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Intraspecific variation in non-selected natural populations of Douglas-fir (Pseudotsuga menziesii (MIRB.)… Fashler, Anita Marie Kvestich 1979

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cop.  I  INTRASPECIFIC VARIATION IN NON-SELECTED NATURAL POPULATIONS OF DOUGLAS-FIR (PSEUDOTSUGA MENSIESII ( M I R B . ) FRANCO)  by  ANITA MARIE KVESTICH FASHLER B.S.F., U n i v e r s i t y of B r i t i s h Columbia, 1976  A THESIS SUBMITTED IN PARTIAL FULFILMENT 0F THE REQUIREMENTS FOR THE DEGREE OF MASTER OF FORESTRY  in  THE FACULTY OF GRADUATE STUDIES THE FACULTY OF FORESTRY  We accept t h i s thesis as conforming to the required standard  THE UNIVERSITY OF BRITISH COLUMBIA A p r i l , 1979 ©' Anita Marie Kvestich Fashler, 1979  In presenting t h i s thesis in p a r t i a l f u l f i l m e n t of the requirements, f o r an advanced degree of the U n i v e r s i t y of B r i t i s h Columbia, I agree that the Library s h a l l make i t f r e e l y a v a i l a b l e f o r reference and study.  I f u r t h e r agree that permission f o r extensive copying of t h i s  thesis f o r s c h o l a r l y purposes may be granted by the Head of my Department or by his representatives.  It i s understood that copying or  p u b l i c a t i o n of t h i s thesis f o r f i n a n c i a l gain s h a l l not be allowed without my w r i t t e n permission.  Department of  ho  re.shy  The U n i v e r s i t y of B r i t i s h Columbia 2075 Wesbrook Place Vancouver, B. C. V6T 1W5 Canada  -iiABSTRACT  Seedlings from the International Union of Forestry Research Organizations (I.U.F.R.O) c o l l e c t i o n s i n 1966 and 1968 sampling the range of Douglas-fir from northern C a l i f o r n i a to B r i t i s h Columbia were used to e s t a b l i s h a provenance-progeny Research Forest in Haney.  t e s t at the U n i v e r s i t y of B r i t i s h Columbia  The study analyzed height growth i n a t o t a l of  384 h a l f - s i b f a m i l i e s representing 48 provenances.  The objectives were  1) to estimate the degree of genetic v a r i a t i o n between and within, provenances f o r height growth,  2) to estimate the a d d i t i v e genetic variance,  3) to estimate narrow sense h e r i t a b i l i t y , mature c o r r e l a t i o n , and  4) to estimate j u v e n i l e x  5) to s e l e c t the best provenances and progenies  f o r the Haney p l a n t i n g s i t e . Results from seed zone analysis showed that the most s i g n i f i c a n t differences in the genetic v a r i a t i o n i n height growth of j u v e n i l e Douglasf i r trees was found i n the r e l a t i v e s i z e s o f the variance between provenances (Vp) and the variance between f a m i l i e s w i t h i n provenances  (Vp^p).  Provenances most adapted to Haney conditions e x h i b i t e d Vp^p and V.^as the dominant contributors to the phenotypic v a r i a t i o n . provenances, Vp was of greater importance.  For less adapted  In a l l zones the greatest  component of variance was due to the variance of i n d i v i d u a l trees w i t h i n f a m i l i e s (V^.)- Variance due to block x provenance i n t e r a c t i o n (Vp g) was x  the next l a r g e s t variance between blocks (Vg).  Estimates f o r a d d i t i v e  genetic variance and h e r i t a b i l i t y f o r seed zones were quite h i g h .  Low  values f o r t h e i r respective standard errors indicated high r e l i a b i l i t y in the r e s u l t s .  High values of h e r i t a b i l i t y indicated that there are  opportunities f o r s i g n i f i c a n t improvement by s e l e c t i o n in D o u g l a s - f i r . Results from the j u v e n i l e x mature c o r r e l a t i o n analysis indicated that r e l i a b l e s e l e c t i o n of the best and d e l e t i o n of the poorest provenances and f a m i l i e s may begin at age f i v e . y e a r s . I t i s recommended that s e l e c t i o n in seed zones 2 and 3 and the best provenances from seed zone 1 would y i e l d the best r e s u l t s f o r the Haney site.  As an example of genetic gain, a s e l e c t i o n i n t e n s i t y of only one  in four (25%) of the top i n d i v i d u a l s was chosen.  Using t h i s low s e l e c t i o n  i n t e n s i t y , figures obtained f o r genetic gain at Haney varied from 17.90% f o r two year to 10.96% f o r eight year height growth.  S e l e c t i o n in the  best provenance, of the best seed zone could increase t o t a l height growth by almost 33%.  An a d d i t i o n a l increase of 70% i s suggested i f the best  i n d i v i d u a l w i t h i n the best provenance i s chosen.  Further gains in  height growth are possible i f higher s e l e c t i o n i n t e n s i t i e s are used.  - iv TABLE OF CONTENTS Page INTRODUCTION  1  LITERATURE REVIEW  5  1.  Components of Variance  5  2.  Heritability  7  3.  Juvenile x Mature C o r r e l a t i o n  .  10  MATERIALS AND METHODS  14  RESULTS AND DISCUSSION ;  28  1.  Genetic V a r i a t i o n i n Height Growth  28  2.  Estimation of Additive Genetic Variance (V^)  37  3.  Estimation of H e r i t a b i l i t y  39  4. 5.  Juvenile x Mature C o r r e l a t i o n S e l e c t i o n of the Best Provenances and Progenies f o r the Test S i t e  45 46  SUMMARY  54  LITERATURE CITED  57  APPENDICES: I. II. III. IV. V. VI.  Least-Squares Analysis of Variance (Seed Zones 1, 2 and 3 and I n t e r i o r ) Components of Variance (Seed Zones 1, 2 and 3 and Interior) Regression Analysis f o r Maternal E f f e c t s (Seed Zones 1 and 2) 2 H e r i t a b i l i t y (h ) and Standard Errors (S.E.) f o r Total Height Based on Individual Provenances ... Ranking According to Mean 1975 and 1978 Total Height (Seed Zones 1, 2 and 3 and I n t e r i o r ) Mean (m) and Standard Deviation (S.D.) of 1972 to 1978 Total Height f o r Provenances i n Seed Zones 1, 2 and 3 and I n t e r i o r  61 63 67 69 70 74  -  V  -  LIST OF TABLES  Table  1 2  Page  E l e v a t i o n , L a t i t u d e , Longitude and Thousand Seed Weight (TSW) of the Douglas-fir Provenances ... ... ... Analysis of Variance and Expected Mean Squares f o r Analysis of Between Provenance V a r i a t i o n  :  17 21  3- Analysis of Variance and Expected Mean Squares f o r Analysis of Within Provenance V a r i a t i o n  23  4  Total 1978 Height Differences. Between Seed Zones  29  5  Maximum, Minimum, 1978 Total Height Maximum, Minimum, 1978 Total Height  30  6  Average i n Seed Average in Seed  and Standard Deviation of Zones 1 and 2 and Standard Deviation of Zone 3  31  7  Maximum, Minimum, Average and Standard Deviation of 1978 Total Height in the I n t e r i o r Provenances  32  8  A d d i t i v e Genetic Variance (VA), Standard Errors (S.E.) (S.E. (V )/V )100 and V as a Percent of Total Phenotypic Variance f o r Total Height: Seed Zones 1 and 2  40  A d d i t i v e Genetic Variance (VA), Standard Errors (S.E.) (S.E. (V )/V )100 and V as a Percent of Total Phenotypic Variance f o r Total Height: Seed Zone 3 and I n t e r i o r Provenances  41  A  9  A  10  A  A  A  A  H e r i t a b i l i t y (h ) and Standard Error (S.E.) f o r Total 2  Height Based on Seed Zones  43  11  J u v e n i l e x Mature C o r r e l a t i o n :  Seed Zone 1  47  12  J u v e n i l e x Mature C o r r e l a t i o n :  Seed Zone 2  13  J u v e n i l e x Mature C o r r e l a t i o n :  Seed Zone 3  49  14  J u v e n i l e x Mature C o r r e l a t i o n :  I n t e r i o r Provenances  50  15  Estimates of Genetic Gain f o r Height Growth in Seed Zones 1 , 2 and 3 and I n t e r i o r Provenances  ...  48  53  - vi LIST OF FIGURES  Figure  1  Page  Location of Douglas-fir Provenances  16  ACKNOWLEDGEMENTS  I am very g r a t e f u l to Dr. 0. S z i k l a i , Faculty of Forestry and c h a i r man of my thesis committee f o r valuable assistance and continuous encouragement in the preparation of t h i s t h e s i s .  Gratitude i s also expressed to the  other members of my committee - Mr. Bruce D e v i t t , P a c i f i c Logging Co. L t d . ; Dr. C F . Wehrhahn, I n s t i t u t e of Animal Resource Ecology and Dr.  D.H..  Williams, Faculty of Forestry - f o r reviewing the thesis and c o n t r i b u t i n g t h e i r useful advice. In a d d i t i o n , acknowledgement i s extended to Drs. J . Hodges and R.G. Peterson, Faculty of A g r i c u l t u r e , Department of Animal Science f o r recommending the least-squares computer program used h e r e i n .  I am also  g r a t e f u l to Mr. M. Hoque, PhD. candidate and Mr. D. Lee, research a s s i s t a n t , Department of Animal Science f o r guidance i n handling the program. to thank Ms. G. Ho and Ms. S. Phelps  I wish  f o r t h e i r assistance i n computing.  I am g r e a t l y indebted to the B r i t i s h Columbia M i n i s t r y of Education and to P a c i f i c Logging Co. L t d . f o r t h e i r f i n a n c i a l support i n the form of the Graduate Research Engineering and Technical (G.R.E.A.T.) grant. Thanks are given to Mrs. M.A. DeVescovi f o r her t e c h n i c a l assistance and advice and to Ms. J . K e l l y f o r the typing of the manuscript. And f i n a l l y , I wish to thank my husband, Robert, f o r his support and cooperation.  INTRASPECIFIC VARIATION IN NON-SELECTED NATURAL POPULATIONS OF DOUGLAS-FIR (PSEUDOTSUGA MENZIESII (MIRB.) FRANCO)  INTRODUCTION  The study of natural v a r i a t i o n w i t h i n a tree species of commercial value i s required to derive basic information on population s t r u c t u r e . The information of greatest i n t e r e s t includes the various components of v a r i a n c e . c o n t r i b u t i n g to the t o t a l phenotypic variance and estimates of the h e r i t a b i l i t y of characters of i n t e r e s t .  The components of variance  are useful in assessing the degree of genetic d i f f e r e n t i a t i o n among the trees.  H e r i t a b i l i t y values are useful i n c a l c u l a t i o n s to determine the  amount of possible genetic gain.  An understanding of both the amount  and nature of components of variance and h e r i t a b i l i t y w i l l allow the tree breeder to increase the e f f i c i e n c y . o f tree improvement programs by evaluating the s e l e c t i o n procedures and p r e d i c t i n g what improvements can be a n t i c i p a t e d by various breeding methods (Campbell, 1972; Meier and Goggans, 1978; Rink and Thor, 1976; Owino, 1977 and Stonecypher, 1966). For example, based on information on geographic and i n d i v i d u a l tree v a r i a t i o n w i t h i n the l o b l o l l y pine range Zobel and Dorman (1973) have defined seven d i f f e r e n t provenances that could be used i n d i f f e r e n t areas as e x o t i c s . Estimates of variance components and h e r i t a b i l i t y have been previ o u s l y c a l c u l a t e d f o r several tree species (see Hattemer, 1963 f o r a review),  - 2 although few studies were designed s p e c i f i c a l l y to estimate genetic v a r i ance, the majority i n i t i a t e d as progeny and provenance t e s t s .  Such progeny  t e s t s and provenance studies designed to minimize the variance of a mean are not necessarily good f o r estimating of genetic and environmental v a r i ance (Goggans, 1962).  Therefore, experiments designed e s p e c i a l l y f o r  estimating variance components w i l l generally y i e l d more r e l i a b l e and useful information (Stonecypher, 1966).  However, the high cost of estab-  lishment, maintenance and evaluation of long term tree breeding experi=ments requires a m u l t i p l i c i t y of objectives to j u s t i f y t h e i r usefulness. S a c r i f i c e s in maximum r e l i a b i l i t y in a l l estimates f o r many tree breeding experiments must, t h e r e f o r e , be made. Recent work on the c a l c u l a t i o n of genetic parameters f o r Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) has been published.  Campbell (1972)  studied the genetic v a r i a t i o n in seedling height increment of Douglas-fir and estimated a d d i t i v e and dominance genetic variances.  Results showed  that the dominance portion of genetic variance was quite important accounting f o r 12 to 65 percent of the t o t a l genetic variance; estimates of a d d i t i v e genetic variance were low with h e r i t a b i l i t i e s ranging between 0.08 and 0.22.  L a t e r , B i r o t (1974 and 1976) studied 26 Douglas-fir prov-  enances from the f i r s t c o l l e c t i o n made by International Union of Forest Research Organizations (I.U.F.R.O) (Barner, 1973; Fletcher and Barner, 1978).  Several c h a r a c t e r i s t i c s were measured and c a l c u l a t e d h e r i t a b i l i -  t i e s ranged from 0.21 to 1.0 f o r cotelydon number, 0.05 to 0.75 f o r growth c e s s a t i o n , 0.15 to 1.51 f o r f l u s h i n g time, 0.19 to 1.0 f o r t o t a l f i r s t year height and 0.14 to 0.79 for t o t a l second year height.  