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Study of the needle and cone tissue of alpine fir (Abies lasiocarpa (Hook.) Nutt) 1966

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A STUDY OF THE NEEDLE AND CONE TISSUE OF ALPINE FIR ( ABIES LASIOCARPA ( HOOK.) MJTT). by KALMAN JOSEPH ROLLER FOREST ENGINEER, UNIVERSITY OF SOPRON , 1937 DOCTOR IN FORESTRY,ACADEMY OF SCIENCES, BUDAPEST, 1953 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN THE DEPARTMENT OF BIOLOGY AND BOTANY We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA May, 1966 In presenting t h i s t h e s i s i n p a r t i a l f u l f i l m e n t of the requirements for an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree that the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e for reference and study. I f u r t h e r agree that per- mission for extensive copying of t h i s t h e s i s for s c h o l a r l y purposes may be granted by the Head of my Department or by h i s representativeso I t i s understood that copying or p u b l i - c a t i o n of t h i s t h e s i s for 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 The U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8, Canada. i i ABSTRACT • Alpine f i r ( Abies lasiocarpa ( Hook ) Nutt. ) needles and cones were studied in natural stands to determine the v a r i a b i l - i t y of several traits such as needle length, width,stomatal die tance in a row line on the abaxial surface, difference between cone scale and bract lengths, seed-wing areas, etc. Additional histological characteristics of needle, cone scale and bract are presented,, Sixty-five stands, located on the West Coast of North America, were sampled. Continuous distribution of v a r i - able characteristics has not been found in the 65 stands. Correlation analysis shows that stomatal distance i s directly associated with precipitation.Therefore, stomatal frequency increases with higher precipitation.Analysis of v a r i - ance shows that the difference between cone scale and bract lengths i s highly significant between the different stands. These differences indicate actual variations in the cone collec tion. Varieties and clines are suggested to exist in the natural range of alpine fir„ i i i TABLE OF-CONTENTS Page TITLE PAGE . . . . . . . . . . ..' i ABSTRACT . . . . i i TABLE OF CONTENTS i i i LIST OF TABLES . . iv LIST OF ILLUSTRATIONS v ACKNOWLEDGEMENT . . . . . . . . . . . . . v i i INTRODUCTION . . . . . . . . . . . . . 1 ABIES LASIOCARPA - BACKGROUND 4 METHODS, MATERIALS AND PROCEDURES . . . . 11 NEEDLES: Morphology : 14 Analysis of data on stomata . . .' 20 Histology . .. 28 Summary of observations 35 CONES: Field procedures, technique of preparation and measuring of cone particles 37 Evaluation and analysis of the difference between scale and bract length 41 Histology . . . . . 47 DISCUSSION AND CONCLUSIONS 56 SUMMARY • • • • • • • • •• • 66 LITERATURE 68 TABLES . . . . . . . . 72 FIGURES 8 6 i v LIST OF TABLES Table 1 D i s t r i b u t i o n of measured data on alpine f i r on the Vest Coast. 2 Basic elements of s t a t i s t i c a l analysis on stomatal distance. 3 Correlation c o e f f i c i e n t s of stomatal distance. 4 Regression c o e f f i c i e n t s of cl i m a t i c factors, elimination of ( Y ), stomatal distance. 5 Anatomical characteristics i n needle cross sections of some selected alpine f i r samples. 6 Geographical d i s t r i b u t i o n of a l p i n e - f i r samples collected and some char a c t e r i s t i c s of the resin canals i n t h e i r needles. 7 Measured data of scales and bracts i n the alpine f i r cones collected on the ̂ est Coast. 8 Analysed data on seeds and cones of alpine f i r . 9 Basic elements of s t a t i s t i c a l analysis on scale and bract. 10 Correlation c o e f f i c i e n t s of scale and bract. 11 ^-egression c o e f f i c i e n t s of the variables. Elimination of Y, the difference between the length of scale and bract. V LIST OF ILLUSTRATIONS Figure 1 D i s t r i b u t i o n of alpine f i r stands sampled on the West Coast. 2 Regression l i n e of p r e c i p i t a t i o n on stomatal distance. 3 Comparison between means of stomatal distances on the lower surface of needles. Diagram of Duncan's test. U Outlines of needle cross-sections. D i s t r i b u t i o n patterns of hypodermal c e l l s under the needle epidermis. 5 Shape and areas of seed wings sampled from various stands. 6 D i s t r i b u t i o n of vascular strands i n the cone-scale. 7 Comparison between the means of the cone scale and bract lengths. Diagram of Duncan's test. 8 Delicate cones. Pendleton ^ay, B.C. (36). 9 Husky cones. Mt. Idaho, New Denver, B. C. (57). 10 High-alpine type alpine f i r group. G l a c i e r National Park, Montana (3). 11 Sub-alpine type alpine f i r group. Hurricane ridge, Washing- ton (24.). 12 Yellow tinged alpine f i r stand on la v a bed. LAVA CAMP Lake, ^regon (52). 13 Stand., located 'at Willson Creek,New Denver B.C. ( 57 ). Bracts of some selected samples. The code numbers of samples, as f o r Table 7 are from above l e f t to r i g h t : igure 14 (cont'd) 1st row: 5, 37, 60, 7, 26 2nd row: 20, 12, 25,.17, 38, 24 3rd row: 24, 38, 3, 3, 22, 2 4-th row: 31, 31, 19, 9, 31 15 Smallest seeds and wings. Aleza Lake,; B.C. • (31) 16 Largest seeds without .ving^Will'son Cr., New Denver BC. (31). 17 Stracture of sunken stomata. 18 Stomatal pattern on both surfaces of the alpine f ir needle. 19 Sunken stomata, well defined hypodermal layer under the epi- dermis and extensive tracheid. tissue in the vascular bundle (at the right corner). Garibaldi Lake trail, B.C.,(l). 20 Cross-section of bract. Immature cone. 21 Cross-section of welI4developed needle. 22 Cross-section of scale. Immature cone. 23-26 Different location, size and structure of resin canals in needle cross sections of alpine fir . Fig. 23: Alaska Hwy., Mi. 4.8O, B.C., (41) j Fig. 24.: Hurricane Rdg., Wash., (24); Fig. 25: Graves Mt., Wash., (54-); F i g « 26: Seedling needle, P.F.S.S. Chalk River, Ont. 27-31 Vascular bundles. Cross-section. Fig. 27: Kicking Horse Pass, B.C.,(37) Fig. 28: Alaska Hwy., Mi. 735, B.C.,(5$ Fig. 29: Garibaldi Lake Trail, B.C.,(l) Fig. 30: Wallowa Mt., East Peaks, Ore., (65>, Fig. 31: Alaska Hwy., Mi. 48O, B.C.,(4$ 32 Vascular bundle in longitudinal section. Prince George. 33 ' Sub-alpine type bark with well-developed 1 en tic els. Moraine Lake, Alt a. v i i ACKNOWLEDGEMENT ' I wish, to express my appreciation to the National Research Council of Canada and the Canada Department of For- estry for granting me financial aid during the period in which this study was conducted„ Thanks are due to my advisors, Dr. A.H. Hutchinson and Dr. T.M.C. Taylor professors of Botany at the University of British Columbia for much assistance and suggestions on many phases of the study. I owe a special debt of gratitude to Mr. H . Sweet for helping to prepare microslides in I960 and 1961.Appreci- ation i s also expressed to many persons, too numerous to men- tion, for much help in locating study areas, f o r provided trans- portation and for direct assistance in the f i e l d and with office records. INTRODUCTION. In November 1958, the Forest Genetics Re- search Foundation,Berkeley,California,published i t s f i r s t survey of forest genetics research on the West-Coast.None of the ninety-four current projects was concerned with the Genus Abies.However.increasing demands from industry for good quality true f i r s for lumber and other purposes empha- size the need for such research. In B r i t i s h Columbia l i t t l e attention has been paid to the research of true f i r s . A.H.Hutchinson(1924) has done some work.; on the embryogeny of amabilis f i r ( A b i e s amabilis (Land) Forbes) and R.L.-Schmidt published a paper on the s i l v i c s and plant geography of the Genus Abies (1957). Moreover,some scattered data were available from several authors concerning the d i f f e r e n t species of Abies occuring i n B r i t i s h Columbia but a comprehensive study has not yet been published. For t h i s reason and i n response to the i n - creasing interest i n the use of this species f o r di f f e r e n t s i l v i c u l t u r a l and i n d u s t r i a l purposes,Dr.Hutchinson sug- gested that the writer undertake a project on alpine f i r (Abies lasiocarpa (Hook.)Nutt.) There are four species of Abies i n B r i t i s h Columbia: amabilis or s i l v e r f i r , (Abies amabilis (Land) For- bes ) ; grand or white f i r , (Abies grandis Lindl.) j balsam f i r , ( Abies balsamea ( L} Miller) and alpine f i r , ( A b i e s lasiocarpa ( Hook. )Nutt.). Commercially amabilis f i r i s the most important species.Alpine f i r i s less important but i t - 2 - has by f a r the g r e a t e s t a l t i t u d i n a l and g e o g r a p h i c a l d i s t r i - b u t i o n i n B r i t i s h C o lumbia.Since a m a b i l i s f i r had been s t u d i e d by H u t c h i n s o n , the p r e s e n t s t u d y was u n d e r t a k e n t o s t u d y t h e s p e c i e s o f second g r e a t e s t i m p o r t a n c e , t h e a l p i n e f i r * The a u t h o r began the problem i n the f a l l o f 1959 and c o n t i n u e d s t u d y i n g i t d u r i n g the f o l l o w i n g y e a r s . The f i e l d work was c a r r i e d out i n t h e summers o f I960 and 1961.However,since 1961, more s p e c i f i c o b j e c t i v e s , d e s i g n e d t o l e a r n as much as p o s s i b l e about the a l p i n e f i r i n the West Coast,extended the i n v e s t i g a t i o n t o Washington,Oregon and n o r t h e r n C a l i f o r n i a . The main purpose o f t h i s s t u d y was t o de- termine the e x t e n t o f the v a r i a t i o n i n a l p i n e f i r w i t h r e - s p e c t t o s e v e r a l h i s t o l o g i c a l and m o r p h o l o g i c a l c h a r a c t e r - i s t i c s and t o de v e l o p procedures f o r f u r t h e r s t u d i e s which might l e a d t o some r e s e a r c h i n t r e e improvement. O b v i o u s l y , i t would t a k e a team o f s p e c i a l i s t s t o o b t a i n a l l p o s s i b l e d a t a i n the morphology o f a l p i n e f i r , so the problem becomes a m a t t e r o f a r b i t r a r y s e l e c t i o n based on the d i f f e r e n t t r a i t s o b t a i n e d d u r i n g the p e r i o d o f f i e l d and l a b o r a t o r y e x a m i n a t i o n . i n the f i r s t t h r e e y e a r s . The f o l l o w i n g n e e d l e and cone c h a r a c t e r i s t i c s have been con- s i d e r e d : l e n g t h , w i d t h and arrangement o f n e e d l e ; a n a t o m i c a l s t r u c t u r e o f the needle,cone s c a l e and cone b r a c t ; d i s t a n c e between stomata i n a row l i n e on the lo w e r s u r f a c e o f the - 3 - n e e d l e ; d i f f e r e n c e between cone s c a l e and b r a c t l e n g t h , a n d s i z e o f the seed-wing. There were t h r e e p r i m a r y o b j e c t i v e s t o t h i s s t u d y : 1. t o determine the v a r i a b i l i t y o f needle and cone c h a r a c t e r i s t i c s f o r two m o r p h o l o g i c a l t r a i t s : d i s - t a nce between stomata i n a row l i n e and d i f f e r e n c e between cone s c a l e and b r a c t l e n g t h , 2. t o become f a m i l i a r w i t h o t h e r t r a i t s , 3. to a s c e r t a i n whether a l p i n e f i r i s e s s e n - t i a l l y one p o p u l a t i o n and whether i n i t s p h e n o t y p i c v a r i a t i o n s i t f o l l o w s c e r t a i n e n v i r o n m e n t a l g r a d i e n t s . - 4 - ABIES LASIOCARPA - BACKGROUND. Alpine f i r belongs to the genus Abies i n the family Pinaceae. Ten species are included i n the f l o r a of North America, eight of which are scattered through the f o r - ests of the West ( Harlow and Harrar, 1950) .. GEOGRAPHY: Alpine f i r i s endemic to the western coast of North America and to the Rocky Mountain areas from Arizona to Alaska as fa r north as 62° of l a t i t u d e . I t grows under various ecological conditions i n dif f e r e n t habitats where the p r e c i p i t a t i o n i s from 10 to 140 inches. I t exhibits a wide range of tolerance to l i g h t and occurs sometimes under the canopy of other species ( Engelmann spruce,mountain hemlock and Douglas f i r ) i n protected valleys or i n open stands and pure groups at timberline.Its a l t i t u d i n a l d i s - t r i b u t