h e r i t a b i l i t y  to  be  negligible. Calcium  has  been r e p o r t e d  studies  (Kitchenham  Bettini  e t a l . 1S75).  heritability involved, was  Rowlands  in  not its  be  and,  heritable  1976,  These  f o r calcium  could  clear  and  t o be  i n view  of  indicated the  considered secure. that  three  Rowlands e t a i » /  reports  statement  by  previous 1974  and  a  low  standard This present  calcium  was  errors study  negligibly  heritable. Inorganic present the  p h o s p h a t e was  study.  literature.  calves,  reported  not  This finding  heritable  was  both  supported  Rowlands e t a l . (1974), a  low  on t h e  in  a  t o moderate h e r i t a b i l i t y  b a s i s of and  this  refuted in  study  using  for inorganic  53 phosphate, while Dinkel  (1968) r e p o r t no  Total and  Kitchenham  and Rowlands  Hilscc  and  significance.  p r o t e i n , shown s i g n i f i c a n t l y  Rowlands  (1976) and  (1976),  was  not  h e r i t a b l e by  found  Kitchenham  t o be s o i n t h e p r e s e n t  study. Uric this  acid  study.  was  n o t found  further corroboration  this  study.  being  of b i l i r u b i n  T h i s was t a k e n Furtmayr  Ho  was e s t i m a t e d  as an i n d i c a t i o n  as n e g a t i v e i n  that  heritability heritability reported  a  b u t one w h i c h was l o w enough t o be o f  value.  Heritabilities bicarbonate  T h e r e was no  Bettini etal.,(1975)  heritability  practical  was g i v e n .  (1975) r e p o r t e d b i l i r u b i n  insignificant,,  significant little  of r e l i a b i l i t y  heritability  available.  heritability  was n e g l i g i b l e . as  he.rita.ble i n  B e t t i n i e t a l . (1975) r e p o r t e d a h i g h  (0.47) b u t no i n d i c a t i o n  The  t o be s i g n i f i c a n t l y  of  triglycerides,  sodium,  and t h y r o x i n e were a l l e s t i m a t e d  publications  to  the  contrary  have  as  been  chloride,  insignificant. l o c a t e d by t h i s  author.  A summary among  of  a l l traits  phenotypic is  given  and  i n Table  a g r e e m e n t between t h e r e s u l t s o f t h i s be  given  at  correlations in  this  point.  to d i s c u s s .  the l i t e r a t u r e  environmental  One  V.  No d i s c u s s i o n o f t h e  and p r e v i o u s  reason  correlations  studies  i s t h e l a r g e number o f  Another i s t h a t c o r r e l a t i o n s  a r e seldom  will  o f t h e same s t a t i s t i c a l  reported derivation  T.  1  TRAIT  H=== = ==  1  Phenotypic  I  (below  Hilkl  diagonal)  Fat 1  and e n v i r o n n e n t a l  Protl  (above  HilkC  + " = = = = = = = « " SC=3=ES3SI1ISZIUSZXSS3 3S3ZSZS  Hilkl  .7561.019  diagonal)  Fat C  c o r r e l a t i o n s aaong  ProtC  traits  Ca  »ith  associated  P01  Glue  Ctt 3 e x 3 3S1S s s cs ESS£=£SS*XSS  .8661.021  . 4301.092  . 2 0 9 1 . 100  .3561. 105  .0551.083  -.1131.088  -.1341.082  .7521.017  . 2 9 7 i . 120  .1081.091  . 3 1 2 1 . 124  .0781.086  - . 1901.091  -.0951.088  . 3 5 0 1 . 120  . 2 0 8 1 . 111  . 3 9 1 1 . 120  .0591.094  -.1581.096  -.0791.095  .7691.050  .9121.023  -.0001.101  -.0611.096  - . 1081.098  .7851.015  .1041.092  -.0811.090  -.1471.087  .0451.115  -.0591.102  -.0711.125  I Fat 1  1  .6981.039  I  Protl  1  .8761.016  .7551.032  I H i IkC  I  .4111.061  .2171.078  .3351.077  I Fat C  1  .2781.067  .1591.060  .2921.073  I  ProtC  1  .1091.066  .3111.077  .1281.072  . 9231.012  I  Ca  1  .0501.062  .0691.063  .0521.068  -.0001.066  .0831.063  .0321.071  I  POI  1  -.0971.065  -.1291.066  - . 1381.068  -.0521.066  -.0741.061  -.0521.069  .0081.052  I  Glue  1  -.1221.061  -.0831.064  -.0691.068  -.0981.066  -.1131.062  -.0621.075  .1011.049  -.1021.051  I  BOB  I  .0561.065  .0821.066  .0111.075  . 0611.068  . 1201.065  .0621.077  .0481.052  .1131.054  I  Oric  1  .0831.067  .0371.065  .0421.071  . 0301.067  .0191.062  .0951.070  . 1691.049  . 1841.050  Chol  1  .0161.062  -.0111.061  -.0121.068  . 2751.063  .2201.061  •217t.069  .1331.018  -.0961.051  -.0071.049  T P  I  -.0191.068  -.0141.070  -.0511.071  .0151.061  -.0161.062  .0191.068  .1951.018  -.0561.052  -.0151.050  Alb  I  .0981.065  .0371.069  .0521.076  .0351.072  . 1321.066  . 1011.078  .3611.047  -.0731.056  .0331.053  Bili  1  - . 3621.051  -.0121.061  - . 1091.067  .0001.066  .0111.061  - . 0 1 9 1 . 072  .0161.049  .0631.052  .3011.045  Alk P  I  -.0191.071  -.0721.076  -.0061.082  - . 1951.080  -.2111.074  -.2021.086  .0271.062  .0581.064  .0721.062  SCOT  1  .0191.061  -.0011.061  .0871.066  .0671.070  .0131.068  .0131.073  -.0181.053  .0781.054  .0031.052  Creat  I  -.0931.071  -.1121.077  -.0551.085  -.0231.087  -.  -.1191.095  .0061.063  . 1021.066  .0301.062  Tr iq  I  -.1671.060  -.1331.063  -.1691.067  - . 1021.065  -.0801.063  -.1161.072  .2831.045  .0481.053  .1471.048  Na  1  - . 0391.062  -.  .0021.069  .0551.065  .0581.062  .0161.071  .0671.019  .1341.052  -.0761.051  -.0081.071  .0131.076  .0291.071  .0581.069  .1021.078  .1291.056  .3751.051  -.0321.100  .0101.061  .0131.069  - . 0691.067  -.0631.065  -.0701.072  .0001.052  i  K  .1131.06 8  Cl  .0081.063  0391.061  .7891.029 .8321.023  1321.081  HC03  -.0931.061  -.0381.066  -.0561.069  -.0301.066  -.0161.063  -.0231.072  T4  -.0191.061  .0201.063  .0001.068  -.0111.065  -.0291.063  -.0991.071  Anyl  - . 0 1 9 t . 07<t  .0611.075  .0151.073  -.0131.080  -.0261.083  .0121.079  .0081.065  -.0171.019  . 1031.062 - . 1051.065  -.1991.051 .2121.051  -.0421.052 .1091.050  . 1001.051 -.0311.051  . 1201.043  -.0691.051  . 1631.048  . 1201.058  -.0911.062  .1541.058  T a b l e V<cont.).  P h e n o t y p i c (belov d i a g o n a l ) and e n v i r o n m e n t a l (above d i a g o n a l ) c o r r e l a t i o n s among t r a i t s with a s s o c i a t e d SE.  r  i  TBAIT  BON  ——————34  Oric  Choi  T P  Alb  Bili  Alk P  SGOT  Creat  ==SSSSS3BSS333333333 — — —•— 3— — 3 ISSSS3tK 3S333SCSISS33333 3=3 33 33332333333333 33 33 3S3E3 3333 33  ==============  Hilkl  -.029±.090  -.0991.095  .0181.084  .0221.095  .0381.090  -.3951.071  .1241.114  .02 81.082  -.0031.112  Fat 1  .0341.093  -.0491.094  -.0161.089  .0371.100  -.0221.096  -.0481.088  .0521.117  -.0021.089  .0311.122  Protl  -.0781.111  - . 1611. 110  -.0141.096  . 0591. 107  .0881.110  -. 1231.093  .1911.128  .1021.092  .0951.140  .056*.100  -.0371.104  .3641.096  -.0191.094  -.1401.115  -.0001.102  .1161.159  .0671.105  . 