In  contrast to Campbell, these r e l a t i v e l y high estimates suggested that the  - 3 above c h a r a c t e r i s t i c s are under strong genetic c o n t r o l .  Working on 10 to  15 year o l d r a c i a l crosses of D o u g l a s - f i r Orr-Ewing and Yeh (1978) also found low estimates of h e r i t a b i l i t y suggesting that both growth and s u r v i v a l were dependent on weak a d d i t i v e genetic variance.  The apparent  contradictory r e s u l t s of these studies i n d i c a t e that the estimation of genetic parameters of D o u g l a s - f i r populations require greater e f f o r t in the f u t u r e . The material used i n t h i s study ( K v e s t i c h , 1976) i s unique since i t samples the range of D o u g l a s - f i r from northern C a l i f o r n i a to B r i t i s h Columbia, and provides the opportunity to assess the genetic c h a r a c t e r i s t i c s of several d i f f e r e n t populations of Douglas-fir simultaneously.  A  combined provenance-progeny t e s t of 48 non-selected provenances of Douglasf i r including.384 open-pollinated f a m i l i e s were used to t e s t v a r i a t i o n in height growth. The objectives of the study were: 1.  To estimate the degree of genetic v a r i a t i o n f o r height growth among various D o u g l a s - f i r provenances when grown at Haney, B r i t i s h Columbia.  2.  V a r i a t i o n i n height was studied at two l e v e l s :  (a)  between provenance  (b)  w i t h i n provenance  To estimate the a d d i t i v e genetic component f o r open-pollinated progeny from d i f f e r e n t provenances.  3.  To estimate h e r i t a b i l i t y .  4.  To study the consistency of height growth of f a m i l i e s in each year since 1971 ( i . e . to estimate " j u v e n i l e x mature" c o r r e l a t i o n of progeny).  - 4 5.  To s e l e c t the best provenances and progenies f o r the Haney p l a n t ing s i t e .  - 5 -  LITERATURE REVIEW  1.  Components of Variance The phenotypic variance of an i n d i v i d u a l c h a r a c t e r i s t i c may be p a r t i -  tioned into two major categories; the genetic variance and the environmental variance (Falconer, 1960).  I t has long been recognized that the  t o t a l genetic variance could be subdivided i n t o three d i f f e r e n t portions: a f i r s t due to the average e f f e c t s of genes (V^), a second due to the i n t e r a c t i o n s of a l l e l i c gene e f f e c t s (dominance) (Vp) and a t h i r d due to i n t e r a c t i o n s of n o n - a l l e l i c gene e f f e c t s ( e p i s t a s i s )  (Vj) (Fisher, 1918).  General t h e o r e t i c a l expectations f o r f u r t h e r subdivision of the t o t a l genetic variance into portions have been c a r r i e d out more recently by several authors (Cockerham, 1954; Henderson, 1954; Kempthorne, 1955; and L i , 1954).  Their additions included i n t e r a c t i o n s among average e f f e c t s of  two or more n o n - a l l e l i c genes ( V ^ + V ^  A  more n o n - a l l e l i c dominance e f f e c t s ( V  + V  D D  + e t c . ) , i n t e r a c t i o n s of two or + e t c . ) and i n t e r a c t i o n s  D D Q  between dominance and average e f f e c t s of two or more n o n - a l l e l i c genes ^ DA V  +  DDA  V  DAA  +  V  +  -)-  e t c  T n e  di vi s i on of the t o t a l phenotypic variance  (Vp) can then be expressed as V  P  =  V  A  +  V  +  D V  +  V  DA  I  < AA  + V  DDA  +  V  +  V  AAA V  +  +  DAA -> +  where Vj equals e f f e c t s due to e p i s t a s i s , V  V  +  £  V  DD  +  V  DDD  E  +  V  +  J  equals environmental l i n e a r  e f f e c t s and Vj equals environmental non-linear e f f e c t s (genotype x environment i n t e r a c t i o n ) (Toda, 1958).  - 6 -  The components of t o t a l genetic variance may be derived from analysis of variance of h a l f or f u l T:.sibs.  For example, Lacaze and Arbez (1971)  studied w i t h i n provenance v a r i a t i o n i n Pinus s y l v e s t r i s L. and subdivided the variance into between family and w i t h i n family components.  The v a r i -  ation between family expresses the a d d i t i v e genetic variance of mother trees and the variance w i t h i n family expresses various genetic (additive or not) and environmental e f f e c t s .  With the assumption of h a l f - s i b s , the  genetic e f f e c t s include three-quarters of the a d d i t i v e genetic variance and a l l the dominance v a r i a t i o n .  Eiche and Anderson (1974) provided a good  explanation of some of the a p p l i c a t i o n s of the components of variance. the a d d i t i v e genetic variance i s high with respect to the t o t a l  If  phenotypic  variance, the breeder should aim f o r mass s e l e c t i o n and cross breeding. High dominance variation.means that family s e l e c t i o n and h y b r i d i z a t i o n of trees with high s p e c i f i c combining a b i l i t y would y i e l d the best r e s u l t s from breeding.  I f both the a d d i t i v e and dominance genetic variance e f f e c t s  are low and the genotype x environmental i n t e r a c t i o n i s high, breeding e f f o r t s should produce separate l i n e s f o r each e c o l o g i c a l region. overdominance i s important, inbreeding with the object of producing  When hybrids  between unrelated l i n e s i s the recommended program aim. Any model developed f o r the estimation of genetic variance involves various b i o l o g i c a l assumptions.  Common genetic assumptions concerning  parent populations and progenies include: a)  normal d i p l o i d and Mendelian i n h e r i t a n c e ,  b)  population in linkage e q u i l i b r i u m ,  c)  r e l a t i v e s not inbred,  - 7 d)  r e l a t i v e s random members of a non-inbred population (no selection),  e)  r e l a t i v e s are h a l f - s i b s ,  f)  no maternal or cytoplasmic e f f e c t s ,  g)  no m u l t i p l e a l l e l e s ,  h)  no e p i s t a s i s (Stonecypher, 1966 and Sprague,  1966).  Attention should be given to the v a l i d i t y of such assumptions  in the t e s t  considered and the e f f e c t s of any such i n v a l i d assumptions on the estimates of genetic variance should be discussed.  2.  Heritability H e r i t a b i l i t y (h ) i s the f r a c t i o n of the observed or phenotypic variance  which was caused by differences between the genes or the genotypes of the i n d i v i d u a l s , that i s , the r e l a t i v e amount of v a r i a t i o n passed on to the next generation by each i n d i v i d u a l (Lush, 1949).  H e r i t a b i l i t y values can  be reported in two ways; i n the "narrow" sense or in the "broad" sense (Toda, 1958).  When material i s propagated by sexual means, the non-  a d d i t i v e e f f e c t s of the genotypes are not passed on to the progeny.  This  r a t i o of the r e l a t i v e amount of a d d i t i v e genetic variance (V^) to the t o t a l phenotypic variance (Vp) i s the narrow sense h e r i t a b i l i t y (h  = n  s  ^/V )/ p  When material i s propagated v e g e t a t i v e l y , the. e f f e c t s of dominance and e p i s t a s i s are passed on since the genotypes are t r a n s f e r r e d unchanged.  In t h i s case  the r a t i o of the r e l a t i v e amount of t o t a l genetic variance (VQ) to the t o t a l 2 phenotypic variance, that i s , the broad sense h e r i t a b i l i t y (h ^ = V^/Vp)./ 2 2 Narrow sense h estimates are frequently less than h in the broad sense s  so possible gains from breeding organisms that reproduce sexually are usually smaller than gains from organisms reproducing asexually.  - 8 H e r i t a b i l i t y can be c a l c u l a t e d i n several ways (Hattemer, 1963): a) ' sib analysis (estimates h^  )  n  2 b)  clonal analysis (estimates h ^  )  c)  parent-offspring regression (estimates .h  ).  2 However, although h  estimates the r e l a t i v e importance of the genetic com-  ponents of variance, they are most useful i n the assessment of genetic gain. 2 Several factors should be kept in mind when dealing with h estimates. 2 First, h  estimates are not constant f o r a l l conditions (Zobel, 1961).  Any-  thing that causes an increase in v a r i a t i o n (eg. environmental e f f e c t s ) w i l l 2 cause a change i n the h estimate. In general, the more.uniform the s i t e 2 the greater the h value. Even the method of c a l c u l a t i o n can change the 2 2 h estimate. With such l i m i t a t i o n s , h can almost be considered a parameter of a f i e l d t r i a l rather than a genetic parameter since i t r e f e r s to s e l e c t i o n in a d e f i n i t e population under d e f i n i t e environmental conditions 2 (Hattemer, 1963).  Therefore, whenever a value i s stated f o r h  i t must  r e f e r to a p a r t i c u l a r population under p a r t i c u l a r conditions (Falconer, 1960 and Zobel, 1961). 2 In the second case, h  i s a population concept and a property of a  character and the environment.  It.does not measure the c o n t r i b u t i o n of  the genotype and the environment to the phenotype of the i n d i v i d u a l (Suzuki and G r i f f i t h s , 1976). The t h i r d f a c t o r to consider includes several problems associated 2 2 with h (Falkenhagen, 1972). The h model depends upon a s i m p l i s t i c s t a t i s t i c a l model which i s based on the a d d i t i v e genetic e f f e c t s model. However, there i s no b i o l o g i c a l reason to believe that most of the genetic  - 9 e f f e c t s r e s u l t from the a d d i t i v e e f f e c t s of many i n d i v i d u a l genes.  In f a c t ,  the dominance genetic variance (V^) i s r a r e l y found to be greater than the a d d i t i v e genetic variance (V^)(even though i t may be) because of the method used i n c a l c u l a t i o n of the components of variance. breeding value and dominance deviation are defined.  That i s , the way that Kimura and Crow (1964)  i l l u s t r a t e d the r e l a t i v e amount of a d d i t i v e genetic variance and dominance genetic variance f o r one locus i n d i f f e r e n t s i t u a t i o n s and showed that even when most of the variance i s due to dominant gene e f f e c t s , the c a l c u l a t i o n of the components of variance continues to represent, rather low dominance variance except i n a narrow range of gene frequency.  Evaluation of  2 2 2 i n d i v i d u a l h , family h and w i t h i n family h together with the components of variance w i l l provide a more accurate assessment.of what gene action i s a c t u a l l y o c c u r r i n g . For instance, i f v a r i a t i o n w i t h i n f a m i l i e s i s high and v a r i a t i o n between f a m i l i e s i s low and i f i t i s proven that most of the phenotypic v a r i a t i o n i s due to genetic e f f e c t s , then V 1979).  Mendelian genes may not form  Q  i s high  (Wehrhahn,  a representative sample of the t o t a l  genetic material although normal Mendelian inheritance i s an important assumption i n the c a l c u l a t i o n of the components of variance.  Selection  a l t e r s gene frequencies which in turn cause changes in genetic parameters so that p r e d i c t i o n of future gains based on h  estimates of parent populat-  ions may not be a p p l i c a b l e to the o r i g i n a l population.  F i n a l l y , estimates  of variance components often have very large standard errors since tests are usually r e s t r i c t e d to sampling small numbers of r e l a t i v e s and very few environments. sizes.  Reduction i n standard errors would require large sample  Pirchner (1969) estimated that a sample s i z e f i v e times l a r g e r  was required to decrease the standard e r r o r by one-half. ever,useful h  Therefore, how-  estimates may be in assessing genetic g a i n , one must be  - 10 aware of the assumptions made, and the complexities of h  2  when applying or  2 discussing h values.  3.  J u v e n i l e . x Mature C o r r e l a t i o n Juvenile x mature c o r r e l a t i o n refers to the interdependence between  q u a l i t a t i v e or q u a n t i t a t i v e data c o l l e c t e d at d i f f e r e n t : i n t e r v a l s during the l i f e cycle ( S z i k l a i , 1974) and depends upon the strength of genetic control of a character.  It i s a useful measurement to p r e d i c t the perform-  ance of the same or other a t t r i b u t e s at a more advanced stage.  Schmidt  (1963) was one of the f i r s t workers to recognize the importance of e a r l y testing.  