i o n i s from sea l e v e l to 3,000 feet i n Alaska; 2,000 to 7,000 feet i n B r i t i s h Columbia; 2,000 to 8,000 feet i n Washington and Oregon and 3,500 to 9,500 feet i n the Rocky Mountains (Harlow and Harrar , i 1950 ) .The d i s t r i b u t i o n of alpine f i r i s broken up by lowlands, and stands are thus isolated from each other i n some regions of i t s natural range.The segregation of stands may have brought about some inherent v a r i a t i o n of cha r a c t e r i s t i c s within the species during the geological eras. SITE REQUIREMENTS: Trees of largest dimension have "been found by the author on f a i r l y deep, loose podsolic s o i l at about 4000 feet elevation at Willson Creek,near New Denver B.C. and the poorest trees on bedrock covered by a thin mantle of s o i l at high elevations i n the Rocky Mountains.Trees of moderate size were sampled on a sandy beach at Bear Lake,north of Prince George B.C., on well-drained loamy s o i l around Babine Lake, B.C. and on g l a c i a l t i l l s on the Cascade Mountains and at Crater Lake,Oregon. According to Harlow and Harrar (1950):"This species i s r a r e l y ever found on heavy clays but not i n ~ frequently grows on s o i l s too wet f o r Engelmann spruce." This quotation indicates that the alpine f i r commonly occu- pies moist s o i l s and occurs often i n alpine meadows and along lakeshores at high elevations. Schmidt (1957) pointed out that, on Vancouver Island,alpine f i r i s most abundant on well-drained,shallow, rocky s o i l s . I n addition to growing on these s o i l s i n the northern coastal f o r e s t , i t grows also on s o i l s characterized by impeded drainage and with a standing water-table near the s o i l surface during the growing season. In summary: alpine f i r i s adapted to a wide vari e t y of s i t e s between elevations of 3,000 to 9»500 f e e t . However i n the North the extension of i t s range into higher a l t i t u d i n a l zones i s lower than i n the South.Under poor con- - 6 - ditions at high elevation i t may be a pioneer on rock outcrops, as for instance on the s t e r i l e lava bed at McKenzie Pass, Eastern Oregon. Its adaptability to d i f f e r e n t ecological conditions makes alpine f i r a very worthwhile species f o r s i l v i c u l t u r e .and emphasize i t s role i n the mixed forests of the spruce-balsam type regions i n the northern and southern Interi o r of B r i t i s h Columbia, which areas form one of the p r i n c i p a l timber reserves of the Province.- TAXONOMIC DESCRIPTION: Species of Abies are distinguished mainly by two c h a r a c t e r i s t i c s : ( a ) smooth twigs with c i r c u l a r needle scars ( b ) the erect disintegrating cones* [Native Trees Of Canada ( 1961 )T] Alpine f i r has s p i r a l l y arranged acicular leaves which appear to be two ranked on the twigs,those on lower branches crowded and directed upward; upper crown needles directed forwards,often appressed on leading shoots (Harlow and Harrar 1950 ).The length of needles varies between 2 to 4 c e n t i - metres.The sunken stomata are both dorsal and ventral.The r e s i n canals, usually two,are medial.The hypoderm i s highly variable, uniform or multiform; the thick wall of the endodermis and the double fibrovascular bundles are c h a r a c t e r i s t i c of t h i s f i r . According to P u l l i n g ( 1934) ,presence or absence - 7 - of stomata on a l l surfaces of the needles,number of stomata per square unit,contrast i n shape of cross sections of nee- dles,fused, s l i g h t l y separated or d i s t i n c l y separated nature of the two vascular strands - are a l l heritable,although the v a r i a b i l i t y of species of Abies i s very high i n t h e i r morpho- l o g i c a l and anatomical characters. Stover ( 1944 )finds that the size of needles varies from year to year. The mesophyll shows palisade-like c e l l s , and r e l a t i v e l y large a i r spaces.Epidermal c e l l s are thickwalled except those of the current, growing season;xylem and phloem are separated into two bands within the vascular bundle; there i s a difference i n the size of the r e s i n canals not only i n di f f e r e n t needles but also i n those from d i f f e r e n t habitats,being smaller on the more mesic habitat,, Laing ( 1956 ) warns that the form of needle t i p i s often an uncertain character and has to be used with d i s - cretion. The apex of needle may be acute,rounded or marginate. He also stated that the cross section of the needle i s impor- tant because often i t has to be decided whether the r e s i n canals, of which there are usually two, are at the edge of needle (mar- gin a l ) or some distance from the edge ( medial) » I t has been noted that the r e s i n canals i n the needles on non-coning shoots are sometimes i n a d i f f e r e n t position to those on coning-shoots. Alpine f i r i s monoecious and has separated staminate and ovulate cones on the same tree. The staminate cone i s dark indigo blue,the ovulate cone i s dark purple i n the i n i - t i a l stage of development. - 8 - Both the staminate and ovulate cone are borne on twigs of the - previous year.The f i r s t occurs on the lower branches i n the a x i l of the needles, the l a t t e r stands erect on the top branches of the crown«( Collingwood and Brush 1955) . The ovulate cone matures during a single grow- ing season.When mature i t i s from three to f i v e centimetres long, ( Pig.8 and 9 ) , c y l i n d r i c a l and purplish grey to nearly black. The cone frequently drips with s i l v e r y r e s i n . When ma- ture i t s seeds are released and the scales are shed. The cone axis remains on the tree for some seasons.Bracts are shorter than the scale.The bract i s mahogaay-red and has erose,round- ed shoulders and a long,black, spinelike t i p ( Harlow and Harrar 1950 ). Boivin ( 1959 ) emphasizes that there i s a con- stant r a t i o between scale length and bract length within cer- t a i n variations of alpine f i r . This r a t i o i s discontinuous between d i f f e r e n t variations and species.The bract/scale r a - t i o i s about 5/10 - 6/10 for the alpine f i r . Seeds are ivory brown,from four to s i x m i l l i - meters long, with a large lustrous purplish or v i o l e t - t i n g e d wing ( Sudworth 1908). Figure 15 and 16 . Winter buds are rounded,consisting of l i g h t orange-brown scales more or less covered with r e s i n 0 Twigs are f a i r l y stout,orange-brown,and when immature are covered with a fine rusty-red pubescence. The bark of the trunk i s usually grey,but some- times chalky white. The b l i s t e r - l i k e r e s i n pockets on the bark are c h a r a c t e r i s t i c of a l l the f i r s or "balsams". On large trees the bark i s l i t t l e broken, and usually narrow shallow cracks are present near the base of the trunk. VARIETIES AND HYBRIDS, Some color differences of the foliage are noted and described by several authors: Collingwood and Brush ( 1955), Sudworth (1908 ) , Duffield. ( 1962 ) .However color differences of the foliage are d i f f i c u l t to d i s t i n g u i s h since they vary with season, s i t e , health of trees and with the age of f o l i a g e . Figure 11,12 and 13. An alpine f i r hybrid was reported i n a personal communication by Heimburger to KLaehn and Winieski( 1962 ). He had observed a natural hybrid with balsam f i r i n northern Alberta. Harlow and Harrar ( 1950 ) l i s t a \ a r i e t y as cork- bark f i r , ( Abies arizonica ( Merr.) Lemm. ). This has thick,corky bark which i s free of r e s i n b l i s t e r s . Furthermore,in alpine f i r the cone scale are mostly wedgeshaped while i n corkbark f i r they are often halberdlike at the base. The corkbark f i r occurs i n the southern Rocky Mountains at elevations from 8,000 to 10,000 fe e t . Dallimore and Jackson ( 1961 ) mention another va- r i e t y of alpine f i r , ( Abies lasiocarpa var. compacta (Beissner ) Render ). This i s a dx-zarf form of compact habit occuring on poor rocky s o i l . The i r r e g u l a r arrangement of the foliage,the pointed needles of the terminal shoot,and the conspicuous s t o - matic l i n e s on the upper surface of the needle are i t s d i s t i n - g u i s h i n g f e a t u r e s . KLaehn and W i n i e s k i (1962 ) and C h i a s s o n ( I960, 1962 ) r e p o r t e d an easy c r o s s a b i l i t y f o r female balsam f i r and male a l p i n e f i r • Three s u c c e s s f u l a r t i f i c i a l p o l l i n a t i o n s were c a r r i e d out by Mergen e t a l . (1964 ) on female A b i e s p r p c e r a , i A. Koreana and A. s a c h a l i e n s i s w i t h a l p i n e f i r p o l l e n . The 1 r e s u l t i n g h y b r i d s have ' been s e t out f o r progeny t e s t i n the R e s e a r c h F o r e s t a t Y a l e U n i v e r s i t y . BIBLIOGRAPHY. . The f i r s t b i b l i o g r a p h y o f the t r u e f i r s was p u b l i s h e d i n 1963 by Lanner and Krugman and i n c l u d e s 373 a r t i c l e s . The a u t h o r of t h i s paper i n t e n d s t o b r i n g t h i s b i b l i o g r a p h y up t o date and a f u r t h e r e x p l o r a t i o n o f a r t i c l e s on the Genus A b i e s i s underway; about 1200 a r t i c l e s have been ag g r e g a t e d . - 11 - METHODS, MATERIALS AND PROCEDURES. . Most of the f i e l d work for the project was car- r i e d out i n the summers of I960 and 1961, although a few spec- imens were sampled i n the following years, when opportunities allowed the c o l l e c t i o n . Intervening periods were u t i l i z e d i n making laboratory and s t a t i s t i c a l analyses. A l l research and analyses were completed by June, 1964. It was considered that specimens should be c o l - lected for investigation i n some s p e c i f i c areas where alpine f i r stands were known to be present. In addition the c o l l e c t i o n of specimens was sometimes partly casual and p a r t l y determined by acce s s a b i l i t y of some areas during the f i e l d t r i p s . Sampling was most intensive i n the northern areas where i t was desirable to include a number of prominent o u t l i e r stands and others having overlapping areas with balsam f i r . The location of a l l sampled areas i s shown on the map i n Figure 1. The names and code number of the locations of the areas are given i n Table 1. Sampling procedures were c a r e f u l l y considered with a s t a t i s t i c i a n concerning the number of samples desirable from a certain area and appropriate s t a t i s t i c a l methods f o r analyses. I t was suggested that trees be selected f o r specimens at random from each stand as follows: trees to be sampled must be distributed equally i n the area and located i n uniform p o s i - t i o n as much as p o s s i b l e . l t means Uniform shade,density and mixture of other species.This way, the extraneous v a r i a b i l i t y - 12 - of needle c h a r a c t e r i s t i c s which could jeopardize the confidence of the s t a t i s t i c a l analyses,can be avoided. It was decided that ten trees per stand should be selected f o r sampling. These provide suffi c e n t data to c a l - culate mean values of needle characteristics for s t a t i s t i c a l analyses. The number of trees were limited by the available time and funds because of the wide range of alpine f i r from Oregon to the Yukon T e r r i t o r y . In a given stand the distance between the trees to be sampled was 150-250 yards, depending upon the extent of the sample-stand. However, the size of a sampling area was purposely l e f t f l e x i b l e to accommodate l o c a l differences i n stand conditions which could not be predetermined. . Where alpine f i r trees occupied a larger area at dif f e r e n t altitudes or various slopes and aspects,two or more sample-divisions were made and ten trees were sampled from each d i v i s i o n , e.g. Mt.Rainier and Olympic N.P. This method of sam- pl i n g was followed where both the mixed and pure stands formed a continuous forest area. At timber l i n e where the trees form small i s o l a t e d clumps on the alpine meadows,the method of c o l l e c t i o n was modi- f i e d and needle samples were taken from the scattered groups following the same isobar and so contur l i n e i n the middle be l t of the meadow. Sometimes, alpine f i r , mixed with other species was found i n long s t r i p s beside the highways and roads. Trees were - 13 - selected from the s t r i p about 200 yards from each other along the road. The t o t a l number of selected stands was 65 and 650 trees were sampled. Needle samples were collected from healthy , sexually mature trees. This means 20-40 years of age i n alpine f i r . Needles were picked from 3 year-old shoots of branches on the south side of the tree and at a height of 150 centimeters. It was found that a 3 year old needle has a well*-differentiated tissue complex i n i t s cross-section but i t can s t i l l be cut e a s i l y . The number of needles collected was limited by the 16 m i l l i - l i t e r capacity of v i a l s used as containers f o r t h e i r storage. PAA preservative was used for f i x i n g and storing materials i n v i a l s . - 14 - NEEDLES; MORPHOLOGY. The following measurements and observations have been made i n the laboratory from material that was c o l - lected i n the field„ A l l measurements were made on preserved needles selected at random from the v i a l s . LENGTH. The length of needles i s highly variable and, diffe r e n t lengths are commonly found on the same tree. Accord- ing to several authors ( Collingwood and Brush 1955fBoivin 1959 ) those of the lower branches are r e l a t i v e l y longer than those on the higher limbs and branches.Stover ( 1944) finds that the needles are longer on mesic and shorter on xeric habitats. The following needle lengths are reported by various authors: Boivin( 1959 ) 2.5-4.0 centimeter Collingwood and Brush ( 1955 ) 2.5 - 4.5 centimeter F u l l i n g (1934 ) . 