1071.148  c  .123±.092  .0521.090  .2741.088  -.0191.088  .0391.098  .0551.092  .0111.138  -.0011.099  .0061.136  Protc  .0411.119  .1011. 114  .3401.110  .0531. 103  -.0231.126  -.0271.115  .1211.178  .0761.110  .1751.198  Ca  .050±.066  . 1731.062  .1341.060  . 1991.061  .3861.060  .0161.061  .0341.089  -.0511.067  .0081.093  P04  .1191.069  .1891.063  -.0991.065  -.0581.065  -.0801.074  .0651.066  .0611.094  .0861.070  . 1221.099  -.0441.066  .1111.062  -.0071.062  -. 0451.062  .0351.069  .3031.056  .0911.089  .0031.067  .0381.091  .1871.064  . 1891.064  -. 1371.065  -.0111.075  .2001. 063  -.0211.091  .0391.070  -.0511.093  .0311.061  -. 0521.063  -.0761.069  .4321.050  .0121.086  .0091.067  -.0411.090  .0601.063  .2861.065  -.0981.061  -.1221.089  .0431.067  -.0021.093  .0031.070  -.1791.062  -.1151.086  -.0571.067  -.0561.090  -.0371.070  -.0641.095  -.0901.076  .2181.103  -.0111.089  .0071.067  .0001.093  .27«t. 104  .005±.127  flilkC Pat  Glue BOH Oric  . 18U1.050  Choi  .1811.050  .0311.049  T P  -.1301.051  -.0511.050  .0591.050  Alb  .0801.056  -.0691.054  .2651.051  .0031.054  Bili  . 1921.050  .4281.040  -.0971.049  -. 1761.049  -.0341.054  .0111.061'  -.0971.061  -. 1021.061  -.0641.066  -.0091.062  Alk P  -. 0231.061*  SGOT  .0371.054  .0081.052  .0411.052  -. 0541. 053  -.0201.057  .0071.053  .0871.069  creat  -.0661.064  -.0371.061  -.0011.063  -. 0411.061  .1191.070  .0001.063  .1051.079  .0311.067  Trig  -.0031.053  .0661.051  .0901.049  . 1011.050  . 1751.053  .0521.049  .1611.061  .0241.053  -.4411.056  . 1571.054  .0651. 062  .0041.062  .1201.054  .1191.063  .0591.061  •034t.055  -.0841.068  .0341.060  .1051.073  Na  .0041.053  .0531.051  .0371.050  -. 1271.050  K  . 1221.058  .1701. 054  .0331.056  . 0681.056  Cl  -.0091.098  |  -.0921.053  -.0261.052  -.0751.052  -.0651.051  . 1281.054  .0221.052  .0391.062  .0601.054  .0681.061  HC03 I  -.0041.053  .0551.050  -.0101.050  -. 1551. 049  -. 1301.054  ,0791. 049  -.0231.062  -.0041.054  -.0071.063  .0121.052  -.0181.050  .0501.049  -. 0541.050  .2581.050  -.0341. 049  .0861.061  .0041.052  .0261.062  -.0261.062  .0301.061  .0931.058  . 0831.059  . 1461.063  . 0371. 059  -.0091.075  .0291.063  .0631.079  T4  |  Amyl  |  Table V(cont.).  i  Phenotypic  (belov d i a g o n a l )  and e n v i r o n m e n t a l  (above diagonal)  c o r r e l a t i o n s aiong t r a i t s » i t h a s s o c i a t e d  r  TRAIT 1  Trig  Na  33SCSSSE3S2S3  ^ — — — — — — — ——se.— — —  BilJM  1  K  Cl  3ZXX3S  HC03  T4  Amyl  3isss:s3:z3=x XSS3SZXXX333X————————————  ————=——-——-—  -.1861.081  -.0431.084  -.0051.099  . 0091.085  -.1391.087  -.0211.082  .0691.108  Fat 1 I  -.1521.086  -.0451.089  .0041.101  .0121.089  -.0471.091  .0231.036  . 1431. 127  Protl  1  -.1961.095  .0021.097  -.1181.112  .0491.097  -.0641.097  .0001.094  .1291.130  HilkC  1  -.1371.100  .0711.096  -.1161.114  .0261.101  -.0401.100  -.0541.097  -.0131.114  Fat C I  -.1011.091  .0701.088  .0431.098  -.0871.096  -.0201.090  -.0361.090  .0081.106  Protc  I  -.1661.118  .0611.111  .0091.123  . 1091.118  -.0311.115  -.138i.114  .0371.123  Ca  I  .2831.056  .0671.062  .1421.073  - . 0001.066  -.0471.062  .1201.060  . 1391.079  P04  1  .0491.067  .1381.066  .4121.066  - . 2061.065  .2191.064  -.0711.065  - . 1081.086  Glue  1  .1481.060  -.0781.064  -.0401.051  . 1041.065  -.0321.064  . 1641.060  .1781.073  BOH  1  -.0031.068  .0041.068  .1881.077  -.0961.067  -.0041.068  .0131.066  . 1051.087  Oric  1  .0681.064  .0541.064  .1761.070  -.0261.065  .0551.063  -.0191.063  .0501.084  Choi  I  .0901.061  .0371.062  .0361.074  - . 0771.066  -.0101.062  .0501.061  . 1071.078  T P  1  .1041.064  -.1281.062  .0761.074  -.0661.065  -.1571.062  -.0551.063  .0881.079  Alb  1  .1901.068  .1701.070  -.0251.083  . 1391.070  -.1401.070  .2771.064  . 1691.085  Bili  I  .0521.061  .0651.062  .0371.073  .0221.066  .0791.062  - . 0 3 4 1 . 061  .0431.080  Alk P I  .2031.088  .0041.088  .0361.102  . 0411.087  -.0281.088  .1091.089  .123*.115  SGOT  1  .0251.068  .1281.069  -.0601.080  .0641.070  -.0041.069  .0041.067  -.1151.088  Creat  I  -.5761.086  .1551.093  .2451.111  .0741.090  -.0091.093  .0341.092  - . 1351. 131  -.0481.062  .0541.074  -.1381.064  Trig  -.0531.062  .1001. 060  . 1721.084  . 2561.061  .2881.056  .0051.061  -.0421.082  . 0501.074  -.0061.074  - . 0 5 7 1 . 073  -.0211.093  -.3511.057  .0031.065  . 1241.082  -.0421.061  - . 1641.081  Ha  1  -.0481.049  K  I  .0491.056  .2171.054  Cl  1  -.1341.051  .2491.048  .0481.056  HC03  I  -.0531.049  .2881.045  -.0051.056  - . 3411.045  T4  I  .1001.048  .0051.049  -.0521.056  .0021.051  -.0421.049  Aayl  1  . 1421.060  -.0361.060  .0311.066  .1141.059  -.1401.059  .2421.071  .-971.080 .0831. 058.  SE.  57 nor  were t h e d a t a  the  present  adjusted  study.  covariance zero.  lower  were s e t t o  for  considered  the  zero.  of  zero.  greater  than  zero.  of  theoretical  lack  traits  range,  these  the exact  with  genetic  was  t h a t were or  of  a  this  value  no  be  was  slightly  low g e n e t i c  genetic c o r r e l a t i o n s , were s u b j e c t t o  o f which may g r e a t l y e x c e e d  f o r these  to  was  genetic  the correlation s t a t i s t i c .  attached  of  appropriate  only  very  correlations  but l a r g e s t a n d a r d  traits  may f a l l  Cue t o i n the  e r r o r s were i n d i c a t i v e o f correlations  between  two  e x t r e m e l y low g e n e t i c components o f v a r i a n c e .  On a t h e o r e t i c a l no  that there  value  many  o f a parameter t h a t has a  negative  genetic  limits  component  negative,  The same r a t i o n a l e c o u l d  a  genetic  noted that  genetic  was used t o e s t i m a t e  of c r e d i b i l i t y  i s  and  obtaining of a negative  chance, c o r r e l a t i o n s generated acceptable  in  that h e r i t a b i l i t i e s of  was  estimation  indication  If  extreme f l u c t u a t i o n ,  the  It  the  components o f v a r i a n c e  component o f v a r i a n c e estimate  done  c o r r e l a t i o n s were s e t t o  trait  The  of variance.  some g e n e t i c  If  individual  as a s t a t i s t i c a l  component  the  V I .  Table  as a s t a t i s t i c a l  limit  treated  the  in  T h i s was done f o r t h e same r e a s o n  variance  for  given  components o f c o v a r i a n c e  components and many g e n e t i c  some t r a i t s  is  section.  