Nanson (1965, 1968, and 1970) c a r r i e d on t h e o r e t i c a l work on the  value of e a r l y t e s t i n g concluding that at equal s e l e c t i o n i n t e n s i t i e s , e a r l y s e l e c t i o n permits higher genetic gain than l a t e s e l e c t i o n i f the c o r r e l a t i o n between e a r l y and l a t e measurements of a t r a i t are greater than that t r a i t ' s 2 • • • broad sense h . An advantage of the e a r l y t e s t i s that i t allows the evaluation of a greater number of f a m i l i e s or provenances which would give a higher s e l e c t i o n i n t e n s i t y and f u r t h e r increase the genetic gain.  Because  trees are l o n g - l i v e d organisms, f o r e s t g e n e t i c i s t s e s p e c i a l l y hope that j u v e n i l e x mature c o r r e l a t i o n s would be high t o . a l l o w the r e s u l t s of e a r l y tests to be used p r o f i t a b l y in s e l e c t i o n work.  However, i t i s important to  r e s i s t the temptation of overestimating the value of e a r l y t e s t s .  The  l i t e r a t u r e includes examples of cases both accepting and r e j e c t i n g the e f f i c a c y of e a r l y t e s t i n g i n f o r e s t t r e e s . For example, Toda (1964 and 1972) found that the diameter of ages less than s i x years c o r r e l a t e d negatively with diameter at 30 years i n L a r i x leptolepis.  (Sieb. and Zucc.) Gord.  Evidence gives varying r e s u l t s f o r  Pinus s y l v e s t r i s L. Nanson (1968) found that growth data c o l l e c t e d at age  - 11 10 c o r r e l a t e d well with estimates made l a t e r in mature stands and P a t l a j (1973) c a l c u l a t e d a p o s i t i v e c o r r e l a t i o n c o e f f i c i e n t of e a r l y to l a t e r height growth of 0.75.  However, Gierytech (1974) reported low c o r r e l a t i o n  between various growth characters of e a r l y and l a t e r years also in P. s y l v e s t r i s L. Four d i f f e r e n t pine species in Louisiana were studied over a 30-year period by Wakely (1971).  His r e s u l t s suggested that s e l e c t i o n should not  be done before age 10 and that r e l i a b i l i t y of r e s u l t s remained low at 15 years, not increasing appreciably; u n t i l approximately age 20.  Working  with Pinus ponderosa Laws and P_. monticola Dougl. Steinhoff (1974) also found that evaluation of r e s u l t s from provenance or progeny t r i a l s was not very re.liable^at ages of less than 15 to 20 years.  He said that a  minimum c o r r e l a t i o n c o e f f i c i e n t (R) of 0.70 was required f o r good r e l i a b i l i t y of e a r l y t e s t r e s u l t s since the c o e f f i c i e n t of determination (R ) would account f o r at l e a s t h a l f of the v a r i a t i o n present although s e l e c t i o n to c u l l the poorest and favour the best f a m i l i e s or i n d i v i d u a l s could begin e a r l i e r (approximately age 15).  Namkoong and Conkle (1976) showed a  decline i n the r a t i o of family v a r i a t i o n to t o t a l v a r i a t i o n between ages f i v e to eight years in P_. ponderosa. Laws.  Such rapid changes in the  r e l a t i v e s i z e of variance components are consistent with low c o r r e l a t i o n between e a r l y height and those taken at l a t e r ages.  Results with P_.  v i r g i n i a n a M i l l . have shown strong c o r r e l a t i o n between heights at very early ages and l a t e r years.  Genys and Forbes (1973) found that at l e a s t  64 percent of the best s t r a i n s f o r height at age 16 could have been predicted at age seven.  Meier and Goggans (1977) also found p o s i t i v e  c o r r e l a t i o n s between annual growth and t o t a l height although i t was not s i g n i f i c a n t u n t i l age eight suggesting that s e l e c t i o n should not proceed  - 12 u n t i l the material was at l e a s t age e i g h t . In D o u g l a s - f i r , Namkoong e t a l _ . . (1972). studied age r e l a t e d v a r i a t i o n in genetic control of height growth of a 53 year o l d p l a n t a t i o n .  Results  indicated that the genetic v a r i a t i o n changed during the d i f f e r e n t stages of the l i f e cycle ( j u v e n i l e , e a r l y reproductive and l a t e r p e r i o d s ) .  Additive  genetic variance (and therefore h ) was highest from the e a r l i e r l i f e stages of establishment to the e a r l y reproductive years. the r e l a t i v e amount of  A f t e r t h i s period  decreased as trees.reached dominance and height  growth became more uniform.  The rapid change in the r e l a t i v e s i z e of  variance components was s i m i l a r to that observed in P_. pohderosa Laws (Namkoong and Conkle, 1976) and suggests low c o r r e l a t i o n between e a r l y and l a t e r height measurements. Work i n human genetics has also pointed out that trends evident in e a r l y l i f e stages may not be c o r r e l a t e d to those at other stages.  Bock  et al_. (1963) developed a model to describe human growth that includes two l o g i s t i c f u n c t i o n s ; the f i r s t accounts f o r a component f o r p r e p u b e r t a l growth which continues in reduced degree u n t i l maturity and the second accounts f o r the c o n t r i b u t i o n of the adolescent spurt. From the v a r i e t y of evidence presented, i t can be concluded that the r e l i a b i l i t y of e a r l y tests d i f f e r s between species as well as w i t h i n species.  Generally, p o s i t i v e j u v e n i l e x mature c o r r e l a t i o n s were observed  in older provenance experiments where material was g e n e t i c a l l y very diverse. But when material was not very d i v e r s e , r e s u l t s were not as dependable (eg. f o r progeny t e s t s ) .  Longer t e s t periods seem to be required f o r progeny  tests with s i g n i f i c a n t family x s i t e i n t e r a c t i o n or f o r tests that do. not contain a wide range of v a r i a t i o n .  Also much experimentation on j u v e n i l e  -  13  -  x mature c o r r e l a t i o n assumes t h a t the r e l a t i v e r a t e s o f growth f o r e n t genotypes do not vary g r e a t l y w i t h age. ations  T h i s assumption  has i t s  s i n c e the examples g i v e n above show t h a t p a t t e r n s o f growth  change as organisms mature.  differlimit-  rate  Given t h i s e v i d e n c e , i t i s c l e a r t h a t f u r t h e r  e x p e r i m e n t a t i o n i s needed t o d e f i n e the j u v e n i l e x mature c o r r e l a t i o n o f forest  trees.  - 14 -  MATERIALS AND METHODS  Seedlings representing the open-pollinated progeny of 48 provenances from the f i r s t and second I.U.F.R.O. seed c o l l e c t i o n s i n 1966 and 1968 established i n a provenance-progeny  t e s t at the U n i v e r s i t y of B r i t i s h  Columbia Research Forest i n Haney i n 1971 provided the material f o r t h i s study.  Details on the l o c a t i o n of the provenances are given in Figure 1 and  Table 1.  The t e s t was designed as a randomized complete block r e p l i c a t e d  three times, each provenance represented by eight f a m i l i e s including f i v e seedlings per family per block.  The mother trees may be considered as  representing the natural v a r i a t i o n w i t h i n the stand and as non-related since there was no s e l e c t i o n applied and trees were separated by an i n t e r v a l that exceeded normal s e e d - f a l l distance ( B i r o t , 1974 and Lines, 1967).  Further information on experimental layout and the t e s t s i t e are  given by Kvestich (1976). The trees i n the p l a n t a t i o n were measured to the nearest centimetre i n September, 1978 f o r 1976, 1977 and 1978 height increment as well as 1978 t o t a l height.  Growth data f o r e a r l i e r years had been previously c o l l e c t e d  in 1973 and 1975 (Kvestich, 1976).  Very l i t t l e m o r t a l i t y occurred since  the 1975 measurements were taken with the average m o r t a l i t y to May 1978 being 9.8%.  Only four provenances e x h i b i t e d m o r t a l i t y figures greater than 15%.  These were provenance 99 (seed zone 1) at 29.21, provenance 115 (seed zone 2) at 40.8%, provenance 103 (seed zone 3) at 34.2% and provenance 66 (seed zone 7) at 24.2%.  Because of the high s u r v i v a l , competition e f f e c t s w i t h i n  the p l a n t a t i o n were minimal f o r a l l years, each tree having approximately the same a v a i l a b l e growing space.  However, e f f e c t s of competition are  - 15 becoming more important as the trees mature and plans are being made to t h i n the t e s t during the summer of 1979. S t a t i s t i c a l analyses were c a r r i e d on f o r i n d i v i d u a l provenances and f o r provenances arranged i n groups according to seed zones. Grouping was made according to four c a t e g o r i e s :  seed zones 1, seed zone 2, seed zone 3  and a fourth representing i n t e r i o r provenances (seed zones 4, 5, 7 and 8 ) . A l l expected mean squares were derived using a random e f f e c t s model.  The  analysis f o r v a r i a t i o n between provenances was based on the f o l l o w i n g l i n e a r model:  FIGURE 1  TABLE 1 E l e v a t i o n , L a t i t u d e , Longitude, and Thousand Seed Weight (TSW) of the Douglas^fir Provenances Climatic Region  Provenance No.  la  23 Cassidy 32 Duncan  laa  55 Gard Station  lb  90 95 96 99  lc  104 111 117 124  01 a l l a Corvallis M i l l City Roseburg  KJ  Ashland* Hawkinsville Big Bar Lower Lake  Elev. ( f t )  Latitude deg. min.  .Longitude deg. min.  TSW  650 200  49 48  03 45  123 123  57 45  9.1725 10.2133  1500  48  00  123  05  9.0224  1100 250 550 900  43 44 44 43  05 42 48 19  123 123 122 123  34 13 24 30  14.2130 11.5786 11.2457 11.4927  4900 3500 4300 3100  42 41 40 38  05 47 47 50  122 122 123 122  39 40 12 42  15.8762 16.1968 16.7025 16.0461  550  50  27  127  27  10.0874  2a  12 Jeune Landing  2b  51 Humptuli ps 52 Matlock 53 Matlock  450 1650 400  47 47 47  19 18 15  123 123 123  54 26 25.  8.8665 10.0426 9.9037  2c  67 Naselle 79 P r i n d l e 83 Vernonia  150 1500 700  46 45 45  22 37 46  123 122 123  44 08 13  10.5983 11.7812 11.7173  2d  87 Waldport 89 C o q u i l l e  200 :.250  44 43  24 12  123 124  52 10  9.5850 10.4077  * Provenances excluded from the a n a l y s i s , (see page 21)  TABLE 1 (continued) Climatic Region 2d  Provenance No. 91 Brookings 92 Burnt Woods*  Elev. ( f t . )  Latitude min. deg.  Longitude deg. min.  TSW  1000 1100,  42 44  07 36  124 123  12 42  12.0091 9.8074  2e  115 Areata*  2900  40  54  123  46  13.9353  3a  29 Caycuse  . 700  48  55  124  26  10.1977  3b  25 27 40 42  Squamish Chilliwack Darrington Perry Creek*  50 3000 500 2000  49 49 48 48  47 06 16 03  123 121 121 121  09 42 28  9.0917 7.3223 10.9612 10.9612  3c  60 61 73 76  Denny Creek Gold Bar* Cougar Packwood  1800 400 1650 2150  47 47 46 46  24 51 05 34  121 121 122 , 121  32 39 18 40  10.9442 9.6996 10.9332 8.4889  3d  86 97 101 103  Pine Grove* Detroit Oakridge Wolfcreek*  2400 1600 2900 1400  45 44 43 42  06 44 54 41  121 122 122 123  23 10 22 23  14.2043 11.1186 12.4379 12.5340  3e  112 Dunsmuir 113 Burney*  3300 3350  41 41  12 05  122 121  18 39  15.1707 18.0451  10  51  01  125  36  10.3733  2600 2100 1600 1650  48 47 46 45  23 13 00 48  120 121 121 121  24 07 10 41  13.2947 10.3615 13.5443 12.2576  4 5b  6 Klinaklini 46 64 77 80  Twi.sp Cle El urn Glenwood Willard  * Provenances excluded from the a n a l y s i s , (see page 21)  •-38  TABLE 1 (continued) Climatic Region  Provenance No.  Elev. ( f t . )  Lati tilde deg. min.  Longitude deg. min.  TSW  7a  10 Revel stoke 18 Salmon Arm  2000 1550  51 50  00 44  118 119  12 13  8.8204 8.7206  7b  28 Nelson 16 Spokane  2700 2000  49 47  30 47  117 117  16 12  9.2489 13.5056  8  11 Golden  2700  51  23  117  00  9.0777  - 20 -  Y  ijkm  =  H  +  i  P  +  w h e r e  Y  E  +  B  j - (PB)^-  +  F/P  k ( i )  the j  t  (BxF/P)  J k ( i )  m(ijk)  the mean measurement of the  ijkm  +  tree in the k^* family in 1  block  h  mean of a l l trees over a l l f a m i l i e s , blocks and provenances provenance e f f e c t B. = block e f f e c t (PB) F/P  = provenance x block i n t e r a c t i o n  1 d  = family w i t h i n provenance  k(i)  (BxF/P)j ^.^ = block x family w i t h i n provenance i n t e r a c t i o n (sampling error) k  ^m(ijk)  =  e x  P  e r i m e n  '  t a  l  error  A s t a t i s t i c a l t e s t f o r the s i g n i f i c a n c e of the highest order i n t e r a c t i o n (BxF/P) was performed by using ANOVAR, a computer package a v a i l a b l e from the U n i v e r s i t y of B r i t i s h Columbia Computing Centre, and was found to be i n s i g n i f i c a n t at the one percent l e v e l .  Therefore, a revised model was  used f o r the genetic analysis in the form: Y. ..  ijkm  where E^-j  =  M + P. + B. + ( P B ) . . + F/P. / .> + E ,. .. s ** i j ij ' k(i) m(ijk)  includes both the sampling and experimental e r r o r .  pooled e r r o r term increases the p r e c i s i o n of the F t e s t s .  