2 . 0 centimeter and longer Harlow and Harrar ( 1950 ) . 3.0 - 4.0 centimeter Khuchel (1954 ) 3.0-4.0 centimeter Native trees of Canada, (1961) 2.5 - 4.4 centimeter The authors have not made any notes about the position of branches,except Harlow and Harrar (1950)who chose needles f o r measuring from s t e r i l e side branches. How- ever, the data of the quoted authors are very similar„Perhaps, they used the same source for the lengths of the needles. - 15 - The author of thi s paper measured the length of needles on s t e r i l e branches at the south side of the trees. The measurements were based upon the lengths of 10 needles, selected at random from thirty-nine stands' specimens. The lengths range from 0.9 to 4.3 . centimeters. The diagram below shows the frequency d i s t r i b u t i o n of needle lengths with a mean value ( X) of 2.13 centimeter and the standard deviation (s- ) 0.25 centimeter.This mean length i s shorter than the minimum, reported by the above authors. The explanation of the s i g n i f - icant differences between those of the quoted authors and of the authors of thi s paper might be the following: As i t was mentioned above, needle length i s strongly affected by habitat, aspect and the position of branches along the stem. In t h i s project, the aspect and the position and location of branches were i d e n t i c a l therefore i t could be assumed that the habitat was c h i e f l y responsible f o r the higher means and the larger range l i m i t s of the needle lengths. This theory i s supported by the fact that the stands sampled are very widely distributed i n t h e i r natural range and s o i l , climate and elevation also varied remarkably at the di f f e r e n t locations. Frequency d i s t r i b u t i o n of needle length FREQUENCY u ^ ° 1201 10 0 90 " 80-70 60-50 - io- 390 needles of 39 stands ro 15 20 1 30 50 LENGTH -j m m The change of needle lengths i s discontinuous i n the series of obser\£;ions on d i f f e r e n t habitats. Therefore the obtained data are not appropriate f o r s t a t i s t i c a l analysis of continuous variations and no conclusion can be drawn as to the cause of the needle length differences between the specimens. However, i t could be noted here that the longest needles Were found at Wallowa lake (43) , the shortest were collected at Aleza lake ( 28 ) and beside G-iscome Road (15 ) • WIDTH. Ten needles per stand were measured at three points of each needle.: ; M P '/4 The average value of these three measurements give the data f o r the computer.The results of computing are shown under X^ i n Table 1. In Table 2. under the column of X^ , the following values of needle-width are l i s t e d : average width of needles of 65 stands, X = 1.76 millimeter,standard deviation .17 millimeter and range: 1.35 - 2.20 millimeter. Table 3. presents the correlation c o e f f i c i e n t s with precip- i t a t i o n (X^ , altitude (X^) , and la t i t u d e (Xg).The c o e f f i c i e n t of altitude has the highest numerical value among the other factors analysed although i t does hot indicate a s i g n i f i c a n t c o r r e l a t i o n between the needle width and a l t i t u d e . - 17 - APEX. Four hundred needles,selected at random from the vials,were used for t e s t i n g the v a r i a b i l i t y of the apex form.. One h a l f of the needles were notched, one fourth were acute and one fourth were round.ed.This r a t i o of d i s t r i b u t i o n of the dif f e r e n t apex form i s casual,although i t shows that the apex i s usually notched on the sterile,shaded branches. Another observation was made on the form of needle apex.Needles were collected from d i f f e r e n t areas of the crown, i . e . upper,middle and lower areas. It was found that needles were acute or pointed on f e r t i l e branches from the upper crown at high elevation and rounded at low elevation. Trees, at lower elevations have pointed needles only on the leading shoots. A general statement could be drawn from the above observations: Needles are acute or pointed i n the upper crown and on the leading shoots while notched and rounded i n the lower crown. GROOVE AND RIDGE. The upper surface of the needle i s deeply grooved at the base and s l i g h t l y grooved at the apex. The lower surface i s ridged along the midrib.These char a c t e r i s t i c s are common on/ some true f i r s which have flattened needles i n cross section, e.g. balsam,silver and grand f i r s . I t was observed that groove and ridge appear more sharply on the needles at high elevation than at low elevation. - 18 - COLOR. Colour of the needles varies from dark green to yellow green. Those collected on lava beds or from high e l e - vations on bed-rock which had arisen from volcanic deposit, are yellowish green, Figure l o and 12 ; those of understorey trees, located i n well-drained,podzolic s o i l , a r e darker green, Figure 11 and 13. STOMATA. Two bands of stomata are on the abaxial surface of the needle consisting of 10-15 regularly distributed stomatal rows, Figure-18. One band of stomatal rows follows the midrib from the t i p to the base on the adaxial surface of the alpine f i r needle. The band forms a t h i n s t r i p e of two or three rows of stomata. Sometimes,this st r i p e disappears before i t reaches the base of needle. The d i s t r i b u t i o n of stomata i s i r r e g u l a r i n t h i s band,and the number of stomata i n a square unit i s ( highly v a r i a b l e . Stomata on both adaxial and abaxial surfaces i s a unique feature of the alpine f i r among the true f i r s of B r i t i s h Columbia. While species associated with alpine f i r i n the Province do not have stomata generally distributed on the adaxial surface of needles,they may have a few at the apex which do not extend beyond the upper.quarter of the needle. - 19 - T h i s f e a t u r e p r o v i d e s a good b a s i s f o r d i s t i n - g u i s h i n g the a l p i n e f i r from o t h e r t r u e f i r s o c c u r r i n g i n the P r o v i n c e , but i t cannot be used i n Washington,Oregon and C a l i - f o r n i a where s e v e r a l s p e c i e s o f t r u e f i r p o s s e s s n e e d l e s t h a t a r e s t o m a t i f e r o u s on b o t h s i d e s . The i r r e g u l a r i t y and h i g h v a r i a b i l i t y o f s t o m a t a l appearence on the a d a x i a l s u r f a c e o f a l p i n e f i r n e e d l e l e d t h e a u t h o r t o examine the s t o m a t a l d i s t a n c e on the a b a x i a l s u r f a c e . D i s t a n c e s between two n e i g h b o u r i n g stomata were measured on 20 n e e d l e s per s t a n d . Ten. measurements on each n e e d l e g i v e 200 d a t a per stand.Needles were s e l e c t e d randomly from each v i a l w hich r e p r e s e n t s a s t a n d . The measurements were made i n a one m i l l i m e t e r wide s t r i p e c r o s s i n g the c e n t r e p o i n t o f the upper h a l f o f the needle where the s t o m a t a l d i s t r i b u t i o n and the co n s t a n c y o f s t o m a t a l number proved t o be f a i r l y r e g u l a r <• A u n i t , e q u a l t o 0.2 m i l l i m e t e r was used f o r measuring the s t o m a t a l d i s t a n c e s . The a r r a y o f d a t a has been c o n s o l i d a t e d by g r o u p i n g measurements i n t o u n i t c l a s s e s . The c l a s s e s were summarized t o determine the t o t a l v a l u e o f Y, c a l l e d " d i s t a n c e i n d e x " . Two s e p a r a t e d measure- ments were made on each s t a n d : Y^ and , t o t e s t t h e v a r i a - b i l i t y o f s t o m a t a l d i s t a n c e s w i t h i n the s t a n d . The average o f and 1^ was used as d i s t a n c e i n d e x ( Y ) o f a sampled s t a n d f o r computing and a n a l y s i n g the d a t a i n d i f f e r e n t ways as p r e s e n t e d i n the f o l l o w i n g pages. - 2 0 , - Scoring system of 100 measured data Y 1 for Mount Rainier ( 25 ) : ' Units 3_ 4 5 §___7___ PreQuencjr 4 26 48__15_ 7 Total Y 1 12, 104 240 90 49 equal to 495 units. This distance index ( Y )shows more about the distribution of stomatal distances within a stand than the average value of measurements, and different stands can be distinguished from each other by the different values of"Y". • ANALYSES OF DATA ON ST0MATA. To analyse the interrelationships among the traits measured more data would have been needed. But there are sufficient measured data to find variations of the stoma- t a l distances in the different stands and show relationships between the distance indices (Y) and some independent varia- bles which affect the appearence of stomata on the needle surfaces. The mean (X )of the stomatal distances ( based on the data in Table 1 )is 0.104 millimeter in the 65 alpine f i r stands sampled and the average number of stomata 9.21 per millimeter in a single row line. The average number of rows i s 12 and so the average number of stomata is 115 i n a 1 m i l l i - meter wide stripe which lays on the lower needle surface perpen- dicular to the needle axis, Figure 18. . - 21 - The details of analysis of variance for the distance index (Y) are the following: Source Df Sum SQ . Mean SQ F Stand 64 2.64564E+05 . 4.13381E+03 38.429 Residual 65 6.99200E+03 1.07569E+02 Total 129 2.71556E+05 The F value exceeds.the value for F = 1.80 indicated in the body of the table for the 0.1 fo level with 64 an 65 degrees of freedom and shows that differences between the distance indices are highly significant. This result i n - dicates that there are significant differences between the sampled stands and the differences are not a matter of chance,'. While the analysis shows that the means of stomatal frequency in the investigated 65 stands are not equal, i t does not imply that a l l the stand. means differ significantly from each other. In order to determine which of the 65 stand means are significantly similar and-which are different, Duncan's multiple range test was used as employed by L i ( 1959) as the "least significant difference" between two stand means. In this case ISD (least Significant Difference ) at the 1 fo level is LSP.0,r. V i ̂  > ̂  w h i c h t = the tabulated 't* value from Duncan's Table ( 1955 )• f = number of observation for each stand that i s equal to 2,i.e. Y^ and Y^ are the observations. 