A summary o f g e n e t i c is  as  A discussion of c e r t a i n r e l a t i o n s h i p s w i l l  be u n d e r t a k e n i n a l a t e r  correlation  f o r t h e same e f f e c t s  b a s i s i t c a n be seen t h a t i f a  component,  then  i t cannot  exhibit  trait a  has  genetic  Table  VI.  TRAIT  Genetic  1  ====== U =  correlations  Hilkl = =r = = = = = = = = = =  tlilkl Fat  1  1  I  diagonal)  Protl  and genetic couponents  MilkC  Fat  of covariance  C  ===================================================== 102111822  268512710  .3141.493  110.198.5  9. 67E » l 6 . 8 0 E  (above  ProtC  diagonal)  with  Ca  = = = X = S3S ESEX  associated SE.  P04  G  ==================.  386612206  102612600  0. 0  0.0  0.0  917. ± 1 1 1 0  199.1125.  131. 185.5  0.0  0.0  0.0  2197±1762  153.188.8  139. 1 8 5 . 2  0. 0  0.0  0.0  1.38E1.75B  1.16E1. 8 2 E  0. 0  0. 0  0.0  516. 1290.  0.0  0.0  0. 0  0.0  0. 0  0.0  0.0  0.0  Protl  I  .9411.101  .7781.215  UilkC  I  . 5881.371  . 1 0 7 i . 393  Fat  C  I  . 6 8 0 1 . 130  . 6 5 2 1 . 283  .6131.371  ProtC  1  . 7 0 1 1 . 358  . 1 3 7 1 . 339  .5571.298  .9191.035  .9661.053  Ca  1  0.0  0.0  0.0  0. 0  0.0  0. 0  P04  1  0.0  0. 0  0.0  0.0  0.0  0. 0  0.0  Glue  I  0.0  0. 0  0.0  0.0  0.0  0. 0  0. 0  0.0  . 9411.797  .1911. 652  .7681.561  . 1311.589  .1581.610  . 1581. 448  0.0  0.0  0.0  0.0  1. 7 1 i 1 . 6 7  1 . 8 1 i 1 . 12  O.C  0.0  0. 0  0.0  0.0  0.0  0.0  0. 0  0.0  0.0  0.0  0. 0  0. 0  0.0  0.0  BOS  1  Dric Chcl  0.0  0. 0  0. 0  0.0  0.0  .637i.661  . 4 5 6 t . 583  -.1381.519  .8961.513  .6971.484  . 5 6 0 1 . 393  0.0  0. 0  0.0  0.0  0. 0  0. 0  0.0  0. 0  0. 0  0.0  0.0  Bili  -.9611.505 0.0  Creat  1  0. 0  0.0  Alb  SGOT  . 8391.101  -.7991.601  -. 5 9 7 1 . 6 0 2  P  .3021.389  - . 5 3 7 t . 516  T P  Alk  -.1901.132  0.0  - . 5 7 0 1 . 396  - . 9061.192  0. 0  0.0  - . 6 3 6 1 . 357  -.5161.397  -. 9051.190  -.9111.193  -.8681.203  0.0  0.0  0.0  .1001.508  .1071.532  -. 1 2 1 1 . 6 1 9  0.0  0.0  0.0  -. 2801.297  -.4481.281  -.5911.244  0. 0  .0671.712  0.0  Trig  O.C  0. 0  0.0  0.0  0.0  0. 0  0. 0  0.0  0.0  Na  0.0  0. 0  0.0  0.0  0.0  0.0  0. 0  0.0  0.0  K  . 9 8 3 1 . 530  .8361.597  .6161.391  .1371.150  . 4 4 4 1 . 399  0. 0  0.0  0.0  CI  0.0  0. 0  0.0  0.0  0.0  0.0  0.0  0.0  HC03  0.0  0. 0  0.0  0.0  0.0  0. 0  0. 0  0.0  0.0  TU  0.0  0. 0  0.0  0.0  0.0  0. 0  0.0  0.0  0.0  . 3281.368  .0391.103  . 0 6 9 1 . 432  0. 0  0.0  0.0  Aayl I  Fat  (below  -. 1 9 5 ± . 526  -.078t.525  -. 7 3 2 1 . 368  I  'E'xIO*  -.6531.451  -  f o r covariance  and  SE.  - . 8661.990  00  VI(cont.).  Table  Genetic  c o r r e l a t i o n s (below diaqonal) and q e n e t i c components o f co v a r i a n c e  (above diaqonal) with a s s o c i a t e d  SE.  •—  i BUN  TB AIT  Oric  ssxs  Chol  E SS3333EE3=33  .3^3 33=3:3= = ES  ESSE S3  SSXSXS  Alb  T P  Bili  Alk P  SGOT  Creat  ===========================================================================  Hilkl  na.±73.7  0.0  0.0  -26.3H1.2  13. 9t6.71  0.0  -252211013  0.0  -7.6913. 42  Fat  4.3912.12  1.021.128  0.0  -1. 151.479  .4961.209  0.0  -75.5131.8  0.0  -.4631.199  -.1581.051  0.0  -92.7137.1  0.0  -.3541.150  1  -1.331.569  Protl  7.52±3.16  1.221.503  0.0  HilkC  52.H30.8  0.0  0.0  0.0  47. 0H8.8  0. 0  -615312523  451.1290.  - 1 1 . 7 H . 47  Fat  2.18±1.73  0.0  0.0  0.0  1.291.589  0.0  -220.193.5  17.3t7.31  -.6621.279  -207.188.9  -15.311.69  -.8901.369  C  ProtC  I  3.0H1.56  0.0  0.0  0.0  1.231.539  0. 0  Ca  [  0.0  0.0  0.0  0.0  0.0  0.0  0.0  0.0  0.0  P04  I  0.0  0.0  0.0  0.0  0.0  0. 0  0.0  0.0  .0011.001  Glue  I  0.0  0.0  0.0  0.0  0.0  0. 0  0.0  0.0  0.0  0.0  0.0  0.0  .0611.026  0.0  0.0  0.0  0.0  0. 0  0.0  0.0  0.0  0.0  0.0  0. 0  0.0  0.0  0.0  0.0  0. 0  0.0  0.0  0.0  BOH  -.2981.178  0.0  -.0061.003  Uric  I  0.0  Chol  I  0.0  0.0  T P  1  0.0  0.0  0.0  lib  I  0.0  0.0  0.0  Bili  I  0.0  0.0  0.0  Alk P  I  0. 0  0.0  0.0  -.0841.468  0.0  SGOT  I  0.0  0.0  0.0  0.0  .9191.955  0.0  -1.011.660  Creat  |  -.2111.598  0.0  0.0  0.0  -.2601.116  0.0  .3151.299  .2661.510  Triq  I  0.0  0.0  0.0  0.0  0.0  0.0  0.0  0.0  0.0  Na  |  0.0  0.0  0.0  0.0  0.0  0. 0  0.0  0.0  0.0  K  |  0.0  0.0  0.0  .7571.546  0.0  CI  I  0.0  0.0  0.0  0.0  0.0  0.0  0.0  0.0  0.0  HC03  I  0.0  0.0  0.0  0.0  0.0  0.0  0.0  0.0  0.0  T4  I  0.0  0. 0  0.0  0.0  0.0  0.0  0.0  0.0  0.0  Auyl  I  0.0  0.0  0.0  .0081.516  0. 0  1.16H.03  .5941.283  1.45H.09 0.0 -.0U81.585  -. 7011.998  -1.29t. 944  0.0  -.0721.047 0.0  0.0  .4181.117 0.0 -65.3122.1  -.6571.391  -.4161.346  -.0011.000 0.0 .2081.087 .0951.039  1. 101.793  -.4011.400  U3  Table  ?I(cont.). Genetic SE.  i  r  I  TBAIT I  I  Fat 1  (belov diagonal)  and q e n e t i c components o f c o v a r i a n c e  (above d i a q o n a l ) with a s s o c i a t e d  •  Trig  H*"====4 1 Hilkl |  correlations  Ha  Cl  HC03  S BXSSSXSZSSSC3SS X2S3CZXXEEZSnm s x z s xs x e s xzxzzss xcxt  0.0  0.0  | 0.0  T4 — ——  —  Anyl  |  mi  34.0*15.0  0.0  0.0  0.0  -4426*1770 I  0.0  -.130*.055  0.0  0.0  0.0  -354.*142. I  0.0  0.0  0.0  -252.±104. |  0.0  0.0  6438*2710 |  •  1 Protl |  0.0  0.0  1.07*.l»36  1 BilkC |  0.0  0.0  07.8*18.5  I  0.0  0. 0  .367*.191  0.0  0.0  0.0  27.8*13.4 |  J Protc |  0.0  0.0  1.24*.565  0.0  I  Fat C |  -234.496.6  0.0  0.0  41.9*24.5 I  Ca  |  0.0  0.0  0.0  0.0  0.0  0.0  0.0  I  I POtt  |  0.0  0.0  0.0  0.0  0.0  0.0  Q.O  |  I Glue  |  0.0  0.0  0.0  0.0  0.0  0.0  0.0  I  1 BOH  |  0.0  0.0  0.0  0.0  0.0  1 Uric  |  0.0  0. 0  0.0  0.0  0.0  0.0  0.0  I  1 Choi  |  0.0  0.0  0.0  0.0  0.0  0.0  0.0  I  1 T P  1  0.0  0. 0  0.0  0.0  0.0  0.0  0.0  1  1 Alb 1  0.0  0.0  .007*.003  0.0  0.0  0.0  .023*.225 I  1 Bili  0.0  0.0  0.0  0.0  0.0  0.0  0.0  1 Alk P |  0.0  0.0  -.715*.292  0.0  0.0  0.0  1 SGOT |  0.0  0.0  .723*.281  0.0  0.0  0.0  234.±89.5 |  I Creat |  0.0  0.0  -.002*.001  0.0  0.0  0.0  1.20*.466 |  0.0  0.0  0.0  0.0  0.0  |  0.0  0.0  0.0  0.0  0.0  1  0.0  0.0  0.0  1.21*. 487 |  0.0  0.0  0.0  1  0.0  0.0  I  0.0  I  |  I Trig  I  1 Ha  |  0. 0 0.0  -.043*.012  0.0  0. 0  |  0.0  0.0  0.0  I HC03 |  0.0  0. 0  0.0  0.0  I  0.0  0. 0  0.0  0.0  0.0  1 Aiyl  |  0.0  0. 0  .317*.453  0.0  0. 0  1  X.  1 K 1 Cl  |T<*  1  -22.6*8.60 I  |  -150.*56.4 (  0. 0 i  61 correlation trait  with another t r a i t ,  has a g e n e t i c  regardless  component.  