This  The expected  mean squares of the analysis of variance are given in Table 2.  - 21 TABLE 2 A n a l y s i s of V a r i a n c e and Expected Mean Squares f o r A n a l y s i s of Between Provenance V a r i a t i o n  Source of Variation  df  Provenance  P-l  Block  b-1  Provenance x Block  V  e p  + btV  p / p  where  V  V  B  + bftVg  PB  V. ep  (t-1)  p = No. b = No. f, = No. t = No. = van P  PB V  ftV  +  p B  V + btV ep F/P  P(b-l)(f-l) + bpf  ftV  +  +  P'(f-D  (Pooled)  Squares  + ftVp  ep  (p-D(b-l)  Family/Provenance Error  Expected Mean  B  F/P  = van" ance due to b l o c k x provenance = v a n ' ance between  n  blocks  = v a n ' ance between f a m i l i e s  = residual V ep  variance  interaction  (pooled)  within  provenance  bftV  r  - 22 -  The analysis f o r w i t h i n provenance v a r i a t i o n was based on the following 1inear model:  Y  ijk  +  B  i  +  F  j  +  (  B  F  )  i j  +  E  k(ij)  where Y... = the mean measurement of the i jk i block t  tree in the  family in the  h  y = mean of a l l f a m i l i e s over a l l blocks B. = block e f f e c t Fj = family e f f e c t (BF)..j = block x family i n t e r a c t i o n F-k(ij) = experimental  error.  Table 3'rishows the appropriate a n a l y s i s of variance showing the expected mean squares.  -  23 -  TABLE 3 A n a l y s i s of V a r i a n c e and Expected Mean Squares f o r A n a l y s i s o f Within Provenance V a r i a t i o n  Source of Variation  df  Block  b-1  Family  Expected Mean  f-1  Block x Family Error  where  V  g  + tVg  (b-l)(f-l)  V  g  +  bf(t-l)  V  Vp = v a r i a n c e between  tV  Dr  V = residual e  variance  + btV  p  B R  g  families  V - = v a r i a n c e due to block x f a m i l y Dr  F  Squares  interaction  - 24 I f the i n t e r a c t i o n term was found to be i n s i g n i f i c a n t , the model was s i m p l i f i e d to include the e f f e c t of block x family i n t e r a c t i o n i n the pooled error term.  Corresponding s i m p l i f i c a t i o n of the analysis of variance and  expected mean squares was also done. The analysis of variance was performed by l e a s t squares and maximum l i k e l i h o o d procedures to obtain unbiased estimates of the constants and the sums of squares f o r the t e s t s of s i g n i f i c a n c e (Harvey, 1975).  This was  required since the designrwas unbalanced due to missing trees caused by mortality.  The procedure u t i l i z e d the Mixed Model Least-Squares and Maxi-  mum L i k e l i h o o d computer programs LSMLGP and LSML76 of Harvey (1968 and 1976).  A l i m i t a t i o n of these programs was using a maximum of ten provenances  f o r each seed zone grouping..; ^Provenances were eliminated i f the data included missing subclasses ( f o r example, missing f a m i l i e s w i t h i n b l o c k s ) .  Further  omissions were made to obtain a sampling of provenances throughout each seed zone range.  Omitted provenances are marked i n Table 1.  The programs  were designed f o r use in animal breeding research and compute various parameters including least-squares analysis of variance, estimates of variance and 2 covariance components, estimates of h  and i t s standard e r r o r as well as  genetic, phenotypic and environmental c o r r e l a t i o n s . Assumptions made f o r the analysis of variance included momogeneous variances, independent errors and normal d i s t r i b u t i o n of observations. Through p l o t t i n g of the data, the homogeneity of variances and normality of observations appeared to be v a l i d .  The assumption of independence  of errors was assumed c o r r e c t . Assumptions associated with the genetic i n t e r p r e t a t i o n of the components of variance included those mentioned i n the l i t e r a t u r e review.  The  - 25 assumptions  that r e l a t i v e s are not inbred and are random members of a non-  inbred population are v a l i d .  The maternal e f f e c t s were tested by estimat-  ing the c o r r e l a t i o n c o e f f i c i e n t between thousand seed weight (Yao, and t o t a l height in the f i r s t and second years. assumptions  1971)  Deviation from these  i s most l i k e l y with respect to the r e l a t i o n s h i p of i n d i v i d u a l s  w i t h i n a family.  I f the open-pollinated seeds had only a few pollen parents  then they would be more c l o s e l y r e l a t e d than h a l f - s i b s and the estimates of 2 h would be biased upward. The genetic components of variance are estimated from the analysis of variance as f o l l o w s : a)  variance among outcrossed h a l f - s i b family means (Vp and Vp/p) i s the covariance of h a l f - s i b s and estimates 1/4 V^.*  b)  variance w i t h i n outcrossed h a l f - s i b . . f a m i l i e s (V 3/4 V  A  +  V,.  .)estimates cp  The analysis of open-pollinated progenies can therefore only y i e l d information about a d d i t i v e genetic variance since the dominant e f f e c t s are confounded in  \T. ep  There are some expected problems from the a n a l y s i s .  The  number of  f a m i l i e s per provenance i s rather small (n = 8) leading to possible sampling errors f o r males and females selected from the population ( B i r o t , 1974). (n ^ 5). 1977).  Also the number of open-pollinated progeny f o r each family i s low Such small sample sizes may lead to large sampling errors  (Owino,  F i n a l l y , genetic variance estimates based on data from a s i n g l e  * The standard e r r o r of V was c a l c u l a t e d from the formula derived by Anderson and Bancroft (1952). fl  - 26 environment may be biased upward because of confounding of a d d i t i v e genetic e f f e c t s with the i n t e r a c t i o n of a d d i t i v e genetic e f f e c t s and the p a r t i c u l a r environment i n which the experiment i s conducted (Gardner, 1963). H e r i t a b i l i t y estimates based on i n d i v i d u a l s e l e c t i o n derived from the analysis of variance were c a l c u l a t e d from the formulas: a)  f o r seed zones: variance among f a m i l i e s = covariance h a l f - s i b s cov HS = h V A V  A  then h  =  F/P  4 V  = V  where  —  F/P  V  ep  +  i s the expected mean square f o r the pooled B x F/P and  within p l o t v a r i a t i o n , b)  f o r i n d i v i d u a l provenances assuming s i g n i f i c a n t block x family interaction : V then h  A  2  - 4 V  F  F  4V  = V  e  +  V  and  BxF + V  F  f o r i n d i v i d u a l provenances assuming n o n - s i g n i f i c a n t block x family i n t e r a c t i o n : '*  F  V + V ep  - 27 where V  i s the expected mean square f o r the pooled BxF and  within plot variation. The estimation of j u v e n i l e x mature c o r r e l a t i o n s was done by simple c o r r e l a t i o n s between progeny annual growth and t o t a l height and simple c o r r e l a t i o n s between t o t a l heights at various ages.  Progeny annual  growth was computed from the average of 1975, 1976, 1977 and 1978 height increments.  The l a s t four years were chosen since a comparison of the  standard deviations of height increments f o r each of the years using the c o e f f i c i e n t of v a r i a t i o n (Steel and T o r r i e , 1960) indicated that the v a r i a b i l i t y s t a b i l i z e d by 1975.  - 28 RESULTS AND DISCUSSION  I n i t i a l analysis of the v a r i a t i o n in height growth f o r a l l seed zones i s summarized i n Table 4 g i v i n g the o v e r a l l average eight year height growth f o r the provenance-progeny t r i a l .  Large d i f f e r e n c e s f o r the 1978  t o t a l height are shown between seed zones.  The r e s u l t s f o r a l l provenances  within seed zones are presented i n Tables 5, 6 and 7. between provenances w i t h i n seed zones i s also found.  Significant variation The large range i n  provenance means indicates considerable genetic v a r i a t i o n .  This high v a r i -  a b i l i t y in height growth suggests that s u b s t a n t i a l gains can be made through s e l e c t i n g the most d e s i r a b l e provenances.  These f i n d i n g s are consistent  with those of Kvestich (1976) that i n d i c a t e d s i g n i f i c a n t between and w i t h i n provenance v a r i a t i o n f o r 1972 to 1975 height increment and f o r 1975 t o t a l height. Least-squares analysis of variance f o r 1972 to 1978 t o t a l height i n seed zones 1, 2 and 3 and the i n t e r i o r provenances reveal s i g n i f i c a n t d i f f e r ences between b l o c k s , provenances and f a m i l i e s w i t h i n provenances as well as s i g n i f i c a n t block x provenance i n t e r a c t i o n f o r a l l seed zones (Appendix I). S i g n i f i c a n t family w i t h i n provenance v a r i a t i o n gives the opportunity f o r s e l e c t i o n of the best f a m i l i e s i n the best provenances. s t a t i s t i c a l l y s i g n i f i c a n t at the 99% l e v e l .  A l l tests are  The only exception i s the  v a r i a t i o n between blocks i n the seed zone 3 provenances which i s s i g n i f i c a n t at the 95% l e v e l only f o r 1972, 1975 and 1976 and not s i g n i f i c a n t f o r 1977 and 1978 t o t a l height. 1.  Genetic V a r i a t i o n in Height Growth Studies of the genetic v a r i a t i o n i n the Douglas-fir population were  - 29 TABLE 4 Total 1978 Height Differences Between Seed Zones  Seed Zone  Average 1978 Total Height (cm.)  Standard Deviation (cm.)  1  284.2  97.7  2  347.8  101.4  3  313.4  96.2  4  273.9  80.1  5  253.3  86.8  200.4  68.4  8  164.7  76.0  Average  297.2  104.9  7  ,  - 30 TABLE 5 Maximum, Minimum, Average and Standard Deviation of 1978 Total Height i n Seed Zones'1 and 2 (cm.) Seed Zone  Prov.  Max.  Min.  Average  Standard Deviation  23 32 55 90 95 96 99 104 111 117 124  472.0 480.0 493.0 492.0 500.0 570.0 . 470.0 405.0 557.0 366.0 411.0  100.0 107.0 70.0 31.0 151.0 175.0 92.0 52.0 66.0 71.0 68.0  288.6 345.3 319.5 297.8 327.7 372.1 281.8 211.3 . 227.2 211.8 238.1  93.1 73.4 77.7 102.5 76.3 72.7 88.7 82.9 77.6 64.7 88.4  12 51 52 53 67 79 83 87 89 91 92 115  530.0 564.0 590.0 580.0 570.0 503.0 542.0 533.0 620.0 520.0 585.0 401.0  313.0 335.7 380.4 387.7 377.6 323.1 358.2 363.7 383.4 370.8 315.6 223.7  92.2 87.7 100.3 101.5 104.8 80.9 84.5 82.0 109.1 83.0 106.9 77.3  74.0 140.0 134.0 115.0 115.0 37.0 • 158.0 133.0 68.0 165.0 91.0 84.0  - 31 TABLE 6 Maximum, Minimum, Average an Standard Deviation of 1978 Total Height n Seed Zone 3 (cm.) Prov.  Max.  Min.  29 25 27 40 42 60 61 73 76 86 97 103 101 112 113  458.0 605.0 520.0 475.0 575.0 500.0 500.0 510.0 475.0 410.0 470.0 378.0 485.0 425.0 405.0  72.0 133.0 125.0 100.0 174.0 145.0 149.0 66.0 149.0 57.0 153.0 61.0 126.0 84.0 48.0  Average 324.3 381.3 369.3 307.9 384.3 299.2 341.9 330.0 315.5 237.4 331.9. 228.5 314.9 254.2 237.9  Standard Deviation 68.5 86.1 89.4 79.7 94.1 92.5 75.9 87.9 73.2 86.2 67.7 86.0 82.0 100.5 85.0  - 32 TABLE 7 Maximum, Minimum, Average and Standard Deviation of 1978 Total Height i n the I n t e r i o r Provenances (cm.) Zone  Prov.  Max.  4  6  434.0  5  46 64 77 80  368.0 420.0 490.0 442.0  7  10 18 28 66  354.0 330.0 384.0 330.0  8  11  331.0  >  Min.  Average  Standard De\  273.9  80.1  192.9 263.0 269.1 287.2  63.7 88.5 95.9 64.2  87.0 63.0 78.0 49.0  192.0 183.6 237.7 185.3  63.4 59.1 72.0 64.0  37.0  164.7  76.0  78.0 60.0 58.0 • , 45.0 115.0  - 33 performed to e v a l u a t e the o r i g i n variation  i n h e i g h t growth.  o f the l a r g e amount o f observed p h e n o t y p i c  V a r i a t i o n was s t u d i e d a t two l e v e l s ;  within  provenance a n a l y z i n g provenances i n d i v i d u a l l y and between provenance i n g provenances grouped a c c o r d i n g to seed zone g r o u p i n g s . v a r i a t i o n w i t h i n provenances results  for several  sampling  errors.  is  sample provenances was found to be low due to  (that i s ,  between provenance v a r i a t i o n ) . improved a l l  test  The  seed zone g r o u p i n g s .  provenance v a r i a n c e (Vp)  large  estimates.  is  In seed zone 1 the component f o r  a p p r o x i m a t e l y f i v e times  l a r g e r than the com-  ponent f o r f a m i l y w i t h i n provenance ( V p ^ ) , a c c o u n t i n g f o r an average p  26% o f the t o t a l p  phenotypic v a r i a t i o n  i n the o p p o s i t e d i r e c t i o n i s  1976.  from 1972 to 1978.  from 14.43% i n 1972 -to 32.67% i n 1978 i s  4.54% although  shown.  the components o f v a r i a n c e were s l i g h t l y  of v a r i a n c e f o r b l o c k s  (V ) g  A gradual  A trend to  h i g h e r i n 1975 and  due to the component  which accounted f o r only about 1%.  