2 S = the error mean square of the analysis of variance. - 22 - Since more than two stands are compared the quantity corresponding to the LSD i s called the shortest significant range,(SSR ) „ i The workings of the new multiple range test i s ' given in Figure 3. To perform this test the means of the 65 stands are necessary and these are then arranged according to their magnitudes. The standard error of the mean i s needed in computing the SSR. The standard-error of the means of Y i s : |s 2/n =107.6/2 =7.3 . The 1 ̂  significance level was chosen, the values of t were obtained from Duncan's Table.Each (r.oi) of the tabulated values was multiplied by the standard error to form the SSR,which are given in Figure 3 and in the table of Figure 3. In order to create groups within the series of stands sampled which are as close as possible to each other the smallest limit of each group was determined so that the corresponding SSR value was subtracted from the highest means (Y ) of the 65 stands one after the other. The means (Y), which f a l l into the intervals between the smallest limits and the corresponding means represent identified groups and the d i f - ference of stomatal frequency is not significant between the samples within each group. On the other hand, those of the means which f a l l outside an identified group dif f e r s i g n i f i - cantly from each other. For instance, the mean value of the 65th sample i s 646, after subtracting the SSR value 34,there - 23 - w i l l remain 612, which i s the smallest l i m i t of t h i s group and includes the 620,644 and 646 distance indices as i d e n t i f i e d samples. Figure 3 i l l u s t r a t e s the associated samples with the 65 stands, represented by horizontal lines.Any two • means and a l l the intervening means underlined do not d i f f e r s i g n i f i c a n t l y but the remaining means which are not under- scored by the same l i n e are s i g n i f i c a n t l y d i f f e r e n t . For ex- ample, the s t o m a t a l frequency of No£63, 64 and 65 d i f f e r s s i g n i f i c a n t l y from the frequencies of No.l to No. 62 stands. From Figure 3. i t can be seen that the difference i s highly s i g n i f i c a n t between the f i r s t and l a s t groups i n the series of stands tested. The significance of differences increases gradually from one side towards the other of Figure 3. The number of the samples i n the i d e n t i f i e d groups varies from two to twenty eight.The largest group of i d e n t i f i e d samples f a l l s into the middle part of the figure and includes populations which occur' at various habitats from the Yukon T e r r i t o r y toward South Oregon. The range of the d i s - tance indices of these stands extends from 480 to 514* This group represents the most frequent stomatal number on the ab- a x i a l surface of alpine f i r ' needle. Bach group underlined deviates from i t . To explain the occurrence of such variations ; further s t a t i s t i c a l analysis has been made. Several factors j were analysed by regression to test t h e i r r e l a t i o n to the stomatal frequency and d i s t r i b u t i o n . The basic elements of these - 24 - components are represented in Table 2. These are the only fac- tors available for analysis,other desirable data such as hu- midity,radiation,soil quality,et cetera are lacking. The com- ponents in Table 2, such as precipitation,elevation and l a t - itude generally characterize the ecological condition of the habitats where the samples were collected from. For example, the humidity of s o i l and air are controlled by the precipi- tation; the range of temperature and the radiation by the ele - vationjand latitude indicates some change of precipitation and temperature in British Columbia. It was assumed that one of the components men- tioned above may have some relation to the stomatal frequency represented by the stomatal distance ( Y ) , and the regression analysis promotes to detect i t . At f i r s t , the means ( X ), covariences ( S— ) and standard deviations ( S ) were calculated for each of the above mentioned components in order to obtain the correlation coef- ficients between them. The latter are given in Table 3» The correlation between the various factors at 1 fo level i s found to be as follows ( the meaning of the sym- bols used i s given in the legend of Table 1 . ) : X,correlate significantly with X2 X4and Y X 2 do X, X5 X6and Y X4 do X.and Y X5 do X2 X 6 X 6 do X£ X 5 Y do X, X2 X4 After the evaluation of correlation between - 25 - the distance index and tested factors i t was found that there are highly s i g n i f i c a n t negative c o r r e l a t i o n between Y and X, , X* and X„. The correlation of Y to X2 confirms the f i r s t a- nalysis of Duncan's multiple range test and proves that there are s i g n i f i c a n t differences between the distance indices of the various stands.The distance index i s i n inverse r a t i o to the. stomatal frequency (X,). On the other hand,this produces a positive correlation between stomatal frequency and precip- i t a t i o n both having negative c o e f f i c i e n t s i n the regression equation. Figure 2 shows that the stomatal frequency i s ' the highest and distance index the lowest i n those regions where the average p r e c i p i t a t i o n i s . h i g h over about 95 inches. For example: Code Prec. • No. location Inch ...Freĉ . Y 1 Garibaldi lake,B.C. 120 95 451 4 Black Tusk,B.C. 140 118 466 7 Arrowsmith Mt., V.I. . • • 100 127 468 14 Boulder lake,Olympic IT.P. ,Wash. 100 107 485 19 Mount Rainier,Wash. 120 91 488 20 Forbidden Plat., V.I. 140 92 488 Where p r e c i p i t a t i o n i s low( under 25 inches), low stomatal frequency was found on the needles with the ex- ception of the samples collected i n the Yukon T e r r i t o r y , lakes and swamps surround the mixed alpine f i r stands i n that region and keep the s o i l moist i n spite of the low p r e c i p i t a - t i o n i n t h i s region which probably accounts f o r the high stomatal frequency. Code Prec. No, location Inch Preq. Y 36 Pendelton Bay, B.C. 20 62 514 40 Pine Lake,Yukon . 8 110 526 43 Wallowa Lake,'Ore.' 20 90 529 44 Alaska Hwy.,Mi.852, Yukon 12 99 521 49 Mt. Bonaparte, Wash.' • 20 74 540 50 Divide Cr.,Glacier N.P.,Mont. 17 68 543 54 Graves Mt.,Wash. 17 75 561 The data from the analysis of variance can he treated to regression analysis by computing the regression equation. Y a + b, x, + b2 xs + b6 xs + \ xf + b, x5 + \ xs 66.86-0.367-K). 387+0.611+7.866+0.002-0 »025 The analysis was employed for the distance index as a t r a i t and the six variables presented in Table 1 and 2 . The result of the computation is given in Table 4 • In Table 4, the variables have been ranked in accordance with their impor- 2 tance, i.e. their R values. The sum of the sample regression coefficients , 2 R - 0.712, reflects a f a i r l y close association between the tested t r a i t (Y) and the five independent variables (precipitation, stomatal frequency, needle width, latitude and elevation). More- over, Table 4 shows that precipitation has the highest value of R, 57 of the total 71 percent,against the other variables, 14$ . This result was obtained after omitting the other variables one after the other during the computation. Used Snedecor's (1956) Table for the Vfo fiducial limit of R, i t was found that the value of R equal to 0 . 3 3 1 in the row of 63 degrees of freedom with 2 variables.Since the computed R value i s 0.570 that con- siderably exceeds 0.331, the association of stomatal distance with precipitation is considered to be highly significant. A diagram in Figure 2 expresses visually the relation of Y with precipitation.The dots are l i e i n g along the regression line ( 0-P ) in a relatively narrow band extending from the positive upper l e f t to lower right,not scattered widely over the whole f i e l d . This "significant relationship" means that as the precipitation varies,so the stomatal frequency varies directly. The variation pattern may be an example of a type of c l i n a l va- riation and indicates a non-adaptive value, i.e. a physiological character to which stomatal frequency i s linked. The analysis appears to prove that specific effects on stomatal differenti- ation can be obtained by different ecological conditions and that probably there are some possibilities of controlling stomatal distribution over the epidermal surface of the needle. To confirm this statement only one reference needs to be mentioned here. Zucker(1963) emphasized that envi- ronmental conditions existing at the time of stomatal differen- tiation in the embryonic leaf bud greatly affect this process. Availability of watertight intensity, and temperature have a l l been shown to be important factors.leaves of sunflower grown on a wet s o i l possess 20 times the number of stomata as do corre- sponding leaves from plants grown under very dry conditions. - 28 - EXAMINATION OP NEEDLE CROSS-SECTION . Preparation of cross-section for examination was adapted from the technique described by Johansen (1949) with some modification by Sweet ( I960 )and the author. The modification involves an acceleration of dehydration.Johansen ( p.132 )used a close series of dilutions of ethyl alcohol for dehydration. The author used only 5, 50, 70 and 95 % dilutions for two hours of each. Furthermore,the author used a mixture of Parowax,Fisher tissuement and bees wax, in( 48:48:4) percent ratio respectively for embedding the needles instead of rubber-Parowax used by Johansen. Usually i t is necessary to soften the embedded needles of alpine f i r for sectioning. So, the blocks were soaked in water for 48 hours or more. The transections and longitudinal sections were cut 10 microns thick using a rotary microtome. Sections were stained with safranin and fast green and mounted i n balsam. The cross-section of alpine f i r needles shows the following characteristics: EPIDERMIS has one layer. Cells are cube shaped with very thick c e l l walls. Both surfaces of the needle usually are heavily cutinized which i s a xeromorphic character,nevertheless needles i have been collected at Bear lake, Aleza lake, Summit lake and ' • • ' . I on Mount Idaho in British Columbia which show only a thin cuticle layer rarely visible under the misroscope.High moisture content of s o i l and air may be the reason for this lack of cuticle. STOMATA are deeply embedded in the epidermis even apparently into the hypodermal layers.There are some differences in shape of guard cells in the sections observed due to the degree of opening of stomata at the moment of collection. The arrangement of cells surrounding the openings i s always constant and the guard cells are overtopped by the sudsidiary cells^ Figure 17 • The term 'subsidiary c e l l ' is used in the sense of Esau (I960) . HYPODERMIS shows several patterns under the epidermis. There are discontinuous and continuous layer or layers. One can infer that well-developed hypodermis usually indicates dry habitat and severe climate. It is assumed that the role of thickened hypo - dermis i s to strengthen the needles against wind and snow pressure on upper branches at high elevation. On dry habitats,the continuous layer, or layers, of hypodermis control the rate of water-vapor loss and reduce the injurious effect of was found that needles collected from high mountains had continuous and often doubled layers while needles from lower elevation have discontinuous hypodermis, Figure 4« MESOPHYIL tissue is differentiated into palisade-like and spongy parenchyma cells i n the alpine f i r needle. The feature of 'palisade l i k e ' cells departs from the common palisade in form and size. Palisade cells are elongated and quadrilateral-shaped.Those of alpine f i r needle are shorter and more rounded and usually form one layer "between the hypodermis and the spongy parenchyma, Figure 25. Grand f i r has typical palisade among the true f i r s while palisade cells of the balsam f i r are usually similar to those of alpine f i r . Irregularly distributed parenchyma cells are located under the palisade layer. The parenchyma cells have particularly large numbers of chloroplast and numerous air spaces were found between them in the three year old needles sampled for measurements0 The air spaces are continued as inter- cellular spaces in the spongy parenchyma.The appearance of the intercellular spaces is usually irregular in the cross-section although in the longitudinal section they appear regularly arranged.Spongy parenchyma cells occupy a l l the abaxial region of the needle and include the resin canals medially located between the stele and epidermis sheath. It was observed during the investigation that the spongy parenchyma appears more compact in the young needles than in the