F o r t h i s r e a s o n , when t h e above  situation  occurred  o r was i n d i c a t e d  standard  errors,  the  genetic  correlation  number o f p a i r e d  o f whether t h e s e c o n d  genetic  by e x t r e m e c o r r e l a t i o n s  component o f c o v a r i a n c e a n d t h e  were s e t t o z e r o .  The  appendix  lists  selection f o r the additive  and  i t  breeding  is  traits  To  direction.  indicated  that  another based  that  the  Also,  farmer w i l l  find  range  of  factor  In  on  effect, a  better  selection  t h e framework o f d i r e c t s e l e c t i o n , t h e  of  the  a portion  regards  the genetic a  primarily  component.  the  that  In general,  s e l e c t i o n on  population  mean  context o f t h i s present o f the genotype  effect  of  the  of  level genes  the  that  i n the study, i t cow  of that were  a  was  trait,  a  major  i n t h i s genotype.  approach  result i n  classical  conventional  component i n d i c a t e s  additive  with  significant will  the dairy  f o r e s t a b l i s h i n g t h e measured  controlling  is  gene  the  work w i t h i n  genetic  responsible and  of the t o t a l  c a n r e s u l t i n movement  desired also  that  portion  must have a s i g n i f i c a n t g e n e t i c  significant trait  unlikely  program w i t h i n  theory.  the  o b s e r v a t i o n s used f o r each c o r r e l a t i o n .  G e n e t i c improvement i n d a i r y c a t t l e h a s p r o c e e d e d by  and  i s that premise  to  conventional  selection  of indirect selection. that  the  traits  This  involved  theory, approach have  c o r r e l a t i o n and t h u s s e l e c t i o n f o r one concurrent  interpretation  change of  this  in  another  situation  trait. i s  that  a  trait The the  62 individual  genotypes i n v o l v e d  with the t r a i t s  in  common, t h e r e l a t i v e s i z e o f t h i s p o r t i o n  the  genetic  traits  correlation.  This  significant  of genetic  genetic  correlation  have  basis,  to  albumin,  SGOT,  discussion generally  and  study  BON  there  follows  that  amylase,  only  On  component.  these  be were i t this  discussion  is  These  potassium,  and a l l m i l k p r o d u c t i o n t r a i t s . , correlations  must  If a trait  further  phosphatase,  the estimation also  genetic  examined  of h e r i t a b i l i t y some  production  correlations  In  further  traits  but  correlations  a word o f c a u t i o n  warranted.  The g e n e t i c  analyses  involved  i n the c a l c u l a t i o n . by  relationships traits.  i s given  will  traits,  involving  these  1 summary o f p h e n o t y p i c traits  and  the  i n Table VII.  were o f g r e a t e s t  interest i n  i n i n t e r p r e t i n g these  correlations  halfsib  multiplied  and  o f t h e serum  between t h e p r o d u c t i o n  serum c o n s t i t u e n t s  The g e n e t i c  also  i t  a s i g n i f i c a n t genetic  alkaline  and t h e m i l k  study  that  correlations.  meriting  two  be c o n s i d e r e d .  heritable  is  with  then  genetic  traits  of the v a r i o u s  Beyond  traits  of  those  creatinine,  this  significant  t h e number  reduced are  any  component,  the  by  o f gene e f f e c t . ,  components f o r b o t h t r a i t s .  shown t o have no g e n e t i c cannot  is  portion  being estimated  common g e n o t y p e a f f e c t s  t h r o u g h t h e phenomenon o f p l e i o t r o p y  The i d e a  have some  were  correlations  calculated  m u l t i p l i c a t i o n by a c o n s t a n t Since errors  of  t h i s f a c t o r , the standard  this  of four  measurement error  using was were  associated  Table  VII. Genetic (above) production traits. , —  T  1  I Trait  |  h*  and  phenotypic  (below)  correlations  between  selected  serum c o n s t i t u e n t s  and t h e  r  |  Creat  Alk  P  Atayl  K  Alb  SGOT  BON  |  1-=======+ | .10741.0976 |  -.4901.432 -.0934.074  -.9641.505 -.0491.074  -.4951.526 -.0191.074  .9831.530 .1181.068  .6371.661 .0981.065  0. 0 .0191.061  .9441.797 .0561.065  | |  1 F a t 1 | .1530±.1O78 |  -.636±.357 -.1121.077  -.5701.396 -.0721.076  -.7321.368 -.0431.080  -.0781.525 -.0081.071  .4561.583 .0371.069  Q.O -.0011.064  .4941.652 .0821.066  | |  I P r o t l | .15541.1172 |  -.546l.397 -.0551.085  -.9C6t.492 -.0061.082  -.6531.451 -.0261.083  .8361.597 .0131.076  -. 1381. 549 .0521.076  0. 0 .0871.066  .7681.564 .0411.075  | |  I Milkc  | .3370±.1382 |  -.2801.297 -.0231.087  -.905l.190 -.1951.080  .328l.368 .0641.075  .6161.394 .0291.074  .896l.513 .0351. 072  .1001.508 .0671.070  .1341.589 .0611.068  I |  I F a t C | .28211.1270 |  -.4481.281 -.1321.081  -.911t.193 -.2441.074  .0391.403 .0151.073  .1371.4 50 .0581.069  .6971.484 .1321.066  .1071.532 .0131.068  .1581.610 .1201.065  I 1  I P r o t C | ,3979t.1560 |  -.5911.244 -.1191.095  -.8681.203 -.2021.086  .C691.432 .0421.079  .4441.399  . 1021. 078  .5601.393 .1011.078  -. 1211.619 .0431.073  .1581.448 .0621.077  | |  1  flilkl  64  with  the genetic c o r r e l a t i o n  standard  own s a m p l i n g  with  reliability  and  of these  a  albumin,  correlations  Current  lactation with  a  lactation  silk,  taken  the  much  (0.944). in  their  was between  standard  lower,  error.  Unlike  /  error among  Although that  indicating  a  points  estimates  o r BUN.  estimates  in  time  f o r creatinine  as with  first  (r=-C.905)  with  milk,  current  there  was no  These d i s c r e p a n c i e s  o f t h e same  were  subset  (0.616) and a l b u m i n  correlations  lactation  with c r e a t i n i n e  a smaller  relationship  Potassium  reliable first  with  phosphatase,  strongest  are e s s e n t i a l l y  different  correlation  reflected  above was f o r a l b u m i n .  Alkaline  strong,  relationship  at  was  was c o r r e l a t e d  the  standard  milk.  what  (0.637) and BUN  The p o o r e s t c o r r e l a t i o n  e r r o r was  milk  showed  small  between  phosphatase  estimate.  (0.896) a l s o had  strong  genetic  between c r e a t i n i n e and m i l k was l e s s t h a n  constituents.  lactation  they  i s best  strong  alkaline  with  estimate.  the standard  more r e l i a b l e  albumin  phosphatase,  reliable  correlation  possessed  The s t r o n g e s t g e n e t i c c o r r e l a t i o n  alkaline  serum  since  error.  (-0.490),  (0.983),  set of c o n s t i t u e n t s l i s t e d  the  correlations  The  e a c h one s u b j e c t t o i t s  production  creatinine  errors.  indicating  of  milk  potassium  standard  for  t o i t s standard  lactation  (-0,964),  milk  several statistics,  inflated.  v a r i a n c e . , The m a g n i t u d e o f t h e c o r r e l a t i o n  correlations  the  from  in relation  First  The  appropriately  e r r o r s must be l a r g e f o r t h e s e  were e s t i m a t e d  judged  was  not  correlation  serious.  The  and BUN were a s s o c i a t e d  65 with  large  significant  standard only  for  errors. the  first  the  correlation  importance  the  correlation  were d i f f e r e n t  Literature  was  serum c c n s t i t i u e n t s (1975) milk  study.  correlations estimated  t h i s study  insignificant  First  in  pattern albumin  the  fat  that  appeared  Kitchenham  with  a positive this  e ta l . between  this  present  correlation  present  study.  and  discussed  production  (-0,636),  (-0.732). except  twoe s t i m a t e s o f  a relationship  constituents  were not  that  milk  with Other  production  o r were n o t e d  t o be  literature,  lactation  creatinine  amylase  serum  indicated  of relationships of  i n agreement  by  appeared  by c h a n c e .  reported  reported  they  may h a v e  production.  (1973)  supported  between  in  merely  milk  (1975)  not  lactation  that  i n reporting  and albumin,  SGOT, a f i n d i n g  with  with  Furtmayr  fact  was m a r g i n a l a n d the  sparse  a n dPayne e t a l .  production  The  alkaline  Current amylase as  was g e n e t i c a l l y phosphatase  f a t production was n o t  highly  strongly  correlated  correlated  (-0.570)  followed  and  a similar  correlated.  (0.697) f o r  Also,  current  fat  only. First creatinine  lactation (-0.546),  protein alkaline  (-0.653), p o t a s s i u m  (0.836)  was  a smaller  correlated  consisted  with  of creatinine  yield  was  phosphatase  a n d BON (0.768) .  correlated  (-0.906), Current  group o f constituents.  (-0.591),  alkaline  phosphatase  with amylase  production This  group  (-0.868),  66 potassium  (0.444) and  albumin  (0.560).  T h e r e i s some d i s c u s s i o n w a r r a n t e d of  the  two  groups  lactation  production  Current  lactation  of  correlations,  and  those  records  i n d i c a t i v e of the  population  bleeding.  association  traits  This  and  serum  relationships  on  traits  those  involving  represented as  i t  was  at  This  change,  environmental  and  phenotypic  would be e x p e c t e d caused  be  same  On  serum t r a i t s  genetic  may  this,  incomplete  for  covariance  adjustment  lactation  correlations  age  sire  effects.  a  t o be  be  by  of  present Genetic such  the  and  only  in  correlations  environmentally  comparison of  first  c o r r e l a t i o n s were s u b j e c t  were  discussed first  with  lactation  to  respect to correlations  of  underlying  the  following  reasons.  First  estimates  were  conservative  since  was for  may  serum  overestimated.  more r e l i a b l e i n d i c a t o r s  correlation  environmental  and  be  tend  correlations.  balance, then,  relationships  lactation  as  lactation  events.  unaffected  considerations  of  of current p h y s i o l o g i c a l  should  Beyond  time  these  versus current l a c t a t i o n  heritabilities. with  to  relationships.  lactation the  however,  were  that  correlations w i l l  overestimation,  that  the  in  first  lactation.  between c u r r e n t  important  merits  involving  estimates  T h u s , i f c u r r e n t f e e d i n g programs were a c a u s e production  relative  current  existed  i n time  were r e f l e c t i o n s  the  possibly  overestimated  year of c a l v i n g .  Secondly,  have been i n f l a t e d due  Both s e t s o f  due  correlation  to  to  the  current  confounded  estimates  were  67 used i n t h e  assessment  of  production  and  traits,  involving  first  serum  the  lactation  genetic  relationships  with  reservation that  the  production  were  between those  probably  more  realistic, Futhermore, between  the  obtained  in this  discussed  two  useful  study.  g r o u p s on  The  trends  s i m i l a r f o r both  arose  but  production discerned  by  discernable  of the  estimates  of the c o r r e l a t i o n s trait  none were so s e r i o u s o r  and  discussing  Creatinine highest  correlated  with  correlated  the  patterned  or c u r r e n t  already  groups.  Some as  to  l a c t a t i o n s as  the  and  serum t r a i t s  heritability,  were  the  traits.  production  highly negatively  with  current  particularly  strong  relationship  current  shown t o be  Potassium production  and  correlated  was  by  trait  a pattern  positively  These  with  for milk  two  a  may  the be  heavily  serum  traits  most  highly  constituents  traits  milk.  between  traits.  the  traits  a l l production  the r e l a t i v e l y  another  retained  the  lactation  production  reliable  involving  t h a t w e r e most  a l k a l i n e phosphatase,  with  the  relationships  s e l e c t e d serum c o n s t i t u e n t s  creatinine  It  basis  production  p i c t u r e of the  traits  involved.  and  the  were  standard,  & more g e n e r a l  the  distinctions  in declaring either f i r s t  preferred  with  clearcut  production  were  discrepancies be  no  There  alkaline  except was  a  phosphatase  T h e s e r e l a t i o n s h i p s were low  that  standard was  errors.  involved  both p r o d u c t i o n and  with  milk  estiaates.  protein production  but  68 did  n o t show a s t r o n g r e l a t i o n s h i p The  remaining  strong,  consistent  constituents. measures o f protein,  milk  the  Amylase and BON only  in  serum  correlations  Albumin  was  milk f a t .  constituents with  positively  d i d n o t show  the  production  correlated  production  but,  with  correlations  were  not  e s t i m a t e s and t h u s  tut  selected  with  regards  with  . to  significant  fat for  were c o n s i d e r e d a s n o t p a r t i c u l a r i t y  showed s t r o n g c o r r e l a t i o n s w i t h the  first  lactation.SGOT  and both  strong.  some m i l k  showed  both  traits  no  strong  correlations.  T h i s study  has f o c u s e d  have a s i g n i f i c a n t  on t h o s e t r a i t s  g e n e t i c component.  