to be more important as the t e s t ages.  of  increasing  noted f o r Vp^p which d e c l i n e d from 5.91%  The s m a l l e s t amount o f p h e n o t y p i c v a r i a t i o n i s  appears  of  g i v e s the components o f v a r i a n c e f o r between provenance  variation for a l l  trend f o r V  for  large  t h e r e f o r e , p l a c e d i n the a n a l y s i s  i n c r e a s e i n sample s i z e s u b s t a n t i a l l y Appendix II  The a n a l y s i s  not i n c l u d e d because the r e l i a b i l i t y of the  G r e a t e r e f f o r t was,  pooled i n f o r m a t i o n  analyz-  The amount of  This  component  variation  accounted f o r by b l o c k x provenance i n t e r a c t i o n ( V p g ) shows a s t e a d i l y x  decreasing  t r e n d from 18.78% i n 1972 to 7.21% i n 1978.  c o n t r i b u t i o n to phenotypic v a r i a t i o n (V p) e  (50% to 60%).  Again,  is  The  greatest  made by the pooled e r r o r v a r i a n c e  t h e r e appears  to be a d e c r e a s i n g  trend in V  from 1972 to 1978. The components o f v a r i a n c e f o r seed zone 2 suggest a d i f f e r e n t p a t t e r n of v a r i a t i o n .  The i n c r e a s i n g  trend in V  p  and d e c r e a s i n g t r e n d i n  - 34 Vp^p i s s i m i l a r to that f o r the provenances i n seed zone 1. variance component Vp i s less than Vp^ f o r a l l years. p  However, the  In seed zone 2,  Vp i s about one-half Vp^ on average (3.88% compared to 8.07%) although i n p  1978 Vp and Vp^ are almost equal.  The Vp^ i s s l i g h t l y higher in seed  p  p  zone 2 i n r e l a t i o n to seed zone 1, showing a decreasing trend.  Similarly,  the components Vp g and V  are also higher f o r seed zone 2; the average  amount of v a r i a t i o n i n V  i s 8.07% and 70.98% f o r the pooled e r r o r  x  variation.  p x B  Variance due to pooled e r r o r remains constant throughout the  t e s t period. Observations of i n t e r e s t concerning the variance components of seed zone 3 include that Vp and Vp^p are nearly equal i n 1972 with Vp becoming about twice Vp^p by 1978.  On average, V  p  i s greater than Vp^p but not by the  f i v e - f o l d degree observed i n seed zone 1.  The components of variance f o r blocks  are negative (represented by zeros) f o r a l l years.  Variance due to block x  provenance i n t e r a c t i o n accounts f o r greater v a r i a t i o n than i n e i t h e r seed zone 1 or seed zone 2, averaging about 25%.  The amount of v a r i a t i o n i n  V  constant at an average of 56%.  i s comparable t o seed zone 1, remaining  The i n t e r i o r provenances show variance patterns s i m i l a r to seed zone 1 f o r a l l components except Vp g which accounts f o r about h a l f as much x  v a r i a t i o n as seed zone 1.  However, Vp g remains approximately constant i n x  the i n t e r i o r provenances while i n seed zone 1 i t e x h i b i t s a d e c l i n i n g y e a r l y trend.  Again Vp i s about f i v e times V p . / p  The variance component Vg  accounts f o r the l e a s t and the variance component V p accounts f o r the g  greatest amount of t o t a l v a r i a t i o n .  No trends f o r increasing or decreasing  component s i z e are shown. The preceding discussion has shown that the patterns of v a r i a t i o n  d i f f e r appreciably between the Douglas-fir populations studied, showing differences in the r e l a t i v e s i z e s ' o f some of the components of variance. In a l l seed zones, over h a l f of the t o t a l phenotypic v a r i a t i o n (53.68% to 62.32%) i s concentrated i n the t r e e s - w i t h i n - p l o t s term, V ^ . e r r o r expresses various e f f e c t s ; genetic (includes 3/4  This pooled  + Vp with the  assumption of h a l f - s i b s ) and environmental (measuring e r r o r , e t c . ) .  Because  t h i s value does not d i f f e r s u b s t a n t i a l l y between seed zones, i t may be assumed that e f f e c t s such as measuring errors and V^ also remain constant between seed zones.  The w i t h i n provenance variance, which i s i n f a c t the  difference among f a m i l i e s w i t h i n provenance, also does not vary s i g n i f i cantly between the seed zones accounting f o r 5.12% to 8.07% of the t o t a l variation. However, the r e l a t i o n s h i p of Vp^p with respect to Vp varies among the four groups.  For seed zone 1 and the i n t e r i o r provenances, Vp i s greater  than Vpyp by a magnitude of approximately f i v e with V 13.15% to 25.85% of the v a r i a t i o n .  p  accounting  f o r from  This magnitude i s reduced to approx-  imately a two-fold d i f f e r e n c e f o r seed zone 3.  The r e l a t i o n s h i p i s  reversed f o r seed zone 2 where the part of the v a r i a t i o n due to provenance e f f e c t i s less than the percent due to family w i t h i n provenance v a r i a t i o n . I f s e l e c t i o n i s c a r r i e d out at Haney in the f u t u r e , the approach w i l l be governed by the differences in the r e l a t i v e sizes of Vp and Vp^p.  In cases  where Vp i s greater than Vpyp, provenance s e l e c t i o n f o r increased t o t a l height i s i n d i c a t e d ; in cases where V  p  i s less than Vp^ , family s e l e c t i o n p  would be a more e f f e c t i v e strategy. The above differences i n variance patterns are caused by differences in adaptation of the provenance m a t e r i a l . s i t e (about 400 feet) w i t h i n seed zone 3.  Haney.represents a low e l e v a t i o n  -  36 -  The provenances from seed zone 1 and the i n t e r i o r zone r e p r e s e n t t r e e s adapted to very d i f f e r e n t c o n d i t i o n s greatest  than the Haney area l e a d i n g to the  d i f f e r e n c e s i n performance to be e v i d e n t a t the provenance  These provenances a l s o e x h i b i t e d the g r e a t e s t Vpyp.  There s t i l l  may be s i g n i f i c a n t  these provenances but i t  is  level.  d i f f e r e n c e s between Vp and  f a m i l y w i t h i n provenance v a r i a t i o n  not r e v e a l e d a t the Haney s i t e .  in  The provenances  i n seed zone 3 expressed a reduced d i f f e r e n c e between the magnitude o f Vp and Vp^p.  These provenances o c c u r i n the same seed zone as Haney but  r e p r e s e n t high e l e v a t i o n l o c a t i o n s adapted to the t e s t s i t e . e l e v a t i o n s most s i m i l a r most adapted to Haney  (average  e l e v a t i o n 1895 f e e t ) not w e l l  The provenances o f seed zone 2 r e p r e s e n t low  to Haney (average  e l e v a t i o n 685 f e e t ) and hence are  conditions.  The above r e s u l t s  i n d i c a t e t h a t i n the more adapted provenances, Vpyp  and V p are the dominant c o n t r i b u t o r s g  to the phenotypic v a r i a t i o n .  For  l e s s adapted provenances, Vp i s  o f g r e a t e r importance and becomes  than V p ^ .  o f the g e n e t i c v a r i a t i o n between the  various  The o v e r a l l  Douglas-fir  results  populations  greater  suggest t h a t a combination o f provenance,  f a m i l y w i t h i n provenance and w i t h i n - p l o t s e l e c t i o n would y i e l d the  greatest  i n c r e a s e i n h e i g h t growth.  all  s e l e c t i o n schemes  is  The amount o f improvement p o s s i b l e  s u b j e c t to the l e v e l  v a r i a t i o n within populations  will  of h .  Differences  for  i n the  a l s o i n f l u e n c e the e f f e c t o f s e l e c t i o n  in  f o r e s t t r e e improvement. F l u c t u a t i n g estimates zone 3 provenances) combined i n d i c a t e s  of V  averaging significant  p x B  (7.42% f o r i n t e r i o r to. 25.33% f o r seed  a r a t h e r high 15.67% f o r the f o u r  areas  genotype x environment i n t e r a c t i o n .  Ideally,  the s i t e chosen f o r a provenance o f progeny t e s t should be r e l a t i v e l y homogeneous  l e a d i n g to l i t t l e o r no b l o c k x provenance i n t e r a c t i o n .  The Haney  - 37 s i t e was selected f o r i t s r e l a t i v e homogeniety.  However, other f a c t o r s  such as an invasion of grass from' a nearby experiment has increased the differences between blocks.  S i g n i f i c a n t genotype x environment i n t e r a c t -  ion suggests that s e l e c t i o n from provenances and f a m i l i e s f o r increased height growth, f o r a l l l o c a t i o n s may not be possible,, and i n d i c a t e s the need f o r a greater number of t e s t s i n d i f f e r e n t s i t e s to f u r t h e r i n v e s t i gate genotype x environment e f f e c t s .  That i s , more than one seed orchard  would be required to provide improved seed f o r alT l o c a l i t i e s .  I t should  be noted that the r e l a t i v e s i z e of V g i s diminishing f o r two regions p x  i n d i c a t i n g that the genotype x environment effect.may be decreasing in importance.  2.  Estimating A d d i t i v e Genetic Variance (V^) The c a l c u l a t i o n of V^, according to the r e l a t i o n s h i p Vp/p  <•" =  depends upon several assumptions already mentioned on page s i x of the l i t e r a t u r e review.  The assumptions that the r e l a t i v e s are not inbred and  are random members of a non-inbred population have been shown to be correct.  Further i n v e s t i g a t i o n on the assumption concerning maternal  e f f e c t s was necessary since the reported r e s u l t s r e l y on development in the j u v e n i l e stage.  The existence of maternal e f f e c t s , can have s i g n i f i -  cant consequences on estimated parameters. Maternal e f f e c t s were estimated using the c o r r e l a t i o n , c o e f f i c i e n t ( B i r o t , 1976) between thousand seed weight (TSW) and t o t a l height in 1972 and 1973.  Provenances in ^seed zones 1 and 2 were analyzed.  The amount of  v a r i a t i o n due to maternal e f f e c t s : in seed zone 1 was 2.29% and 2.61% r e s p e c t i v e l y f o r 1972 and 1973 growth.  The figures f o r seed zone 2 were  - 38 3.14% f o r 1972 and 4.16% f o r 1973.  The very low c o e f f i c i e n t s of determin-  2 ation (R ) f o r these provenances suggested that analysis did not require to be continued f o r the remaining provenances.  On the basis of the  r e s u l t s f o r seed zones 1 and 2 .the assumption of no maternal e f f e c t s f o r a l l provenances and progenies was accepted, although the large amount of v a r i a t i o n in TSW w i t h i n provenances reported by Yao (1971) suggests that TSW may s i g n i f i c a n t l y influence height growth. The a d d i t i v e genetic variance, i t s standard e r r o r and  expressed as  a percent of t o t a l phenotypic. v a r i a t i o n f o r t o t a l height (Appendix II) given in Tables 8 and 9. ion i s accounted f o r by  Rather a high percentage of t o t a l height v a r i a t with average estimates ranging from a low of  21.80% f o r seed z o n e ! to a high of 32.27% f o r seed,zone 2. for  are  The component  remains r e l a t i v e l y constant between the years accounting f o r an  average of 24.74% of the t o t a l v a r i a t i o n in height f o r a l l regions combined. These estimates of  are lower than (approximately one-half) the estimates  obtained by B i r o t (1976) who analyzed f i r s t and second year height growth f o r a sample of the I.U.F.R.0. c o l l e c t i o n from Washington State grown i n France. study.  Several of the provenances tested at Haney were included in his The estimates of  54% and 42% r e s p e c t i v e l y .  f o r f i r s t and second year t o t a l heights were Differences i n the estimates of B i r o t and those  presented in t h i s paper may be due to differences in the model, the t e s t s i t e , and the t e s t m a t e r i a l .  B i r o t s analysis did not take account of 1  block e f f e c t s causing.differences due to blocks to be incorporated in.other variance components.  The t e s t s i t e he used was very homogeneous, leading  to decreased environmental v a r i a t i o n and increased estimates of genetic parameters.  The t e s t material included d i f f e r e n t provenances which could  e x h i b i t s l i g h t l y d i f f e r e n t patterns of v a r i a t i o n and only f i r s t and second  - 39 year height growth were s t u d i e s .  I t w i l l be i n t e r e s t i n g to continue compar-  isons of the p a r a l l e l provenance-progeny tests between the two locations i n the f u t u r e . Examination of the r a t i o of the standard e r r o r of  over  multi-  p l i e d by 100 (S.E. ( V ) / V ) 100) given i n Tables 8 and 9 reveal d i f f e r e n c e s A  between seed zones.  A  The average r a t i o s f o r seed zone 1 and the i n t e r i o r  provenances are very s i m i l a r at 14.40% and 15.21% r e s p e c t i v e l y .  Results  f o r seed zones 2 and 3 are also comparable to each other averaging lower at values of 6.86% and 7.21%.  This s i m i l a r i t y of the r a t i o s (S.E.  (V )/ A  V ) 100) f o r pairs of provenance groupings i s s i g n i f i c a n t since i t supports A  the explanation proposed e a r l i e r that the more adapted provenances i n seed zones 2 and 3 e x h i b i t lower v a r i a t i o n i n r e s u l t s while those less adapted provenances from seed zone 1 and the i n t e r i o r express higher variability. 3.  2 Estimation of H e r i t a b i l i t y (h ) 2 The importance of h in the estimation of genetic gain has been pre2  viously outlined.  The h  estimates presented i n t h i s t h e s i s were c a l c u -  lated according to the r e l a t i o n s h i p s given on page 23 of the Material and Methods section and were based on i d e n t i c a l assumptions to those given f o r V  The assumption of open-pollinated progeny representing h a l f - s i b s i s considered 2 accurate since the h estimates are a l l less than one. 2 A>  Estimates of h  according to seed zones f o r t o t a l height growth given  in Table 10 i n d i c a t e that there i s moderate control of the v a r i a t i o n found i n t h i s c h a r a c t e r i s t i c . H e r i t a b i l i t y estimates range from 0.28 f o r 1978 t o t a l height i n the i n t e r i o r provenances to 0.52 f o r 1972 t o t a l height i n the seed zone 2 provenances and average 0.38 over a l l years and regions.  -  40 -  TABLE 8 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 ) , Standard Er as a P e r c e n t of T o t a l Phenotypic V a r i a t i o n  ( S . E . ) , ( S . E . ( V ) / V ) 1 0 0 and V T o t a l H e i g h t : Seed Zones 1 and 2 A  A  A  Location Year  Seed Zone 2  Seed Zone 1 +  VA  S.E.  S.E.(VA) 100(—' —— ) VA  S.E.(V ) A  Pheno. Var.  VA  +  S.E.  -)  100(-  v  %  A  Pheno. Var.  %  1972  .41.12 +  6.01  14.61  23.64  94.86  +  5.82  6.14  43.40  1973  135.26 +  19.67  14.54  20.96  275.29  +  17.76  6.45  36.48  1974  288.35 +  42.05  14.58  20.80  485.14  +  33.12  6.83  32.28  1975  594.67 +  77.06  12.96  25.44  694.04  +  50.24  7.24  28.52  1976  945.28 + 130.88  13.85  24.08  1209.20  +  86.94.  7.19  27.72  1977  1299.47 + 194.86  15.00  19.52  2040.77  +  142.35  6.98  29.08  1978  1911.97 + 291.29  15.24  18.16  2854.15  +  203.66  7.14  28.40  14.40  21.80  6.86  32.27  1  Average  E s t i m a t e s of V are a l l s i g n i f i c a n t at the 0.01 l e v e l . A  - 41 -  TABLE 9 A d d i t i v e Genetic Variance ( V ) , Standard Errors ( S . E . ) , ( S . E . ( V ) / V ) 1 0 0 and V as a Percent of Total Phenotypic V a r i a t i o n f o r Total Height: Seed Zone 3 and I n t e r i o r Provenances A  A  A  A  Location Year  Seed Zone 3 v  + A  S.E. •  I n t e r i o r Provenances —)  100(  v  A  Pheno. Var. %  1972  VA  +  S.E.  -) Pheno. Var.  100(-  .  %  52.30 1+  3.49  6.67  26.76  23.31  +  3.27  14.01  25.24  1973  148.16  +  10.52  7.10  20.76  102.18  +  13.64  13.35  25.72  1974  302.67  +  21.43  7.08  21.32  232.42  +  30.88  13.29  26.96  1975  470.87  +  34.97  7.43  20.92  392.19  +  55.64  14.19  24.32  1976  711.38  +  55.79  7.84  18.56  +  89.14  15.37  20.68  1977  1302.22  +  96.06  7.38  21.00  940.67  +  112.72  12.40  19.12  1978  2209.26  +  153.08  6.93  25.12  1308.30  +  217.02  16.59  17.52  7.21  22.07  15.21  22.80  Average  •'•Estimates of V  A  '  580.12  are a l l s i g n i f i c a n t at the 0.01 l e v e l .  o  - 42 For seed zone 1, h 1972 to 1978.  2  remains r e l a t i v e l y constant varying only s l i g h t l y from 2 There i s a decreasing trend f o r h f o r seed zone 2 from  1972 to 1975 which s t a b i l i z e s to change very l i t t l e i n the f i n a l three years tested.  Provenances in seed zone 3 also e x h i b i t a steady decrease  2 in h  continuing to 1976.  magnitude.  Estimates f o r 1977.and 1978 are increasing i n  H e r i t a b i l i t y estimates f o r 1972, 1973 and ,1974 are increasing  f o r the i n t e r i o r provenances, although values are decreasing in the l a t e r years.  H e r i t a b i l i t y declines in the e a r l y years according to the r e l a t i v e 2  increase of Vp versus Vp^p.  One explanation of the decline in h  disappearance of maternal e f f e c t s not indicated by the .regression  is a analysis  f o r maternal i n f l u e n c e . The r e l a t i v e l y high amount of a d d i t i v e genetic control indicated by 2 the h  values suggest that there are opportunities f o r s i g n i f i c a n t improve-  ment by s e l e c t i o n in provenance and progeny tests f o r D o u g l a s - f i r .  Never-  t h e l e s s , the differences i n genetic variance between regions f o r Vp and Vp^p and the r e s u l t i n g differences i n genetic parameters may cause varying responses to s e l e c t i o n between regions.  This p o s s i b i l i t y should be f u r t h e r  investigated by studying the w i t h i n provenance variance i n d i v i d u a l l y . B i r o t (1976) found s i g n i f i c a n t differences in genetic variance between trees w i t h i n provenance as well as between provenances.  Heritability  estimates f o r i n d i v i d u a l provenances at the Haney s i t e had very large standard errors due to small sample s i z e s . 2 h  Appendix IV gives estimates of  and standard errors f o r several i n d i v i d u a l provenances.  Although these  values have low r e l i a b i l i t y , many are s i m i l a r in magnitude to estimates derived from seed zone a n a l y s i s .  Therefore, emphasis was again placed on  pooled data for'provenances w i t h i n seed zones.  - 43 -  TABLE 10 H e r i t a b i l i t y ( h ) and Standard Error (S.E.) f o r Total Height Based on Seed Zones 2  Location Year  Seed Zone 1 h  2  ± S.E.  Seed Zone 2 h  2  ± S.E.  Seed Zone 3 h  2  ± S.E.  Interior h  2  ± S.E.  1972  0.36 ± 0.10  0.52 ± 0 . 1 1  0.42 ± 0 . 1 0  0.39 ± 0 . 1 0  1973.  0.36 ± 0.10  0.46 ± 0.11  0.37 ± 0.10  0.44 ± 0.10  1974  0.36 ± 0.10  0.40 ± 0.10  0.37 ± 0 . 1 0  0.45 ± 0 . 1 1 -  1975  0.42 ± 0.10  0.35 ± 0.10  0.33 ± 0.09  0.38 ± 0.10  1976  0.40 ± 0.10  0.36 ± 0.10  0.30 ± 0.09  0.33 ± 0.09  1977  0.34 ± 0.10  0.38 ± 0.10  0.34  0.09  0.31  0.09  1978  0.33 ± 0.10  0.37 ± 0.10  0.39  0.10  0.28  0.09  - 44  The magnitude of h  2  -  values c a l c u l a t e d f o r d i f f e r e n t regions are in  agreement with B i r o t (1976) but are greater than those of Campbell (1972) 2 and Orr-Ewing and Yeh (1978).  Campbell estimated h  of j u v e n i l e Douglas-  f i r trees between 0.08 and 0.22 f o r f i r s t and second year height.  These  low estimates were mainly due to high family x l o c a t i o n i n t e r a c t i o n .  The  p o t e n t i a l f o r genetic improvement in seedling height growth nonetheless appeared good because of considerable genetic variance (high dominance genetic variance).  Orr-Ewing and Yeh also suggested that growth was under 2 weak a d d i t i v e genetic control obtaining low estimates f o r h . The h e r i t a b i l i t i e s they presented had low r e l i a b i l i t y because of the unbalanced 2 design and small s i z e of the experiments.  Standard errors f o r h  based on  seed zones at the Haney s i t e are consistent at approximately 0.10 f o r a l l years and seed zones giving a high degree of r e l i a b i l i t y to these estimates The merits of each of the s e l e c t i o n systems and combinations recommend ed according to the r e l a t i v e s i z e of the components of variance can be com2 pared on the basis of expected genetic gains using h  2 estimates.  The h  values c a l c u l a t e d here are based on family s e l e c t i o n . Another concept to be discussed here i s that i t may be more advantageous to c a l c u l a t e genetic gain on a per unit time basis.'  This was suggested  by Namkoong et al_. (1966) since the time to obtain gains may be a c r i t i c a l f a c t o r i n - t r e e improvement programs.  Not only i s the i n t e r v a l between the  i n i t i a l s e l e c t i o n and the time when the next generation i s a v a i l a b l e f o r s e l e c t i o n important, but the a p p l i c a t i o n of s e l e c t i o n w i l l act to change gene frequencies that in turn change estimates of genetic parameters u t i l ized in the gain p r e d i c t i o n , making gain estimates on the basis of i n i t i a l measurements inaccurate.  Although s e l e c t i o n has not been applied yet in  - 45 the I.U.F.R.O. experiment, Namkoong's idea should be considered in the future. 4.  Juvenile x Mature C o r r e l a t i o n Increases in simple c o r r e l a t i o n c o e f f i c i e n t s between t o t a l heights at  various ages are i n v e r s e l y associated with the number of years between measurements  (Tables 11, 12, 13 and 14).  The R  values are s i m i l a r f o r a l l  r e l a t i o n s h i p s i n a l l regions and are a l l s i g n i f i c a n t at the 99% l e v e l . The t o t a l height in 1972 and 1973 accounts f o r almost h a l f of the v a r i a t i o n in 1977 and 1978 t o t a l height.  Between 58% and 63% of the v a r i a t i o n  in 1978 t o t a l height can be explained by v a r i a t i o n in 1974 t o t a l  height.  V a r i a t i o n in 1975 t o t a l height explains approximately 75% of the v a r i a t i o n in 1978 t o t a l height. i s over 0.80.  The c o r r e l a t i o n f o r 1976 and 1977 versus 1978 t o t a l  height  In general, r e s u l t s from height measurements at e a r l y ages  projected forward four to f i v e years can p r e d i c t over 50% of height growth v a r i a t i o n i n l a t e r years.  The strong c o r r e l a t i o n between heights at very  e a r l y ages and l a t e r years indicates that s e l e c t i o n can be made at e a r l y ages to p r e d i c t l a t e r performance with minimal r i s k of losing good i n d i v i d u a l s . Measurements should be continued to determine i f the high r e l a t i o n s h i p of the e a r l y j u v e n i l e to l a t e r j u v e n i l e growth and growth in l a t e r age classes  is  perpetuated. The simple c o r r e l a t i o n s between progeny annual growth (P.A.G.) and t o t a l height are s i g n i f i c a n t at a l l ages with the c o r r e l a t i o n c o e f f i c i e n t 2 increasing throughout the t e s t period.  However, the R  f o r greater than 50% of the v a r i a t i o n u n t i l 1975.  does not account  It i s , therefore,  - 46 suggested to wait u n t i l at l e a s t age f i v e or l a t e r to make s e l e c t i o n s . Nursery e f f e c t s are not considered s i g n i f i c a n t in t h i s t e s t since a l l the trees received s i m i l a r treatment before outplanting.  S i m i l a r i t y in genetic  parameter estimates f o r a l l years also indicates that e a r l y r e s u l t s do not vary s i g n i f i c a n t l y from l a t e r ones. Results from the ranking of provenances according to mean 1975 and 1978 t o t a l height f o r a l l provenances are presented in Appendix V.  These  r e i t e r a t e the close r e l a t i o n s h i p between e a r l y and l a t e r height growth mentioned above showing that f o r a l l seed zones, there i s very l i t t l e change i n the ranking of the best 25% of the provenances i n d i c a t i n g good r e l i a b i l i t y in s e l e c t i o n of the best provenances at age f i v e .  The l a r g e s t  deviation in performance between 1975 and 1978 occurs in seed zone 2, but i t should be noted that a l l of the top seven provenances i n seed zone 2 are greater than 350 centimeters i n t o t a l 1978 height.  The performance of the  remaining provenances also does not change s i g n i f i c a n t l y so the removal of the poorest trees at age f i v e may be f e a s i b l e as w e l l .  Early evaluation  w i l l allow f o r t e s t i n g of a greater number of f a m i l i e s or provenances which w i l l lead to a higher s e l e c t i o n i n t e n s i t y .  Selection i n t e n s i t y i s another  important f a c t o r i n the determination of genetic gain.  5.  S e l e c t i o n of the Best Provenances and Progenies f o r the Test S i t e The f i n a l o b j e c t i v e of t h i s study i s to s e l e c t . t h e best provenances and  progenies on the basis of performance of t o t a l height growth.  From the  information presented in the beginning of the Results and Discussion s e c t i o n , provenances from seed zones 2 and 3 (Washington sources Matlock (52 and 53), Naselle (67) and Oregon sources Vernonia (83), Waldport (87),  - 47 TABLE 11 Juvenile x Mature C o r r e l a t i o n : Seed Zone 1 Variables  Variables R  Dependent  Independent  72 Tot. Ht. vs vs vs vs vs vs vs  73 Tot. 74 Tot. 75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.2  Ht. Ht. Ht. Ht. Ht. Ht.  0.82 0.70 0.54 0.47 0.40 0.31 0.16  73 Tot. Ht. vs vs vs vs vs vs  74 Tot. 75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.  