three year old needles,Figure 26. ENDODERMIS SHEATH lines the different tissue elements of bundle or stele. The sheath i s made up from well differentiated large parenchymatous c e l l s . The nucleus and Casparian strips are not visible and the latter actually are lacking. The absence of Casparian strips i s not unusual in the secondary needles of true f i r s (Esau I960) • Sometimes the c e l l walls of the endodermis sheath are pitted like primary pit fi e l d s . Bundle sheath exten- sions occur at both the adaxial and the abaxial side of the bundle as a narrow channel whose cells are packed with starch. Within the endodermis sheath some thick-walled parenchyma cells line the transfusion tissue. These cells create a discontinuous sheath, very rarely continuous,around the inner part of the stele and are f i l l e d with starch. The complex of parenchyma and tracheid cells i n the bundle of needle i s called transfusion tissue. Large trans- fusion tissue is characteristic for the alpine f i r needle in contrast to the grand f i r and Douglas f i r which have limited transfusion tissue. The finding of a larger transfusion area i n - side the endodermis of the alpine f i r needle may .confirm Takeda's statement (1913 ) that the parenchyma cells of transfusion tissue are water storage cells occurring in xe^phytes. The same exten- sion of transfusion tissue was found by the author in the needle of red f i r and silver f i r collected from dry habitats, on the Sierra Mountains, California, and the Eastern slope of Mount Arrowsmith, Vancouver Island, respectively. In other samples, both red f i r and silver f i r have restricted transfusion tissue in needles collected from moist s o i l and climate. For example, needles of silver f i r collected in the western valleys of Manning Park, British Columbia, have small transfusion tissue. The vascular bundles are separated by parenchyma cells oriented obliquely, Figure 27 '.. As is usual in Pinaceae, the xylem i s adaxial and the phloem is abaxial and both are ar- - 32 - ranged i n 8-10 horizontal rows. Tracheids have h e l i c a l thicken- ings or bordered p i t s i n secondary walls.The h e l i c a l l y thickened tracheids are i n the adaxial side of xylem. The bordered p i t mem- branes have a torus i n a central position which i s c h a r a c t e r i s t i c of Abies species (Esau I960) 0 Both i n the xylem and i n the phloem there are some parenchyma c e l l s with c r y s t a l s . The c e l l s f i l l e d with crystals are usually i n connection with each other and form a v e r t i c a l column of c e l l s i n the .xylem and phloem. The albuminous c e l l s which are p a r a l l e l to t h i s column are e a s i l y seen beside the vascular bundle i n Figure 27. In another project, research has been done by the author concerning the location and tissue•arrangement of vascular bundles i n needle o f balsam,red, Shasta and white f i r . I t was found that distinction.can be made by these q u a l i t i e s f o r several sympatric species 0 For example, alpine f i r needle has more enlarged stele and more v e r t i c a l columns of xylem and phloem than balsam f i r ; transfusion tissue occupies more space i n alpine f i r than i n balsam f i r ; white f i r has one bundle i n the s t e l e , red f i r forms two bundles with a well-developed extension sheath between them. Detailed information on,the above i s to be published i n collaboration with Eugene Parker of Medford,Oregon, who i s working with the author on the d i s t i n c t i o n of some true f i r species occurring on the west coast of North America. RESIN CANAL. A paper Roller(1966a) concerning the number and position of r e s i n canal of alpine, balsam and Fraser f i r i s in press. The evidence is given there and so only the conclusions quoted here. Sixty needles were picked at random from the selected needle material of trees included i n Table 6, and the same number of needles from the seedlings planted in the nursery at Chalk River,Petawawa Forest Experiment Station. For examination, free-hand sections were made and li g h t l y stained in safranin. Features of resin canals in true f i r needles are taxonomic characters of well-established value.Their number and location have been regarded as only slightly less reliable char- acteristics, and many taxonomists have used these features with others to distinguish species in the genus Abies. A more detailed study, of this matter led the author to believe that the position and number of the canals seem to be highly variable in several stages of needle develop- ment. This research confirmed the thesis that resin canal position in alpine f i r and i t s sympatric species varies from peripheral to medial depending on the age of needle and tree, and in certain cases on the height of branch. In the seedlings of alpine f i r , resin canals were found in peripheral location, while needles of mature, trees were characterized by resin canals i n medial position. In some mature trees, needles picked from the lower crown had from none to two or more resin canals partly peripheral partly medial. Examples are shown in Figures 23 to 26. Table 6 presents the measured and evaluated data of some selected specimens of alpine fir.The analysis of these data resulted in the following: The difference in the 'height' of resin canals in the mesophyll is highly significant between the needles collected from adult trees located in various regions. A regression analysis of resin canal position on latitude and elevation shows very slight effect of latitude and no effect of elevation. The regression equation i s Y=7.23 + 0.0196 X,- 0.00000586 X* X, equal to latitude and X2= elevation. The observations and analyses described in the quoted paper confirm the thesis that the change of resin canal position with age, i.e. in seedling and adult stage, i s under genetic control, while the position of resin canals i s modified by environmental factors in adult age. SUMMARY OF OBSERVATIONS.ON ABIES LASIOCARPA NEEDLES. When in due course differences between the needles of various Abies species are examined, the following observations were made in this paper: The surface of both sides of the needle is heavily cutinized; the thickness of the cuticle is variable and thicker on the surface of the upper crown needles at high .elevation than low elevation. The epidermis has one layer. Sunken stomata are embedded into epidermis ( Figure 19) • The f i r s t collenchyma cells arise under the adaxial epidermis. The number of hypodermal layers increases with altitude and many layered hypodermal lines are located at the two margins of needle ( Figure 4 ) o The mesophyll tissue differentiates into palisade-like and spongy parenchyma cells.The palisade-like cells are not true palisade cells ( Figure 21). Endodermis i s usually broken at the adaxial side of stele(Figure 28). The inner bundle sheath gradually becomes visible during aging. A higher number of tracheids than parenchyma cells are in the transfusion tissue complex (Figures 28,29 ). Vascular bundles are poorly developed in the juvenile stage ( Fig.26 ) Two separated bundles are formed in the adult stage(Figure 25 ). In the seedling's needle, there are two peripheral resin canals joining the abaxial epidermal layer.While, in the needle of an a d u l t t r e e , t h e r e are two m e d i a l r e s i n c a n a l s embedded i n the m e s o p h y l l ( F i g u r e 21 ) . H i s t o l o g i c a l c h a r a c t e r i s t i c s o f t h i r t e e n samples are p r e s e n t e d i n Table 5. These samples were s e l e c t e d a t random from the groups formed i n F i g u r e 3.These groups e x h i b i t a s e r i e s o f s t o m a t a l d i s t a n c e s from t h e l o w e s t t o the h i g h e s t v a l u e o f Y. The arrangement o f samples i n T a b l e 5 .follows t h e arrangement of samples i n F i g u r e 3 i n o r d e r t o o b t a i n c o n s i s t e n t v a r i a t i o n s o f h i s t o l o g i c a l c h a r a c t e r i s t i c s a c c o r d i n g t o the i n c r e a s i n g v a l u e s o f t h e s t o m a t a l d i s t a n c e s . Continuous d i s t r i b u t i o n o f v a r i a b l e c h a r a c t e r i s t i c s has not y e t been found d u r i n g the i n v e s t i g a t i o n . O n l y one o b s e r v a t i o n can be noted h e r e , i . e . n e e d l e s from G a r i b a l d i Lake( 1) have h i g h s t o m a t a l f r e q u e n c y and an e n l a r g e d a r e a o f t r a c h e i d c e l l s i n the b u n d l e ( F i g u r e 29 )« C o n t r a r y t o t h i s however n e e d l e s from the Wallowa Mountains (65) e x h i b i t low s t o m a t a l f r e q u e n c y and an en- l a r g e d parenchymatous a r e a w i t h i n the endodermis s h e a t h ( F i g u r e 30 ). The former o c c u r s on moist h a b i t a t w i t h 120 i n c h e s a n n u a l p r e c i p i - t a t i o n , the l a t t e r on dryer habitat w i t h 35 i n c h e s a n n u a l p r e c i p i t a t i o n . The i n v e r s e r e l a t i o n o f s t o m a t a l d i s t a n c e w i t h t h e parenchymatous a r e a and d i r e c t r e l a t i o n w i t h the e n l a r g e d t r a c h e i d elements seem t o be c a s u a l because the changes are d i s - c o n t i n u o u s i n the i n t e r m e d i a t e samples a l t h o u g h t h e s t o m a t a l d i s t a n c e s i n c r e a s e g r a d u a l l y from t h e t o p t o the bottom o f the column i n T a b l e 4. - 37 - CONES : FIELD PROCEDURES, TECHNIQUE OF PREPARATION AND MEASURING OF CONE • PARTICLES. Cones for examination were collected from 40 stands. Most of the samples were collected from the same trees as the needles discussed above and attempts were made to obtain as uni- form a sample of cones as possible. When cones were not available from the same trees as the needles, other trees were used for c o l - lection of cones in the same stand, care being taken to obtain them from similar trees. Sometimes the cones were more than one year old remaining on the trees over the season and sometimes c o l - lection was made in the end of August when seeds were s t i l l im- mature. The distribution of samples and the measured data are given in Table 7. SEED. Only a small quantity of sound seeds was available for ex- amination of seed v i a b i l i t y and weight. The location of the stands and the result of the analysed data are presented i n Table 8. To separate viable from empty seeds ethyl ether was usedo The sound seeds sink while the empty seeds float.This was the method used to select viable seed samples for further test. Cleaned seeds were put into a desiccator for 48 hours to equilibrate the moisture content of the different seed stock. Weighings were made by micro-analytical balance i n grams per 1000 seeds, Table 8. • One portion of the seed specimens was used for germination test. For stratification, seeds were embedded in sand and kept in refrigerator under 37°F temperature for 60 days. Soundness and per cent of f i l l e d seeds were tested by germina- tion test and cutting test respectively. The largest seed was found in the cones from Mt. Idaho ( 24 ) , Figure 16. The sample standard deviation ( S ) is equal to 1.678 gram/1000 seeds. A" t-test shows that there are probably significant differences between the mean value of No. 24 and the individual means of samples,because the calculated t - value does not exceed the value given by the 1 fo probability level for 3 degrees of freedom in the Ficher's Table. The small- est seeds were collected around Aleza Lake,Figure 15. SEED WING. To test the var i a b i l i t y in form and size of the seed wings as many as available were prepared for investigation,also included were the wings of some empty seeds. In this way, eleven of 40 samples were obtainable. Thirty wings were taken out from the middle part of the cones of each sample. The wings were soaked in water for 24 hours then pressed on drawing paper to flatten them. The shape of wings was outlined in f u l l size on the drawing paper and the area of each wing was measured with a planimeter i n square centimeter, Table 8. The standard deviation of sample means is equal to 0.344 sq.centimeter which' indicates some significant d i f - ferences between the individual means. There are conspicuous ex- tremes such as 0.15 for No. 9 » 0.18 for No. 5 and 1.40 for No. 18 and lo05 f o r No. 38 . These d a t a were t e s t e d f o r d i f f e r e n c e s by the t - t e s t and h i g h l y s i g n i f i c a n t d i f f e r e n c e s were found between the means o f each i n d i v i d u a l and the sample m e a n . A l l the t v a l u e s exceed the v a l u e g i v e n by the 0.1 f> p r o b a b i l i t y l e v e l f o r 10 degrees o f freedom. Trees growing on d r y , r o c k y and even on c o o l s i t e s have the s m a l l e s t wings. For example,the sample t a k e n from Mt. R a i n i e r ( N o . 