states  that  genetic  v a r i a n c e and t h e h e r i t a b i l i t y  genetic result with that with  progress  component i n both  a  low  posessed  due  of  heritability  the  to the phenotypic  trait.  theory  A  small  variability T h u s , any  were n o t deemed a s i m p o r t a n t  a significant  selection  a t an a c c e p t a b l e r a t e .  able  more  will  traits as  a d d i t i v e component s i n c e t h e  would n o t be e x p e c t e d  precisely  breeding  to  i s p r o p o r t i o n a l to the  parameter e s t i m a t e s b e i n g low.  low h e r i t a b i l i t i e s  to  Animal  to selection  relative  w h i c h were j u d g e d  those traits  t o progress  under  I n f u t u r e s t u d i e s , i f one were  i d e n t i f y t h i s environmental  component,  the r e l a t i v e i m p o r t a n c e o f t h e a d d i t i v e v a r i a n c e would i n c r e a s e . This  would  usefulness traits.  result  of u r i c  in  acid  a  more  accurate  assessment  and i t s g e n e t i c r e l a t i o n s h i p s  of  with  the other  69 Indirect selection to  selection  of a t r a i t  direct  selection  and  indirectly,  is  alternative  proportional  selection  depends  an  i n modern a n i m a l b r e e d i n g  selection  heritability  offers  on  the  the genetic  to  variance  theory.  of  the  of the t r a i t  selection  success  compared  to  magnitude  of  measurement rational  direct  selection  these  may  factors.  decision  must  will  illumination  Table among  VIII  those  heritable.  standard  considerations  f o r making  this  traits  the  constituents in  this  of  low  its  between  judged  to  table  relative  corroboration  within  A  practical  use. traits  be  correlations significantly  correlations  cf  associated  magnitude,  indicating  some  correlations  involving  the  t h e r e were n o t two  available.  relative  t h e serum  are  milk production correlation  selection  decision.  Unlike the  traits,  the  the strength of the  magnitude t o be o f i n t e r e s t and w h i c h have errors  and  selection.  p r e s e n t s t h e g e n e t i c and p h e n o t y p i c  serum  used  e a s e and economy o f  trait  of  degree of r e l i a b i l i t y .  obtained.  trait  on  be r e a c h e d between  Included  sufficient  in  variance,  indirect  In practice,  o f the relationships  provide support  of depend  become a c o n s i d e r a t i o n  g e n e t i c s y s t e m and t h e Some  will  Response  i n which change i s  between t h e two  The  direct  Response t o i n d i r e c t  sought, the g e n e t i c c o r r e l a t i o n differential.  the  additive  differential., heritability  to  Therefore, this  present  most  heritable  e s t i m a t e s o f t h e same  there study  could of  the  be  no  estimates  ;  Creatinine,  the  of  the  serum  traits.  Table  VIII. Genetic constituents.  (above  diaqonal)  and  phenotypic  (below  diaqonal)  correlations  amonq  selected  serun  1  I Trait 1  i  h*  I  Creat  1 C r e a t I .32481.1328  Alk P  .3451.299  Amyl  K  Alb  .5941.283  -.4011.400  -.2601. 416  -.4461.346  -.6571.394  .3171.453  SGOT  BUN  |  .2661.510  -.2141.598  I  -.0841.468  -1.01t.660  -.0481.585  |  .0081.516  1.46l1.03  -1.291.944  |  .7571.546  1. 101.793  -.7011.998  |  .9191.955  1.45*1.09  I  0.0  I  I A l k P I .29941.1283 I  .1051.079  I Amyl  I . 2024t. 1085 I  .0631.079  -.0091.075  1 K  I .12881.0921 I  .1051.073  -.0841.068  .0311.066  I Alb  I .08521.0810 1  .1191.070  - .0641 .066  .1461.063  .0591.061  I SGOT  I .05221.0739 1  .0311.067  .C871.069  .0291.063  .0341.060  -.0201.057  I BOH  I .04621.0718 1  -.0661.064  -.0231.064  -.0261.062  .1221.058  .0801.056  .0371.054  71 exhibited  few  strong  genetic  constituents.  With  h e r i t a b l e blood  constituents,  with  phosphatase  did  alkaline not  indicate  defined. stronger error.  is  was  genetic  creatinine  degree of  part  s e l e c t i o n would be  as  a m y l a s e and  moderately  was  of  common  related  of  serum  with  standard  with  the  positively  correlation  indicated  error  was  well  amylase  by  the  was  standard  a l k a l i n e p h o s p h a t a s e have with  h e r i t a b l e and  that  other  correlations  creatinine  more r e l i a b l e  in  with  (0.345), a l t h o u g h the  this  I t appeared t h a t  Amylase  to  correlation  (0.594) and  genotype t h a t  of  regards  that  The  correlations  that  the  direct  of  a  creatinine.  most e f f i c i e n t  selection  for  method amylase  level. SGOT, a  trait  statistically with  the  pointed  which  insignificant,  heritable out  that  f o r SGOT  Indirect  selection  located  with  SGOT.  Several  serum  little  selection  larger  with  of  responsible  for  the  less  strong  for  be  This the  an  two  previously under  low  strong  direct  (0.052).  improvement i f t r a i t s  (0.20 2)  large  of  traits.  SGOT  was  be  correlations  was  expected  to  could  correlations  be  with  possessed these c h a r a c t e r i s t i c s .  basis  degree  be  judged  strong It  heritability  heritable  when j u d g e d on high  its  traits  1.46.  a  a number of  h e r i t a b i l i t i e s and  serum  indicated  had  heritability  constituents.  would  moderately  reliable  a  progress could  since  Amylase was SGOT  had  with  of  and  had  correlation i t s standard  similarity Correlations alkaline  in  a  correlation  was  reasonably  error. the  were o n l y  phosphatase  This  genotypes slightly (-1.01),  72 potassium success  (-1.10) and a l b u m i n could  selecting  be  expected  f o r any o f these  BUN  was  significantly  in  much  has  low,  correlation  of  heritability, apparent  in  the  same  traits.  significant 1.45  with  should  involving  SGOT  correlation errors to  and  were s m a l l  traits  caused  large  Amylase* with  level  BUN  that  some  That  of several  t o some  degree.  they  was  factors.  a  genetic  which had a m o d e r a t e of  -1.29.,  of  BUN  It  was  successful  would  of  these  be  to  correlations  However,  the  and  errors  involved  component. in  the  components, t h i s s m a l l i n the estimates.  