Ht. Ht. Ht. Ht. Ht.  0.91 0.71 0.62 0.54 0.41 0.22  Dependent  2  1  Independent  R  2  74 Tot. Ht. vs vs vs vs vs  75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.  Ht. Ht. Ht. Ht.  0.88 0.81 0.72 0.58 0.35  75 Tot. Ht. vs vs vs vs  76 Tot. Ht. 77 Tot. Ht. 78 Tot.- Ht. P.A.G.  0.95 0.87 0.72 0.52  76 Tot. Ht. vs 77 Tot. Ht. vs 78 Tot. Ht. vs P.A.G.  0.95 0.83 0.67  77 Tot. Ht. vs 78 Tot. Ht. vs P.A.G.  0.91 0.81  78 Tot. Ht. vs P.A.G.  0.90  A l l of the simple c o r r e l a t i o n s are s i g n i f i c a n t at the 0.01 l e v e l of testing. P.A.G. = progeny annual growth c a l c u l a t e d as (75 height increment + 76 height increment + 77 height increment + 78 height increment)/4  - 48 TABLE 12 Juvenile x Mature C o r r e l a t i o n : Seed Zone 2  Variables Dependent  Independent  Variables  2 R  72 Tot. Ht. vs vs vs vs vs vs vs  73 Tot. 74 Tot. 75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.2  Ht. Ht. Ht. Ht. Ht. Ht.  0.80 0.67 0.49 0.40 0.34 0.29 0.11  73 Tot. Ht. vs vs vs vs vs vs  74 Tot. 75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.  Ht. Ht. Ht. Ht. Ht.  0.90 0.68 0.57 0.49 0.41 0.18  Dependent 1  Independent  R  ?  74 Tot. Ht. vs vs vs vs vs  75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.  Ht. Ht. Ht. Ht.  0.86 0.76 0.67 0.58 0.31  75 Tot. Ht. vs vs vs vs  76 Tot. Ht. 77 Tot. Ht. 78 Tot. Ht. P.A.G.  0.94 0.87 0.77 0.53  76 Tot. Ht. vs 77 Tot. Ht. vs 78 Tot. Ht. vs P.A.G.  0.96 0.86 0.69  77 Tot. Ht. vs 78 Tot. Ht. vs P.A.G.  0.93 0.82  78 Tot. Ht. vs P.A.G.  0.89  *A11 of the simple c o r r e l a t i o n s are s i g n i f i c a n t at the 0.01 l e v e l of testing. 2 P.A.G. •= progeny annual growth.  - 49 TABLE 13 Juvenile x Mature C o r r e l a t i o n : Seed Zone 3  Variables Dependent  Variables  Independent  Dependent  72 Tot. Ht. vs vs vs vs vs vs vs  73 Tot. 74 Tot. 75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.2  Ht. Ht. Ht. Ht. Ht. Ht.  0.81 0.72 0.58 0.51 0.45 0.38 0.20  73 Tot. Ht. vs vs vs vs vs vs  74 Tot. 75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.  Ht. Ht. Ht. Ht. Ht.  0.92 0.74 0.66 0.58 0.49 0.25  1  . Independent  74 Tot. Ht. vs .75 Tot. vs 76 Tot. vs 77 Tot. vs 78 Tot. vs P.A.G. 75 Tot. Ht. vs vs vs vs  Ht. Ht. Ht. Ht.  76 Tot. Ht. 77 Tot. Ht. 78 Tot. Ht. P.A.G. Tot.Ht  0.87 0.79 0.71 0.61 0.36 0.94 0.87 0.75 0.52  76 Tot. Ht. vs 77 Tot. Ht. vs 78 Tot. Ht. vs P.A.G.  0.96 0.85 0.68  77 Tot. Ht. vs 78 Tot. Ht. vs P.A.G.  0.93 0.82  78 Tot. Ht. vs P.A.G.  0.89  A l l of the simple c o r r e l a t i o n s are s i g n i f i c a n t at the 0.01 l e v e l of testing. P.A.G. = progeny annual growth.  - 50 TABLE 14 Juvenile x Mature C o r r e l a t i o n : I n t e r i o r Provenances  Variables Dependent  Variables  Independent  Dependent  72 Tot. Ht. vs vs vs vs vs vs vs  73 Tot. 74 Tot. 75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.2  Ht. Ht. Ht. Ht. Ht. Ht.  0.83 0.73 0.57 0.51 0.47 0.42 0.26  73 Tot. Ht. vs vs vs vs vs vs  74 Tot. 75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.  Ht. Ht. Ht. Ht. Ht.  0.92 0.71 0.64 0.58 0.51 0.30  1  Independent  74 Tot. Ht. vs vs vs vs vs  75 Tot. 76 Tot. 77 Tot. 78 Tot. P.A.G.  Ht. Ht. Ht. Ht.  0.84 0.78 0.71 0.63 0.40  75 Tot. Ht. vs vs vs vs  76 Tot. Ht. 77 Tot. Ht. 78 Tot. Ht. P.A.G.  0.96 0.88 0.75 0.56  76 Tot. Ht. vs 77 Tot. Ht. vs 78 Tot. Ht. vs P.A.G.  0.96 0.86 0.71  77 Tot. Ht. vs 78 Tot. Ht. vs P.A.G.  0.94 0.85  78 Tot. Ht. vs P.A.G.  0.92  A l l of the simple c o r r e l a t i o n s are s i g n i f i c a n t at the 0.01 l e v e l of testing. P.A.G. = progeny annual growth.  - 51 Coquille (89) and Brookings Squamish (25) and  (91) in seed zone 2; B r i t i s h Columbia sources  C h i l l i w a c k (27) and Washington sources Perry Creek (42)  and Gold Bar (61) in seed zone 3) produce the t a l l e s t trees at the Haney site.  Some provenances from seed zone 1 also e x h i b i t e x c e l l e n t growth  (Duncan (32), B r i t i s h Columbia and M i l l C i t y (96), Oregon). 2 from the information on V^, h  In a d d i t i o n ,  and j u v e n i l e x mature c o r r e l a t i o n , i t i s  evident that i n d i v i d u a l and family s e l e c t i o n in the provenances of the best zones i n 1975 w i l l y i e l d s i g n i f i c a n t improvement i n t o t a l height growth. Applying these r e s u l t s to a Douglas-fir tree improvement program i n south coastal B r i t i s h Columbia, the best provenances in seed zones 2 and 3 would be selected because of t h e i r s i g n i f i c a n t l y greater rate of height growth.  S p e c i f i c i n d i v i d u a l f a m i l i e s or trees may be selected below the  top provenances i n the best zones and exceptional provenances in seed zone 1 would be included.  S e l e c t i o n of some provenances in other seed zones may  also be desireable to increase the genetic base of the r e s u l t i n g breeding program and to provide the material f o r i n t e r r a c i a l crosses to f u r t h e r expand the a v a i l a b l e genetic v a r i a t i o n .  The number of provenances selected  would depend on the s p e c i f i c program objectives of the p a r t i c u l a r agency involved. The response to s e l e c t i o n , that i s , the genetic gain (R) at the Haney 2 s i t e was estimated from the formula R = i V i s the i n t e n s i t y of s e l e c t i o n and V (Appendix VI). chosen.  p  p  h  (Falconer, 1960) where i  i s the phenotypic standard deviation  A s e l e c t i o n i n t e n s i t y of only one in four i n d i v i d u a l s was  Assuming a normal population, the t h e o r e t i c a l value f o r i  25% of the best i n d i v i d u a l s i s 1.23 (Falconer, 1960).  saving  Estimates of genetic  gains f o r a tree improvement program of phenotypic s e l e c t i o n and e s t a b l i s h ment of clonal orchards without progeny t e s t i n g (Squi 1 lace et al_., 1966)  - 52 are given in Table 15.  Results using the r e l a t i v e l y low s e l e c t i o n i n t e n s i t y  of 25% i n d i c a t e genetic gains from 17.90% in 1972 to 10.96% in 1978 f o r a l l regions. The genetic gain estimates suggest that appreciable improvement in growth rate of Douglas-fir i s possible by merely s e l e c t i n g from the top i n d i v i d u a l s from any provenance.  However, the r e s u l t s of the seed zone and provenance  performance show that even greater gains can be achieved by s e l e c t i n g the best i n d i v i d u a l s from the best provenances in the best seed zones. For example, comparing the mean 1978 t o t a l height of provenance 53 (Matlock) at 387.7 centimetres to the mean f o r the whole p l a n t a t i o n at 297.2 c e n t i metres (Tables 4 and 5) an increase in t o t a l height growth of almost 33% i s suggested.  I f the best i n d i v i d u a l w i t h i n provenance 53 is also selected  (1978 t o t a l height of 580 centimetres), a f u r t h e r gain of 70% above the plantation mean is i n d i c a t e d .  TABLE 15  Year  Seed Zone 1 in cm.  % of pop. mean  Estimates of Genetic Gain f o r Height Growth f o r Seed Zones 1, 2 and 3 and I n t e r i o r Provenances  Seed Zone 2 i n cm.  % of pop. mean  Seed Zone 3 i n cm.  % of pop. mean  Interior in cm.  1972  4.74  15.72  8.61  23.10  5.75  16.96  3.69  1973  8.55  15.14  13.83  18.89  9.16  13.81  1974  12.53  14.64  17.08  15.09  13.04  1975  19.02  15.74  19.06  11.99  1976  23.82  14.50  25.69  1977  25.85  11.62  1978  30.93  10.88  % of pop. mean  Average % of Pop. Mean  15.82  17.90  8.25  18.75  16.65  12.99  12.65  18.83  15.39  15.26  10.80  14.94  15.27  13.45  11.88  18.02  9.40  17.15  12.83  12.15  34.09  11.84  26.02  10.15  20.88  11.73  11.34  40.24  11.22  36.27  11.31  23.51  10.41  10.96  ;  co  - 54 -  SUMMARY  The genetic v a r i a t i o n i n height growth of j u v e n i l e Douglas-fir trees i n the I.U.F.R.O. provenance-progeny t e s t at Haney varied between seed zones . in the r e l a t i v e sizes of V greater than Vpy  p  p  and Vpy . p  For two of the four regions, V  p  was  by a magnitude of f i v e (seed zone 1 and i n t e r i o r ) and  d i f f e r e d by a two-fold margin f o r seed zone 3. shown f o r seed zone 2 where V  p  The reverse r e l a t i o n s h i p was  was less than Vpy  p  f o r a l l years.  The con-  c l u s i o n was made that f o r provenances most adapted to the Haney s i t e , Vpy and V  p  are the dominant contributors to the phenotypic v a r i a t i o n and that  f o r less adapted provenances, V than Vp^ . p  p  i s of greater importance becoming greater  By f a r the greatest component of variance was due to the pooled  e r r o r term V  (which includes the v a r i a t i o n of i n d i v i d u a l trees w i t h i n  f a m i l i e s ) accounting f o r 53% to 62% of the t o t a l phenotypic v a r i a t i o n . V a r i a t i o n due to V  p  and V g were the next l a r g e s t sources.  The components  p x  f o r family w i t h i n provenance c o n s i s t e n t l y accounted f o r 4% to 10% of the t o t a l v a r i a t i o n and the smallest component was due to Vg.  The o v e r a l l  r e s u l t s of the study of genetic v a r i a t i o n between the various  Douglas-fir  populations suggest that a combination of provenance, family w i t h i n provenance and i n d i v i d u a l tree s e l e c t i o n would y i e l d the greatest increase i n height growth.  S i g n i f i c a n t genotype x environmental i n t e r a c t i o n suggests  that s e l e c t i o n from provenances and f a m i l i e s f o r increased height growth f o r a l l locations may not be p o s s i b l e .  The r e l a t i v e s i z e of Vp g shows a x  diminishing trend i n d i c a t i n g that i t s e f f e c t may be decreasing in importance. Estimates f o r V^ were quite high ranging from 21.80% to 32.27% and accounted f o r an average of 24.73% of the v a r i a t i o n i n t o t a l height f o r a l l  - 55 regions combined.  The standard e r r o r of  i s r e l a t i v e l y low f o r a l l areas  i n d i c a t i n g r e l i a b i l i t y i n the r e s u l t s and consistent values f o r the r a t i o (S.E.  ( V ) / V ) 100) f o r each seed zone are observed.  The r e l a t i v e l y large 2 amount of a d d i t i v e genetic variance caused estimates of narrow sense h to 2 A  A  be moderately high, ranging between 0.28 and 0.52.  High values f o r h  in-  d i c a t e that there are opportunities f o r s i g n i f i c a n t improvement by s e l e c t i o n in Douglas-fir. The c o r r e l a t i o n analysis between t o t a l heights at various ages and progeny annual growth and t o t a l height f o r j u v e n i l e height growth in Douglas-fir suggest that r e l i a b l e s e l e c t i o n of the best and d e l e t i o n of the poorest provenances and f a m i l i e s may begin at age f i v e .  The r e s u l t s from the  ranking of provenances according to mean 1975 and 1978 t o t a l heights s t a n t i a t e t h i s conclusion.  sub-  Studies should be continued to determine i f the  r e l a t i o n s h i p of e a r l y j u v e n i l e to l a t e r j u v e n i l e height growth can be e x t r a polated to l a t e r age classes.  Early t e s t i n g i s advantageous since i t increases  the s e l e c t i o n i n t e n s i t y , which in turn increases the possible genetic gains through s e l e c t i o n and breeding. Reccommendations a r i s i n g from the analysis presented i n t h i s thesis relevant to a t r e e improvement program f o r D o u g l a s - f i r i n south coastal B r i t i s h Columbia would include s e l e c t i o n of the top provenances i n seed zones 2 and 3 and the best provenances from seed zone 1. B r i t i s h Columbia, Washington and Oregon sources.  These would include  Additional s e l e c t i o n s  from other provenances w i t h i n the selected seed zones and other seed zones are suggested to increase the genetic base and provide the p o t e n t i a l f o r i n t e r r a c i a l crosses. Using as an example a selected proportion of 25% of the best i n d i v i d u a l s at Haney, the figures obtained f o r genetic gain show that an increase  - 56 -  i n mean two to eight year t o t a l height from 17.90% to 10.96% i s p o s s i b l e . I f s e l e c t i o n was made in the best provenance in the best seed zone, an increase in t o t a l height growth of almost 33% i s suggested; s e l e c t i n g the best i n d i v i d u a l w i t h i n that provenance could give:.an a d d i t i o n a l increase of 70%.  Further gains can be derived i f higher s e l e c t i o n i n t e n s i t i e s are used.  - 57 LITERATURE CITED  Anderson, R.L. and Bancroft, T.A. 1952. S t a t i s t i c a l theory in research. 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IUFRO Congress, Sec. 22 - AG 22/24, Munchen, III: 672-718. Lush, J . L . 