9) o c c u r r i n g at 7500-8000 f e e t has the s m a l l e s t wings, F i g u r e 5 • At t h a t l o c a l i t y , the t r e e s are s m a l l and s t u n t e d . I n c o n t r a s t t o them, t r e e s t h a t are g r owing on good p o d z o l i c s o i l , as i n W i l l s o n Creek ( N o . 38 ) or b e s i d e the A l a s k a Highway at 160 m i l e (No. 18) have the l a r g e s t seed w i n g s , F i g u r e 5. CONE AXIS. Unbroken a x e s , 40 o f each s t a n d , were s e l e c t e d , T a b l e 8. The axes were measured from the t i p t o the bottom a t the j o i n t o f t h e twig.The measurement was made i n m i l l i m e t e r . The s t a n d a r d d e v i a t i o n o f means i s e q u a l t o 15 m i l l i m e t e r s . T h e d i f f e r e n c e between the sample mean and i n d i v i d u a l means i s p r o b a b l y s i g n i f i c a n t f o r No. 12,26 and 35 samples. The v a l u e s f o r t h e s e do n o t exceed the v a l u e o f »t' g i v e n by t h e 1% p r o b a b i l i t y l e v e l f o r 6 degrees o f freedom. The «t » v a l u e f o r No. 38 f a l l s between 1 f> and 0.1 f> p r o b a b i l i t y . l e v e l i n the F i s h e r ' s T a b l e and so i n d i c a t e s s i g n i f i c a n t d i f f e r e n c e . The c o r r e l a t i o n between the l e n g t h o f the cone a x i s and the e n v i r o n m e n t a l f a c t o r s i s e v i d e n t . W ith b e t t e r h a b i t a t s t h e r e are l o n g e r cone axes and b i g g e r cones. There seems t o be - 40 - some increase i n the development•of the cone with higher precip- i t a t i o n . I t should be noted that the measured data of cones c o l - lected i n Willson Greek, i n the eastern part of the Interior of B r i t i s h Columbia No. 38 , d i f f e r from the data of other samples i n a l l the three c h a r a c t e r i s t i c s tested. However, the trees also d i f f e r i n crown form, color and growth rate from the other stands. The d i s t i n c t i o n of these c h a r a c t e r i s t i c s may allow one to suppose that the stand i n Willson Creek involves variations which can be either genotypic or ecotypic, Figures 9,13»16 . CONE SCALE AND BRACT. Twenty scales with bracts were picked ran- domly from f o r t y stand samples. Both the scale and the bract were measured separately. The r e s u l t of the measurements i s presented i n tenth of millimeter i n Table 7, under X+ and X5 r e s p e c t i v e l y . The difference of the lengths i s given under Y. Theoretically Y i s equal to ( X̂. - X5) . Actually, Y was found i n a di f f e r e n t man- ner as follows: Ten of the twenty scales with bracts were picked at random.These were measured and the'ten length differences were totaled to obtain Y i n hundredth of millimeter. Then, Y was i n t r o - duced into computer to have the analysis' of variance f o r the d i f - ferences of the ind i v i d u a l stand means and make regression analysis for the factors that may affect the development of the scale and bract. Tenth and hundredth of millimeter were used as a unit of measurement to avoid the decimal without loosing the accuracy of the data i n the tables and the computer records. Some selected bracts from the measured specimens - 41 - were photographed.The picture of these i s given i n Figure 14. Sometimes the scale and "bract were crooked and twisted. In th i s case they were soaked i n water f o r 24 hours, then pressed u n t i l they became dry and f l a t , thus malting for easier measuring. For h i s t o l o g i c a l examination, scale and bract were used i n immature stage a f t e r the tissues had been s l i g h t l y d i f - ferentiated. For this examination specimens were collected i n the Coast Region at Lake Garibaldi, and on Vancouver Island af t e r A p r i l , depending upon the elevation where the trees were located. At high elevation the cone tissue d i f f e r e n t i a t e s at the end of May or some days l a t e r , at low elevation i t usually happens early i n May. The same laboratory method was used f o r preparation, dehydration and staining of material of scale and bract as had been done to the needles mentioned above. EVALUATION AND ANALYSIS OF THE DIFFERENCE BETWEEN SCALE AND BRACT LENGTH.. ' The length of scale and bract can be measured e a s i l y giving adequate data for s t a t i s t i c a l analysis. Therefore, many taxonomists use i t to discriminate the natural hybrid of two clo s e l y related species or d i f f e r e n t variations within one species. Boivin ( 1959 ) suggested a new c l a s s i f i c a t i o n within the species of balsam f i r , on the basis of the r a t i o of cone scale - 42 - and bract. He places alpine f i r as a subspecies of balsam f i r . Myers and Bormann (1963) adopt Boivin's method and use i t to distinguish balsam f i r from the Fraser f i r in some southern regions of North America . The statements and methods of these two authors led the writer to examine the ratio of the cone scale/bract length in the alpine f i r variations. The examination has given a clear picture of the diversity of the scale/bract ratio and confirmed that a wide range of variations could exist within this species. A preliminary t r i a l was made on some reduced specimens, taken from the 40 stands to prove the confidence of the scale and bract length difference against the scale/bract ratio. Both the scale - bract length difference and the scale bract ratio gave the same result i n the discrimination of alpine f i r variations. Because the calculation of length differences (Y) i s quicker, i t was used to measure a l l the samples collected from the 40 stands. Hypothesis for analysis was the following: There are no significant differences between the value of ' Y' between the sampled stands. In connection with this hypothesis i t i s assumed that the scale and bract have constant proportion in size. Both the scale and the bract may change symmetrically i n size under various environmental effects, and during different growing pe- riods, but the ratio between their length remains relatively con- stant.If not, the difference between them should not be due to change alone. RESULT OF THE ANALYSIS OF VARIANCE. The range of Y , from 0.5 to 3.9 millimeter, shows high v a r i a b i l i t y . l t was supposed that there are significant differences between the stand samples and an analysis of variance was computed to prove i t . The model of analysis of variance i s the following: Source Df Sum SQ Mean SQ F Distance 39 3.97049B+04 1.01807E+03 131.57 Residual 40 3.09500E+02 7.73750E+00 Total 79 4.00144E+04 The F value exceeds the value for F indicated in the body of the table for the 0.1$ level with 40 and 39 degrees of freedom. The F test shows that the difference of the value of Y, i.e. the length distances between the scale and bract are highly significant between the different stands. As a conclusion of this analysis i t can be stated that there are actual variations in this cone collection. Some of the samples are closely related to each other,and others are less so. (< The relation between.the stand sample means can be determined through the Duncan's method by the test of least significant differences ( LSD ) . This method was described, above in connection with the analysis of stomatal distances of • needles (pp 21 - 23 ) . LSDc,oT) t,o,VsV f equal to t r o l x ^ 7 ^ 1 3 1 5 / 2 LSDo='tr_oi;.x 1.97 - 44 - The expression of analysis in diagram is given in Figure 7. This illustrates the associated samples as groups underlined by continuous horizontal lines. Since the method of reading of this diagram had been explained in pages 21 and 22 , i t i s only necessary to state the conclusions here. The sample means ( y ) of stands are highly variable and the arrangement of groups, underlined by the same horizontal lines, is not consistent concerning the location of stands i n the f i e l d . The group which has the smallest values of Y, No.l and 2, are located at Garibaldi Barriers,British Columbia and on the northern slopes of the mountains i n the Olypic National Park, Washington, respectively. Between the two stands and close to them, in the f i e l d , there are many other stands, Mt. Arrow- smith (21), Forbidden Plateau ( 32 ) , both i n Vancouver Island; Mt. Rainier, Washington, ( 8 and 9 ) , a l l located at about the same elevation apparently under the same environmental condition on intrusive rocks. But these samples are placed far from the low range limit i n the diagram. They are closer to those groups which were diagrammed in the opposite side and occur under better conditions on gray-brown podzolic soils i n the eastern regions of British Columbia and northern Washington. The highest group of (Y ) involves: Alder T r a i l , Montana, ( 40 ) , East Slope of the Glacier National Park, Montana, ( 39 ) , and Willson Creek, British Columbia, ( 38 ) . The stands, diagrammed in the middle portion of Figure 7 and underlined by the same continuous horizontal l i n e , are widely distributed i n the alpine f i r range and located on different habitats. In Table 7, the data are increasing more or less proportionally with the values of Y except those of the bract lengths which change irregularly in the series of obser- vations from No.l to 40. It i s obvious that the mean of stand samples ( X6) and the length.differences ( Y)exhibit parallel i n - crease because both were calculated from the same measurements. Nevertheless, the proportional increase of the scale lengths (X,,) and length differences (Y) and the irregular change of the bract length ( X5) need further explanation and analysis to re- veal the associations between the variables mentioned above. In the regression analysis, some ecological factors are introduced as independent variables. Precipitation (X,) , latitude ( X2) and altitude (X5) are the only factors avail able for regression analysis. The results are given in Tables 2 9 , 10 and 11 • The sample regression coefficient, R»0.993, reflects close association between Y and the variables mainly because the values of Y involve the individual means of samples (X6) • However, omitting the corresponding variables one after the other i t i s possible to exhibit the relations between them. Herbert and Raymond Tables ( 1950 p. 140 ) of Ifo points for R should be used to evaluate the associations of variables. The computed value exceeds the value in Table for 34 degrees of freedom and 6 variables. It i s assumed that significant associations exist between Y and the variables. Table 10 presents these associations and allows one to conclude the following: Precipitation correlate with latitude,altitude , and bract length Scale length correlate with individual mean of samples and Y Bract length correlate with precipitation and scale length Precipitation has an effect on bract length but does not affect the scale length. The bract may need a longer period for growing than the scale.Higher precipitation indicates a longer growing period for the bract and lower precipitation does the opposite. There i s correlation between scale length and Y but no correlation between bract length and Y. It should be necessary to analyse the relation of Y to the scale and bract length, separately. Two linear equations of X* and X5 on Y are made up to find correlation between X+ , X5 and Y using a graph to show the two regression lines of these variables. Equation of scale length: Y= 42.213 + 0.573 X* . Regression coefficient equals to .573 and an estimated value of Y-50, the value of X,-148.25. Equation of the bract, length: Y-55.919+0.335X5. Regression coefficient: .335, estimated value of Y=50, then X5= 105.60. The divergency of the two lines indicate a poor correlation between the l e n g t h of scale and bract. On the other hand the value of Y depends rather on the scale length than the bract length as the steeper slope of scale line exhibits. The graph of the lines X4 and X5 should he the following: 0 ' : ( H 8 , 4 3 ) P, : (198,77) I r— • • X 100 . 