e r r o r s , these c o r r e l a t i o n s  genotypes o f the t r a i t s  values  One was t h e f a c t t h a t  errors  wide f l u c t u a t i o n s  albumin,  standard estimates for  exceeded t h e t h e o r e t i c a l l i m i t  genetic  genetic  correlations  and  of c o n f i d e n c e i n the high  and c o v a r i a n c e  Not  exceeded t h e t h e o r e t i c a l range of t h e  a small  Other  traits  SGOT.  o f t h i s s t u d y , t h e most  had  standard  as  genetic  BUN  involved  rounding variance  situation  enough, r e l a t i v e t o t h e c o r r e l a t i o n  i n d i c a t e a degree  reflection  o f SGOT by  level  of  o f amylase.  noted  statistic  correlations.  the  be  the  heritability  BUN.  on t h e b a s i s  f o r a lower l e v e l  degree  One o f t h e s e  had a c o r r e l a t i o n  that,  It  moving  reasonable  constituents.,  method o f a l t e r i n g t h e p o p u l a t i o n select  A  h e r i t a b l e , i t possessed  with c e r t a i n h e r i t a b l e which  (0.919).  involved  were h i g h l y  correlations existed  estimation  genetic  were t a k e n  was a  one o f t h e  Combined  Due t o  some  with of  component  the  saaller  to indicate  that  similar.  among serum  constituents  73 and  seemed r e l i a b l e on  were  not  Alkaline  as  the  large  in  phosphatase  was  (-0.446)  and  this  basis  of  magnitude  e s t i m a t e was  were  of  limited  heritable  large  to  this  dairy  these  therefore  use  of  pointed  out  the  direct  in  could the  selection  s m a l l numbers o f rather  and  involved  should  be the  prove  As  an  genetic  on  very  these  traits  to the  frcm  potassium (1.100)  phenotypic  If  a  but  a  correlated  in a trial  merely t o  but  (0.034)  s i m i l a r genotype  were n o t e d due  and  correlation  traits.  be  background  results  SGOT  and  serum  geneticists,  as  of  in  relationships  interest to  example,  animals, i t could t h a n due  of  interesting  a  estimated  genetic  interpretation  indicated  two  correlations  elements f o u n d i n the  being  (-0.060)  environmental input these  the  complex  positive  environmental This  the  Besides  higher  correlations. .  of  compounds  exhibited  anomaly  relationships  i n d i r e c t s e l e c t i o n . <•  discussion  experiments.  in  correlated  not s u f f i c i e n t l y  physiological  response  F i n a l l y , potassium  (-0,657)  was  relationships  large  potassium  among t h e s e t h r e e c o n s t i t u e n t s  relationships  small  Alkaline  magnitude of  cattle.  a  amylase  The  u n d e r l y i n g some o f of  mentioned.  with  three  but  levels.  preceeding  study  with  (0,757),, These l a s t  and  propose the  The  correlated  negatively  i n a l t e r i n g population  correlations  those a l r e a d y  usefulness,,, A l l three r e l a t i o n s h i p s  traits  successful  errors,  relatively reliable,  with a r e a s o n a b l e s t a n d a r d e r r o r . , with albumin  as  negatively  phosphatase a l s o c o r r e l a t e d  positively  t h e i r standard  a  involving sampling  t r e a t m e n t s p r e s e n t i n the  trial.  74 The  same r a t i o n a l e can  estimated results by  in  this  of t h i s  study  forewarning  them  their  present  be  extended  study. would of  experiments.  A  to  a l l  other  preliminary investigation  benefit researchers the  correlations  i n other  of  the  fields  g e n e t i c consequences r e l e v a n t t o  75 CCNCLUSICN  This milk for  study has estimated  production using  traits  these  the  heritabilities  that to  the genetic  traits  parameters estimated  shown  the level  was  breeding  This  would a l l o w  of expression  the  probable  that  these  in  represented  Among  a  t h i s s t u d y was i n p a r t  serum c o n s t i t u e n t s  breeder  of  a small  The  production  of milk,  moderate h e r i t a b i l i t y , serum  traits  creatinine, albumin.  Since  SGOT  and  t o be  reported  progress could these  exhibited  BOH  exhibited  of  possible  in  the  be a c h i e v e d  serum c o n s t i t u e n t s .  of the  in  dairy  ;  exhibited  unexpected.  t h i s degree of h e r i t a b i l i t y  phosphatase,  amylase* low  literature.  potassium  heritability  importance  on  This  the  The were and  b u t were basis  indicated  by d i r e c t s e l e c t i o n on t h e The h e r i t a b i l i t i e s  i t  parameters o f  protein  was n o t  this  genetic  portion  studies,  f a t and m i l k  a finding that  alkaline  hypothesized findings  that  milk  f o r later  to were  in  the  traits.  a report of genetic  as a reference  moderate  trait  terms  and  suggested  possess  animal  of  only  was  r e l a t i o n s h i p s o f i n t e r e s t i n v o l v i n g serum c o n s t i t u e n t s cattle,  the  i f s u c h a change  r e l a t i o n s h i p s among t h e serum and p r o d u c t i o n is  It  should  of a t r a i t  evaluated  and  are necessary  program.  The d e s i r a b i l i t y  further  serum  components o f v a r i a n c e  interest, a trait  t o be d e s i r a b l e .  study  a  of the i n d i v i d u a l t r a i t s .  heritability.,  change  in  f o r some  parameters t h a t  were t h e g e n e t i c  t o be o f g r e a t e s t  high  and e v a l u a t e d  level  estimated  of that of  i n this  76 study  were r e p o r t e d  involving  as  reference  the additive genetic  As an e v a l u a t i o n o f t h e traits  and  the  production  relationships traits,  and  magnitude  of the associated stancard  of  correlation  estimates  were  lactation  and  estimates  one  first  were c o n s i d e r e d Both  were used  parameter  i n  the  of  production  traits  The between level  serum  and  f o r both  possessed  a  production  study,  as separate This  and  production  high,  traits  positive  two  current lactation  of  turn serum  lactation the  same  allowed  the  traits  and  reliablity.  reliable  and c u r r e n t  the  estimates  to current  between  genetic  correlations  were a s f o l l o w s . ,  c o r r e l a t e d h i g h l y and first  to  for  First  i n  serum  traits,  one  estimates  were o f g r e a t e s t  consistent  the  regards  lactation.  relationships  that  of potassium  production also  those  most  this  discussion.  traits,  e r r o r s , ,-• F o r t h e  somewhat s u p e r i o r  estimates.  discerning  i n  studies  c o r r e l a t i o n s were  with  i n v o l v i n g the milk  for  future  between  genetic  particularly  available  for  components of these  estimated  genetic  discussed  values  positively  lactations.  