1949. H e r i t a b i l i t y of q u a n t i t a t i v e characters in farm animals. Proc. 8th Intern. Congr., Hereditas, Suppl. V o l . : 356-375.  - 59 Meier, R.J. and Goggans, J . F . 1977. H e r i t a b i l i t i e s of height, diameter and s p e c i f i c g r a v i t y of younq V i r g i n i a pine. Forest Science 23(4): 450-456. Meier, R.J. and Goggans, J . F . 1978. H e r i t a b i l i t i e s and c o r r e l a t i o n s of the c o r t i c a l monoterpenes of V i r g i n i a pine (Pinus v i r g i n i a n a M i l l . ) . S i l v a e Genetica 27 (2): 79-84. Namkoong, G. and Conkle, M.T. 1976. Time trends in genetic control of height growth in Ponderosa pine. Forest S c i . 22: 2-13. Namkoong, G., Snyder, E.B. and Stonecypher, R.W. 1966. H e r i t a b i l i t y and gain concepts f o r evaluating breeding systems such as seedling;orchards. S i l v a e Genetica 15(3): 76-84. Namkoong, G., Usanis, R.A. and S i l e n , . R.R. 1972. Age-related v a r i a t i o n in genetic control of height growth in D o u g l a s - f i r . Theor. and Appl. Genet. 42: 151-159.' Nanson, A. 1965. Contribution a I'etude de l a valeur des tests precoces. I. Experience i n t e r n a t i o n a l sur I ' o r i g i n e des graines d'epicea (.1938). Trav. Stat. Rech. Groenendaal, Ser. E., No. 1. 60 pp. Nanson, A. 1968. La valeur des t e s t s precoces dans l a s e l e c t i o n des arbres f o r e s t i e r s en p a r t i c u l i e r au point de vue de l a croissance. D i s s e r t a t i o n . Station de Recherches des Eaux et Forets, GroenendaalH o e i l a a r t . 242 pp. Nanson, A. 1970. Juvenile and c o r r e l a t e d t r a i t s e l e c t i o n and i t s e f f e c t on s e l e c t i o n programs. In papers presented at the second meeting of the Working Group on Quantitative Genetics, Section 22, IUFRO, Aug. 18-19, 1969, Raleigh, N.C.: 17-26. Orr-Ewing, A.L. and Yeh, F.C.H. 1978. Survival and growth t r a i t s of r a c i a l crosses with D o u g l a s - f i r . Res. Note No. 85, Province of B.C., M i n i s t r y of Forests. 44 pp. Owino, F. 1977. Genetic divergence in selected populations of L o b l o l l y pine. S i l v a e Genetica 26 (2-3): 64-66. P a t l a j , I.N. 2973. Geograficeskie Kultury sosny obyknoviennoj v Ukrainskoj SSR. Proc. IUFRO Symp. Genetics of Scots pine. Warsaw-Kornik: .1-26. Pirchner, F. 1969. Population genetics in animal breeding. and Co. 274 pp.  Freeman W.H.  Rink, G. and Thor, E. 1976. Variance components and gains i n volume growth in V i r g i n i a pine (Pinus v i r g i n i a n a M i l l . ) . S i l v a e Genetica ... . ;25(1): .17-22. Schmidt, W. 1963. Early t e s t s . F.A.O./FORGEN 63 2a/10, In World Consultation on Forest Genetics and Tree Improvement P r o c , V o l . 1, Rome: 10-20.  - 60 -  Sprague, G.F. 1966. 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R e l a t i o n o f t h i r t i e t h - y e a r to e a r l i e r dimensions southern p i n e s . F o r . S c i . 17: 200-209. Wehrnhahn, C F .  1979.'  Personal  communication.  of  8 February.  Yao, C. 1971. Geographic v a r i a t i o n i n seed w e i g h t , some cone s c a l e measurements and seed g e r m i n a t i o n o f D o u g l a s - f i r (Pseudotsuga menziesii (Mirb.) Franco). M.F. T h e s i s . U n i v e r s i t y o f B.C. 8.8 pp. Z o b e l , B. 1961. Inheritance G e n e t i c a 10(3): 65-70.  o f wood p r o p e r t i e s  in c o n i f e r s .  Silvae  Z o b e l , B . J . and Dorman, K.W. 1973. L o b l o l l y pine as an e x o t i c . G e n e t i c s Resources I n f o r m a t i o n , F . A . 0 . Rome: 3-15.  Forest  - .61 APPENDIX I Least-Squares Analysis of Variance Seed Zone 1 F Values Source of Variation  df  1972  1973  1974  1975  1976  1977  1978  Block  2  18.30*- 15.26* 12.38*15.82*  Provenance  9  2 6 . 7 5 * ' 43.48*' 53.10* 54.92*-59.28* 66.16* 70.53*  Block x Prov.  18  12.25*  Family/Prov.  70  2.35*  15.63*12.62* 2.36*  2.35*  15.90*24.85*;41.26*  8.83*  7.71*  6.65*  6.11*  2.49*  2.53*  2.27*  2.23*  Least-Squares Analysis of Variance Seed Zone 2 F Values Source of Variation  df  1972  1973  1974  1975  1976  1977  1978  Block  2  5..96* 17..40* 18.,42* 13.,37* 12..42* 11..90*  Provenance  9  4.,07*  6..0*  4..79*  5.,49*  Block x Prov.  18  8.,81*  9,.75*  9..10*  8.,96* 10,.37* 10..90*  9..59*  Family/Prov.  70  3.,13*  2,.84*  2..58*  2..37*  2,.39*  2,.42*  * P ^  0.01  7..10*  8..65* 10..58* 10..42*  2,.50*  APPENDIX I (continued) Least-Squares A n a l y s i s o f V a r i a n c e Seed Zone 3  F Values  Source o f Variation  df  Block  2  Provenance  9  Block x Prov. 18 Family/Prov.  70  1972  1973  1974  3.58**15.99* ' 14.05* 20.64*  1975  1976  4.49**  4.39**  1977  1978  1.79NS  0.48NS  28.18*  28.30*  25.49*  28.96*  30.63*  31.45*  18.75* 23.71*  22.25*  17.26*  16.48*  15.99*'  13.47*  2.45*  2.30*  2.15*'  2.31*  2.53*  2.68*  2.44*  Least-Squares A n a l y s i s o f V a r i a n c e I n t e r i o r Provenances F Values Source o f Variation  df  1972  1973  1974  1975  1976  1977  1978  Block  2  23.16*  27.34* . 42.81*  48.30*  46.18*  45.53*  50.67*  Provenance  9  51.81*  62.64*  52.69*  45.60*  43.91*-  50.59*  54.14*  Block x Prov. 18  4.68*  7.33*  6.47*  4.77*.  5.06*  5.59*  6.63*  Family/Prov.  2.49*  2.68*  2.70*  2.44*  2.21*  3.45*  2.03*  * P  ^0.01  **P < NS  70  0.05 not  significant  APPENDIX II Components of Variance: Seed Zone 1  Year V  V  P.  32.66 (18.78)  F/P 10.28 (5.91)  104.27 (59.97)  173.88  138.21 (21.42)  33.81 (5.24)  339.39 (52.60)  645.23  235.47 (16.97)  72.09 (5.20)  728.07 (52.46)  1387.75  V  1.76 (1.01)  Total Pheno. Variance  PxB  V  1972  24.92 (14.43)  1973  133.82 (20.74)  0.0  (0.0)  1974  352.02 (25.37)  0.0  (0.0)  1975  630.75 (26.97)  24.54 (1.05)  274.60 (11.74)  148.67 (6.36)  1206.48 (53.89)  2339.05  1976  1139.94 (29.04)  48.09 (0.34)  393.58 (10.03)  236.32 (6.02)  2107.43 (53.69)  3925.36  1977  2113.86 (31.73)  177.09 (2.66)  550.16 ( 8.06)  324.87 (4.88)  3495.31 (52.47)  6661.29  1978  3437.20 (32.67)  521.28 (4.95)  758.31 (7.21)  477.99 (4.54)  5326.65 (50.63)  10521.44  1  2  Percent of estimated phenotypic variance  Negative component estimate  APPENDIX II (continued) Components of Variance: Seed Zone 2  32 97 ( 15.01)  23.72 (10.85)  157.50 (72.08)  Total Pheno. Variance 218.51  Year V  1972  P  4.32 ( 1 . 9 8 )  V  1  Q.0  B  (o.or  V  PxB  V  F/P  1973  23.62 (3.16)  10.84 (1.43)  124 11 ( 16.42)  68.82 (9.12)  528/67 (69.92)  756.06  1974  36.74 (2.44)  27.06 (1.78)  235 31 ( 15.65)  121.29 (8.07)  1083.25 (72.04)  1503.64  1975  71.65 (2.94)  21.13 (0.87)  381 65 ( 15.67)  173.51 (7.13)  1787.36 (73.39)  2435.30  1976  209.57 (4.80)  16.85 (0.39)  769 75 ( 17.64)  302.30 (6.93)  3064.51 (70.24)  4362.98  1977  411.97 (5.87)  12.99 (0.19)  1276 36 ( 18.18)  510.19 (7.27)  4809.12 (68.50)  7020.63  1978  598.10 (5.95)  0.00 (0.0)  1636 80 ( 16.28)  713.54 (7.10)  7105.36 (70.67)  10053.81  Percent of estimated phenotypic variance  Negative component estimate  APPENDIX II (continued) Components of Variance: Seed. Zone 3.Provenances  Year  V  P  V  B  V  PxB  V  F/P  V ep  Total Rheno. Variance  52..05 (26.64)  13.08 (6.69)  111.03 (56.84)  195.35  0.0 (0.0)  220..60 (30.92)  37.04,(5.19)  367.74 (51.55)  763.40  179.17 (12.63)  0.0 (0.0)  418,.47 (29.50)  75.67 (5.33)  745.45 (52.54)  1418.77  1975  279.47 (12.42)  0.0 (0.0)  556,.84 (24.75)  117.72 (5.23)  1296.27 (57.60)  2250.30  1976  543.20 (14.19)  0.0 (0.0)  902,.20 (23.56)  177.85 (4.64)  2206.19 (57.61)  3829.44  1977  925.61 (14.92)  0.0 (0.0)  1404,.69 (22.64)  325.56 (5.25)  3547.45 (57.19)  6203.30  1978 1384.07 (15.73)  0.0 (0.0)  1700 .75 (19.33)  552.32 (6.28)  5162.56 (58.67)  8799.69  1972  19.20 ( 9 . 8 3 )  1973  88.02 (12.34)  1974  1  0.0 ( 0 . 0 )  2  Percent of estimated phenotypic variance  Negative component estimate  APPENDIX II (continued) Components of Variance: . Interior,Provenances  Year V  1972  P  25.05 (27.12)  V  1  B  V  PxB  V  F/P  V  e  P  Total Pheno. Variance  2.73 (2.96)  5.44 (5.89)  5.83 (6.31)  53.30 (57.72)  92.23  1973  117.75 (29.62)  11.47 (2.88)  36.27 (9.12)  25.55 (6.43)  206.57 (51.95)  397.60  1974  222.12 (25.76)  46.84 (5.43)  70.56 (8.18)  58.11 (6.74)  464.63 (53.89)  862.15  1975  381.25 (23.65)  111.62. (6.93)  96.78 (6.00)  98.05 (6.08)  924.26 (57.34)  . 1611.96  1976  650.33 (23.18)  186.96 (6.66)  184.66 (6.58)  145.03 (5.17) 1638.84 (58.41)  2805.82  1977  1267.18 (25.74)  306.17 (6.22)  352.16 (7.15)  235.17 (4.78) 2763.10 (56.12)  4923.78  1978  2129.03 (28.52)  0.00 ( 0 . 0 )  676.50 (9.06)  327.08 (4.38) 4332.47 (58.04)  7465.06  2  Percent of estimated phenotypic variance  Negative component estimate  - 67 APPENDIX III Regression A n a l y s i s f o r Maternal i n Seed Zone 1  Provenance No.  Effects  1972 R  Percent o f Variation Explained  1973 R  Percent o f Variation Explained  23  0.0112  i:12  0.0175  1.75  32  0.0209  2.09  0.0263,  2.63  55  0.0744  7.44  0.0611  6.11  90  0.0272  2.72  0.0341  3.41  95  0.0188  1.88  0.0081  0.81  99  0.0655  6.55  0.0631  6.31  96  0.0027  0.27  0.0025  0.25  117'  0.0002  0.02  0.0000  0.00  111  0.0003  0.03  0.0001  0.01  124  0.0194  1.94  0.0589  5.89  104  0.0114  1.14  0.0154  1.54  Average  2.29  2.61  - 68 APPENDIX III (continued) Regression Analysis f o r Maternal E f f e c t s in Seed Zone 2  Provenance No.  2 1972 R  Percent of Variation Explained  2 1973 R  Percent of Variation Explained  12  0.0121  1.21  0.0027  0.27  51  0.0516  5.16  0.0385  3.85  52  0.0419  4.19  0.0265  2.65  53  0.0241  2.41  0.0139  1.39  67  0.0124  1.24  0.0268  2.68  79  0.0021  0.21  0.0210  2.10  83  0.0000  0.00  0.0010  0.10  87  0.0543  5.43  0.0308  3.08  89  0.0075  0.75  0.0000  0.00  91  0.0179  1.79  0.0008  0.08  92  0.0091  0.91  0.0280  2.80  115  0.1461  14,61  0.1207  12.07  Average  3.14  4.16  -  69-  APPENDIX IV , H e r i t a b i l i t y ( l / ) and Standard E r r o r s ( S . E . ) f o r T o t a l Height Based on I n d i v i d u a l Provenances 0  1  Provenance No.  Year  42  +  1972  0.00  1973  0.00  1974  0.10  1975  0.26  +  1976  0.39  +  1977  0.43  +  1978  0.39  +  + +  52  0. 00  0.57  0. 00  0.38  0. 21  0.27  0. 29  0.21  0. 34  0.20  0. 36  0.12  0. 34  0.15  Heritabilities  + + + + + + +  55  0.,43  0.41  0.,36  0.37  0.,31  0.39  0.,28  0.41  0. 28  0.49  0.,24  0.55  0. 25  0.40  + + + + + + +  66  0.37  0.76  0.36  0.63  0.36  0.19  0.37  0.06  0.40  0.00  0.43  0.00  0.37  0.00  were c a l c u l a t e d a c c o r d i n g to the f o l l o w i n g  V  F  +  W  + V  ep  + + + + + + +  0.51 0.47 0.30 0.24 0.00 0.00 0.00  formula:  APPENDIX V Ranking According to Mean 1975 and 1978 Total Height: Seed Zone 1 Provenances  1978  lCVT  117  111  124  99  23  90  1975  104  111  117  124  99  90  23  •''Provenances ranked i n increasing height from l e f t to r i g h t , 2 represented to the r i g h t of the l i n e Approximately 25% of provenances  APPENDIX V (continued) Ranking According to Mean 1975 and 1978 Total Height: Seed Zone 2 Provenances  1978  115  1975  115  1  12  92  79  51  83  87  91  67!  12  92  79  51  87  89  53  83  Provenances rank i n increasing height from l e f t to r i g h t .  Approximately 25% of provenances represented to the r i g h t of the l i n e  APPENDIX V (continued) Ranking According to Mean 1975 and 1978 Total Height: Seed Zone 3 Provenances  1978  103  1975  103  1  86  113  112  60  40  76  101  29  73  86  113  112  60  101  76  97  73  29  Provenances ranked in increasing  Approximately 25% of provenances  eight from l e f t to r i g h t .  presented to the r i g h t of the l i n e .  APPENDIX V (continued) Ranking According to Mean 1975 and 1978 Total Height: I n t e r i o r Provenances  2 1978  l l  1975  11  1  18  66  10  46  28  64  77  6  66  18  46  10  77  64  28  80  80  CO  Provenances ranked i n increasing height from l e f t to r i g h t .  Approximately 25% of provenances represented to the r i g h t of the l i n e .  - 74 APPENDIX VI Mean (iii) and Standard Deviation (S.D.) of 1972 to 1978 Total Height f o r Provenances in Seed Zone 1, 2 and 3 and I n t e r i o r (cm.) Location Year  Seed Zone 1  Seed Zone 2  Seed Zone 3  Interior  iii  ±  S.D.  iii  ±  S.D.  iii  ±  S.D.  iii  ±  S.D.  1972  30.15  ±  10.70  37.27  ±  13.46  33.91  ±  11.14  23.33  ±  7.69  1973  56.49  ±  19.32  73.22  ±  24.44  66.32  ±  20.12  44.01  ±  15.24  1974  85.57  ± 28.29  113.18  ±  34.71  100.37  ±  28.66  67.19  ±  22.86  1975  120.87  ±  36.81  158.91  ±  44.28  141.36  ±  37.60  97.81  ±  31.97  1976  164.31  ±  48.41  216.17  ±  58.02  191.64  ±48.83  133.62  ±  42.24  1977  222.47  ±  61.81  287.88  ±  72.93  256.26  ±  62.23  177.96  ±  54.76  1978  284.37  ±  76.19  358.72  ±  88.42  320.79  ±  75.60  225.77  ±  68.26  

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