150 200 The explanation of the relation of X4 and X 5 to the sum of the difference between the length of scale and bract can be purely numerical i.e. summarizing a large and a small figure the larger w i l l increase the sum proportionally higher than the smaller figure. On the other hand, a biological expla- nation, i f i t i s exists, would be more interesting, but i t i s beyond the scope of this study. HISTOLOGICAL CHARACTERISTICS OP SCALE AND BRACT. The origin of the scale and bract as organs of the plant has been debated for a long time; to try to reconcile the various concept i s more d i f - f i c u l t than important. Coulter and Chamberline (L910) present a brief account of the history of the concept of the ovuliferous scale:. 'The ovuliferous scale of Pinaceae has been regarded successively an open carpel, a placenta, a flattened axillary shoot, the f i r s t - 4 8 r two leaves of an axillary shoot, the f i r s t and only leaf of an axillary shoot, a ligule, fused outer integuments and a chalazal outgrowth'(pp. 244-248 ). Schoute (1913) gives some points about the development of cone scale including bract and assumes that at the growing point, bud forming materials may be present which have a restricting influence on leaf-formation. Every part of the vegetative cone in which the influence of the bud forming material ceases, can form a leaf p r i m o r d i u m insofar as other i n - fluences do not prevent i t . The leaf primordium i s not visible through naked eye. Jeffery (1922 )calls the cone scale a repro- ductive f o l i a r organ and points out that the view that the ovuliferous scales in the Abietineae consist of a fused pair of f o l i a r structures has l i t t l e evidence to support i t . It i s as clearly a single leaf as i s the microsporopnyll* Busgen ( 1929) considers the cone scale as a leaf which later in the developmental phase becomes leathery or woody. Poster and Grifford (1959 ) emphasize that doubts have been repeatedly expressed by morphologists regarding the f o l i a r nature of the ovule-bearing structure in gymnospermous plants. The evidence from ontogeny and vascular anatomy i n d i - - cates that this ovuliferous structure is a highly condensed short shoot and not a simple megasporophyll. The following observations confirm Poster and -. 49 - Gifford's statement and show that the arrangement of tissue elements iii the cone-scale are more similar to the stem than to the needle. The author observed that the cross-section of the cone bract has a needle-like structure in inverse arrange- ment, related to the position of bundle xylem and phloem. The different structure in the cone scale and bract may allow one to assume a different derivation of these organs with a subsequent adherence during the development of the strobilus. However, f i n a l conclusions have to await further investigations. SCALE. The hairy and resinous scale consists of epidermis in the cross-section which lacks stomata, but has mesophyll or cortex, numerous strands of vascular bundle, and various forms and numbers of resin canals, Figure 22. The epidermis i s multiple in the structure having 1 to 3 layers of cells and lacks chlorophyll. The cells of outer layer are thick-walled without cuticle on i t s surface. The scale i s covered with resin when the cone matures and dries out. The ; epidermis i s composed of regular epidermal cells and trichomes. Trichomes are built up of one to three c e l l s . They are actively secretory having dense protoplast elaborating resin. Trichomes cover the surfaces of scale on both sides except in those regions where the seed wings and the bract attach to the surface. The author had the opportunity to make some observations on different species of Abies and found that the length of trichomes differs in various species: red f i r has longer,silver f i r has shorter - 50 - trichomes on the scale surface than the alpine fir.Usually the mature scales have longer trichomes in all.cases. The mesophyll does not differentiate into palisade and spongy parenchyma. The tissue complex i s relatively homo- geneous in i t s features, being similar to the cortex i n having hypodermal cells in the outer portion. Intercellular spaces, like subepidermal chambers in the needle mesophyll have been found under the epidermis at the adaxial side. The contents of cortex cells are starch and resin. Leucoplasie were found in the medial cells of cortex. The young parenchyma cells are l o - cated in the horizontal middle sheath rather than i n the marginal region in the cross-section of the scale. In a longitudinal section, elongated parenchyma cells are found immediately under the epidermal layers and have simple pits in the wall of the cells located in the outer layer of the parenchyma tissue. The vascular strands are arranged horizontally in transection. Eighteen to twenty-two strands are present, distributed regularly in one plane parallel with the main axis of the scale. A diagram of scale venation pattern i s given i n Figure 6. The longitudinal strands converge at the base and d i - verge at the apex of the scale. These' terminate in the cortex and are enclosed in transverse parenchyma cel l s . The parenchyma cells have simple pits on their side walls. The vascular bundles are not separated, so form one strand. The xylem element of this united bundle i s located on the adaxial side of the scale and the phloem tissue on the abaxial side. The xylem consists of tracheids - 51 - i with, annular and helical thickenings and parenchyma cells with simple pit fields on the side walls. The sieve cells alternate with parenchyma cells and abundant albuminous cells appear to be located at each side of the phloem. / The transfusion tissue consists of thick walled parenchyma cells and some tracheids. The latter are closer to the bundle than the former. r An ill-defined endodermis sheath lines the vascular and transfusion tissues. The sheath i s interrupted by the extension of parenchyma cells which interconnect the strands laterally throughout the scale. " The resin canals are arranged mostly parallel with the vascular strands. It is possible to distinguish both large canals and some narrow canals. The larger ones are located i n the adaxial half of the scale, close to the strands and more or less of the same number as the strands. The resin canals have an inner sheath that lines the cavity of the canal. This consists of thick- walled and large secretory epithelial cells f i l l e d with resinous material. The thin outer sheath is made up of small parenchyma cells which have a large nucleus. These cells appear to be inter- mediate between the secretory and cortex c e l l s . The smaller canals are distributed irregularly in the region of the vascular strands and are usually between two neighbouring strands. Their structure is much like the large canals but the cavity diameter is less and looks like a gap or an inter- cellular space in the center of a group of epithelial c e l l s . - 52 - BRACT. The bract consists of epidermis with stomata, hypodermis, mesophyll, vascular bundles and resin canals, Figure 20. The epidermal layer is not covered by cuticle. On the adaxial surface stomatal openings interrupt i t s conti- nuity. The wall thickenings of the epidermal cells are uneven and sometimes only the outer walls are thickened. The bract does not show xeromorphic characteristics as does the needle with their sunken stomata and cutinized epidermis. Probably the scale protects the bract ,from drying up. There are no chloro- plasts in the c e l l of the epidermis, not even in the young cones of alpine f i r . Stomata are arranged in 8-10 parallel rows along' the f u l l length of the midrib in one narrow band. The stomatal cells i.e. guard cells and subsidiary cells are located lower than the epidermal cells around the opening but not sunken as in the needle. The occurrence of stomata on the bract surface has not been mentioned in the literature used for references., Their presence here may be evidence, that the bract and scale have the different origins. Several specimens of Abies, Picea.Pseudotsuga and Pinus have been examined for stomata on the adaxial surface of their cone-bracts. It should be noted that the pines have several too many spirally arranged bracts each subtending an ovuliferous scale. With the exception of Pseudotsuga, a l l the genera mentioned have stomata on their bract surface, and each has the same arrange- ment of rows as alpine f i r . - 53 - Hypodermal cells are located under the epidermal layer of the adaxial surface in a discontinuous row. The fiberlike lamellar hypodermal cells are uneven in size and a l l of them have strongly thickened-walls. Sometimes they form two rows toward the edges of the two wings of the tract. There are conspic- uously large hypodermal cells in the second row. Above the midrib the hypodermis occurs in one row, consisting of small cells with well-thickened walls.. Under the epidermal layer of the abaxial surface, no hypodermal layer i s formed. This side of the bract is soft and smooth and adheres to the seeds thus protecting them in the developmental phase. The mesophyll is not well-differentiated in the specimens collected at the period of examination. The c e l l walls seem to be thickened and f u l l y perforated with primary pit f i e l d s . There are shizogenous intercellular spaces between the c e l l s . The cells cling closely together forming outer ridges from their walls which press the wall of the neighbouring cells into the c e l l lumina. The content of the cells i s homogeneous and no plastids appear to be inside the c e l l s . The vascular bundles are separately lined by an i l l - defined endodermis sheath. There is no difference between the cells of the mesophyll and endodermis in the structure of walls and the shape, but only in size. The endodermis cells are smaller than the needle endodermis and turn their tangential walls towards the vascular bundles, apparently having no Casparian strips in their side walls. Some of the walls are pitted. Inside the endodermis, the transfusion tissue con- sists of l i v i n g parenchyma cells and tracheids as does the transfusion tissue complex of the needle. Both the tracheids and the parenchyma form continuous system and the two system interpenetrate each other. Rarely, some sclerenchyma cells appear among them. In the neighborhood of the vascular bundle, like a large many-layered sheath, albuminous cells are located at each horizontal side of the bundle with dense cytoplasm and prominent nuclei. The separated vascular cells are oriented as in the needle, but the xylem complex points towards the adaxial side of the bract. The total number of radial rows are 8 to 10 to- gether in the two separated parts of the bundle. Both the xylem and the phloem have thickened and l i g n i f i e d walls. Annular and helical thickenings are present in the xylem. The parenchyma cell s , located between the two parts of the bundle and around the bundle, have primary pit. fields and simple pits in their secondary walls. Resin canals are two to three in number, embedded in the mesophyll i n medial location as i n the needle cross-section. The resin canals are lined with thinr-walled secretory epithelial c e l l s . Outside these c e l l s , a sheath of parenchyma cells make contact with the mesophyll tissue. - 55 - The author observed that the bract develops f i r s t during the growing season and reaches a slightly differen- tiated form and size earlier than the scale. Therefore, the bract protrudes beyond the scale in the early developmental phase of the strobilus. Some cones have been examined i n early stages of differentiation of strob'ili and have been found to have the bract separated from the s c a l e . This may be further evidence that the origin of scale and bract is not common. - 5 6 - DISCUSSION AND CONCLUSION. Ten of twenty traits were tested by 't' test and an additional two by regression analysis* Significant d i f - ferences of several traits were revealed among stand samples located in various ranges of the alpine f i r on the Vest Coast. But the traits tested do not give sufficient evidence