Serum  with  milk  Albumin  level  correlation  with  milk  production. The s t r o n g e s t and of  c r e a t i n i n e with alkaline  negative  traits.  the production  phosphatase  and  correlations  relationships involved  involved  with the  phosphatase  t r a i t s . , Genetic c o r r e l a t i o n s  a l l three highest  o f t h e h e r i t a b l e serum  alkaline  production magnitude  traits  with  traits  were  of  any of t h e  the  production  C r e a t i n i n e was n e g a t i v e l y c o r r e l a t e d w i t h  milk  pictein  77 and  oilk  fat.  production  was  Correlation not  e f f e c t i v e l y reduced  constituents  into  component a n d / o r has traits.  This  existing  a  subset  exhibited  s t u d y has  heritabilities  creatinine  level  and  the  that  original has  r e l a t i o n s h i p s with  the  i d e n t i f i e d such a subset  genetic  relationships, for  serum c o n s t i t u e n t s .  One  was  in  changing  to assess the  S e c o n d , and an  the  correlations the  benefit  This  of  most i m p o r t a n t  as  of  future  strong  the  genetic  to  be  of  use  alkaline  method o f  negative  phosphatase  h e r i t a b i l i t i e s of should  be of  correlation albumin.,  and  in  w i t h a m y l a s e and BON  had  low  r e s u l t i n an  estimated of  existed  f o r two  among  reasons.  indirect selection nonheritable  trait.  correlation  as  These  use  ether  to  researchers  SGOT e x h i b i t e d with amylase,  possessed  the  serum  exhibited  a  positive  heritability,  potassium  the  these  i n d i r e c t c h a n g e i n BON  Since highest  two  traits  of  SGOT, a  level strong  negative  correlation  selection  were  several  phosphatase..  s e l e c t i o n on  a strong  in  Genetic c o r r e l a t i o n s  alkaline  altering BON  and  genotypic r e l a t i o n s h i p s , ,  amylase  heritability.  Since  should  with  production  studies.  genetic  assessment.  these t r a i t s ,  successful  low  the  correlations - positive  a l b u m i n , and  level  correlated,  in interpreting their results.  a g a i n t a k e n as  trait  was  utilizing  genetic  indications  investigated  of  a  underlying  genotypic r e l a t i o n s could  and  was  posibility  level  assessment o f the  fields  milk  nineteen  shown a  O t h e r r e l a t i o n s h i p s examined were t h o s e t h a t the  with  strong.  These f i n d i n g s serum  of  on  level.  with  amylase  78 G e n e t i c c o r r e l a t i o n s among the not  as  specifically  indicative  of  exhibited  moderate  underlying  p h o s p h a t a s e and  moderate  potassium.  Finally,  the the  serum  has  traits  heritability  o t h e r serum and as  in  studies  future  correlations  negative  but  positive  with  sere were  Creatinine  with  alkaline  correlation  a moderate,  alkaline phosphatase  amylase  correlation  and  existed  albumin.  been s u c c e s s f u l  the  trait  production  tabulated  selection  relationships.  additionally,  measured t h a t of  indirect  positive  between p o t a s s i u m and study  in  genetic  amylase.  exhibited  This  useful  h e r i t a b l e serum t r a i t s  a reference  were o f  and  traits.  s o u r c e as  involving  in identifying a  these  i n t e r e s t on  i t s genetic These well  as  traits.  subset the  basis  correlations  genetic  parameters  a source of  of of  with are  guidance  79  Arave,  C.  Bettini,  Bondar,  T. H., D. M a t a s s i n o , F. C o n s e n t i n o , C. I a n n e l l i , P. Hasina a n d fl. 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S c i . , 27:1092-1096. ,  Appendix.  —  Number o f o b s e r v a t i o n s  1  f o r each  pair  of traits.  —  TRAIT |  llilkl  Fat 1  Protl  Milk  Fat C  ProtC  Ca  P01  Glue  - = « = = = + ================================================ (lilkl | Fat  1  I  450  Protl |  101  101  HilkC I  U26  126  38 2  Fat  C|  126  126  382  512  ProtC I  388  388  38 2  459  459  Ca  |  139  139  395  501  501  452  P01  |  124  121  386  187  487  441  511  Glue  I  440  140  395  502  502  452  533  516  BUN  |  440  140  395  502  502  452  533  515  534  Uric  I  111  111  396  503  503  453  531  516  535  Chol  I  138  438  391  500  50 0  451  531  513  532  T P  I  410  110  396  502  50 2  453  533  515  534  Alb  |  440  140  395  502  502  452  533  516  535  Bili  I  137  137  393  199  499  450  530  513  531  Alk P |  439  139  391  500  500  450  531  513  532  SGOT  |  431  131  391  196  496  418  527  511  528  Creat 1  445  115  398  506  506  454  52 9  511  530  Triq  I  111  144  396  501  50 4  152  526  508  527  Na  t  136  136  388  194  191  441  516  198  517  K  I  136  436  389  191  191  112  516  198  517  Cl  1  435  135  387  495  195  412  517  199  518  HC03  I  135  435  387  195  495  112  517  199  518  TU  |  449  119  401  511  511  158  531  516  535  Amyl  I  4 35  135  388  191  494  112  516  199  518 CO  Appendix(cont.).  Nunber  BON  TBAIT  of observations  Oric  f o r each  Choi  pair  of  T P  traits.  Alb  Bili  Alk P  SGOT  Creat  nilkl Fat  1  Protl HilkC Fat  C  Protc Ca P04 Glue BON Oric  535  Choi  532  533  T P  534  535  532  Alb  534  535  532  534 531  531  Bili  531  532  52 9  Alk  532  533  530  532  532  529  SGCT  528  529  526  528  528  527  526  Creat  530  531  52 8  530  530  527  528  524  Trig  527  528  525  527  527  524  525  521  535  Na  517  518  515  517  517  514  515  511  523  X  517  518  515  517  517  514  515  511  523  Cl  518  519  516  518  518  515  516  512  524  518  519  516  518  518  515  516  512  524  T4  535  536  533  535  535  532  533  529  539  Aayl  517  518  515  517  518  514  515  511  524  HCG3  P  .  CO  Appendix ( c o n t . ) .  Nunber o f o b s e r v a t i o n s f o r each p a i r of t r a i t s . —  r-  i  | TBAIT |  Triq  Na  K  Cl  HC03  T4  i  Anyl  1  !«„-«{« I Milk! | 1 ?at 1 | I  Protl  I  I BilkC | 1 Fat C | | ProtC | 1 Ca  |  | P04  |  1 Glue  1  | BON  |  I  Oric  |  1 Choi  I  1 T P  |  1 Alb  |  1 Bili  |  I  Alk F |  1 SGOT  I  1 Creat | 1 Triq  I  1 Na  |  1 K  i  520  525  1 Cl  I  521  524  524  I HC03  |  521  524  524  527  1 Tt  I  536  526  526  527  527  1 Amyl  I  521  515  515  514  514  1  ,  X  520  526  1  CO CM  ...  1  

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