to prove the existence of regional populations perhaps because an equal number of specimens was not available for a l l t r a i t s , e.g. cones were collected only from two-thirds of the stands sampled for stomatal distances, seeds were available only from four stands. Thus, the question of whether alpine f i r i s to be con- sidered one general population or composed of several populations isolated during the geological eras is one that this study i s unable to answer. The scope of this project has been too limited by time and opportunity to observe continuity and adaptability of quantitative or qualitative characters on a large scale which may indicate the existence of genotypically different or regional populations.The fact that there are significant differences be- tween some specimens concerning the stomatal distance or scale and bract length does not indicate regional differentiation within the population of the alpine f i r . A population can include groups of individuals of almost any phenotypes. Stebbins (1957) defines the population as a group of individuals among which a , larger or smaller amount of interbreeding and gene exchange can occuro Studies in natural stands, i.e. the taxonomic approach, - 5 7 - have the major disadvantage that the method i s purely descrip- tive and no precise distinction can be made of the environmental and genetic components of variations. And so, i t i s not possible to present evidence here for any variations between the stands sampled which seem to be solely under genetic control. The author i s convinced by numerous observations that there are natural hybrids between alpine and balsam f i r i n the overlapping areas (Roller 1966b)••• Several traits such as resin canal position and hypodermal tissue i n the needle cross-section, thickness of the needle, distribution of stomata on the upper surface of the needle, size of the parenchyma ex- tension in the bundle, length and number of the lenticels on the middle-age bark and finely the percentage of the /3 - pinene and & - phellandrene in the resin, show intermediate charac- te r i s t i c s between alpine and balsam f i r in the samples collected from northern Alberta and British Columbia. However, the most d i f f i c u l t i s to identify any hybrid by purely morphological characteristics. In the regions, where these putative hybrids occur, the different species and hybrids exhibit characters which increase or diminish in a given direction under the same environmental conditions and therefore the individuals of these different species show quite similar phenotypic characteristics. Some characteristics may vary on different parts and at various levels of the same tree. Some- times, certain characteristics vary on the same part of a tree during the long period of tree development. Nevertheless, the length of period depends on the aspect, habitat,association etc. This consideration does not support the c l a r i - fication of the hybrids or the ecotypes which may exist in the different regions where habitat differences are found. Certainly, the geographical distribution of alpine and balsam f i r is very wide and trees growing in various regions exhibit high variety i n morphological characteristics which sometimes must be due either to natural selection or other mechanisms such as mutations introgression or genetic. In most cases, i t i s almost impossible to define that the characteristics, selected for testing, are variations or the result of a higher tissue differentiation i n the aging of the plant. Most parts of the inner tissue in the needle /- / • / follow this pattern, particularly those which take part inten- sively in the building up of the tree. For instance, the stele of the needle has a very high variab i l i t y in the arrangement of conducting elements, Figures 23-30 while the mesophyll cells do not vary significantly during the development of the needle either within or between the samples collected, except the chlorophyll content in the palisade c e l l s . The needles of under- storey trees and seedlings have conspicuously higher chlorophyll content in the palisade cells than the needles sampled from trees in open grown stands, or needles from cone-bearing branches which are in a more developed stage. On the basis of observations and measurements, and after considering the high v a r i a b i l i t y of characters within stand - 59 - and between stands, two variation patterns have been revealed. Those stands which include variations such as the stomatal distances, scale and bract length differences,and different size of seed wings are distributed horizontally in different regions of the natural range of alpine f i r and sepa- rated by geographical barriers. The other groups of the variation patterns i n - volves those stands which are distributed vertically i n the same region, being located at different altitudes and related to ecological gradients within a restricted area. They occur i n characteristics such as height, crown and bark color, needle . form, cone size and color, stomatal frequency on the adaxial surface of needle,etc.. ! In the f i r s t case, regression analysis confirms that the va r i a b i l i t y of stomatal indices and scale-bract length differences are under environmental control. Although there i s no evidence against the hypothesis that the stomatal frequency may be adaptive, the analysis has not shown any genetic control of the formation of the stomatal distribution. Obviously, that distinction between environmental-induced modification and genetic variation could not be made. Stomatal distance in a row line was inversely correlated to the precipitation, elevation and latitude. This means that the environment is largely resposible for the stomatal distance and that changes in the environment affect changes in t r a i t values. A l l the traits concerned are discontinuous and divergent in the series of sampled stands. Thus, i t is not possible to establish a systematic classification, at least according to two characteristics like stomatal distance and difference between scale and bract lengths. The horizontally distributed variation pattern should be illustrated by some examples. The extreme stands, which are the ends of this variation pattern, constitute samples at Garibaldi lake (No. 1 ) and on Wallowa Mts. (No. 65) • The data of these two extremes are presented in Figure 5» Signi- ficant difference between the sample means i s proved by analysis of variance, (p. 2 0 ) , but the trees in both stands show simi- l a r i t i e s i n their character combinations. Habitat i s different i n both regions. Garibaldi Lake i s located in the coastal forest region occupying a zone between Douglas f i r , hemlock-cedar for- ests, and the alpine tundra and snow fi e l d s . This forest type i s characteristic of both the Subalpine and the Coast Regions. High precipitation and true podzol predominate. Wallowa Moun- tains are located in the Mountain Forest region of the United States characterized by lower precipitation and f a i r l y deep and loose podzolic s o i l . The two regions are far enough from each other to be quite separate. It is concluded that the difference be- tween needle characteristics from these areas arose after their regional isolation and might be due to adaptation during the past geological eras. Between the two regions, which are almost - 61 - the western and eastern range; limi t of alpine f i r occurrence, there are several intermediate types of alpine f i r and the value of stomatal distance shows some decreasing tendency from east to west. But the decrease of stomatal distance i s irregular and discontinuous, not constant, therefore, i t would not be reason- able to draw conclusions for introgressive populations, / However, in many instances there are gradual changes i n the stomatal distances from one stand to a distant one, A • gradation of character occurs, called cline by Huxley (1933) who l i s t s many of them. This cline may be classified as intra group cline because the mean values of character change gradually through the continuous population* The author does not consider the continuous variations of one tr a i t to be sufficient to estab- l i s h a cline even i f they have been followed i n the series of stands. The author considers that the second variation pattern, i.e. the vertical segregation of different stands, i s a cline and demonstrates i t by an example. In the Glacier National Park, Montana altitude ranges from 3500 to.7000 feet on high mountain slopes. Trees located at the higher altitude around Logan Pass and at lower altitude along Alder T r a i l possess characteristics that change continuously from the high to the low elevation. These may be parallel the example of Clausen, Keck and Hiesey (1940 ) adopted from Stebbins ( 1957) i n connection with two ecotypes of Potentilla glandulosa subs. nevadensis. The quotation i s the following: * It consists of two ecotypes, one i s a dwarf, early-flowering alpine that occurs - 62 - above 2600 m., while the other is subalpine and may be d i s t i n - guished i n garden cultures by i t s t a l l e r stature and later flowering.' The occurrence of the alpine f i r from Alder T r a i l up to Logan Pass i s analogous the existence of two Potentilla ecotypes at low and high elevations. Baur ( 1932 ) gives another similar example of Antirrhinum glutinosum which was found i n the Sierra Nevada of southern Spain at altitudes ranging from 800 m. to 2800 m. on high mountain slopes. Plants taken from higher altitudes show different characters i n frost resistance and habit from those grown at lower altitude. ; No doubt, two types of alpine f i r occur in the Glacier National Park from Alder T r a i l to Logan Pass and their separation i s due to the response of the selective effect of ecological and climatic differentiation. This means differences i n the upper horizon of soi l s , growing season, radiation,annual average of temperature, -humidity and number of frosty days* The annual average of precipitation does not differ significantly but the distribution of snow and rain f a l l do so. At high ele- vation, snow covers the slopes u n t i l June while along Alder T r a i l snow i s usually melted in late A p r i l . The question may be justly asked i f these two types of alpine f i r could be classified as cline. If Huxley's definition of the cline i s taken into consideration i.e. 'internal intra group clines occur when the mean value of the characters changes gradually through a continuous population* the two types of alpine f i r would be internal clines because the means of the tested t r a i t s , such as stomatal distance, scale/bract length difference, needle length and width, bark formation, crown - to - color, height of tree, branch formation, etc. vary gradually from Logan Pass to Alder T r a i l . These two extreme types of Abies lasiocarpa might be called as 'high-alpine type' at high elevation and 'subalpine type' at lower elevation. The taxonomical description of these two types i s the following: Mature bark: High-alpine type: ashy grey or whitish, slightly fissured on the lower and smooth on the higher part of the trunk. Sub-alpine type: dark, grey, furrowed and superficially scaly. Bark on the upper stem usually exhibit well-developed horizontal lenticels, Pigure 33/ which do not appear so de f i - nitely on the bark of high-alpine type trees. Mature needle: High-alpine type: cross-section slightly rounded, less than 2.5 cm. long and about 1.60 mm. wide, yellowish or light yellow green. Stomata on the adaxial side appear as a narrow band along the midrib but abundant at the apex, f u l l y covered on the needles of top branches; on the abaxial side the mean of stomatal frequency i n square millimeter i s 103* Apex i s acute and pointed, the latter occurs particularly on the leading shoot; slightly rounded i n the lower part of the crown. The crown con- sists of ill-developed, short branches in i t s upper part. Sub-alpine type: cross-section flattened, longer than 2.5 cm., and about 1.80 millimeter wide, long, dark blue and blue green to silvery green. Adaxial surface i s covered by i r - regularly distributed stomata. On the abaxial surface,.the mean - 04 of stomatal frequency in a square millimeter i s 70. Apex i s notched and marginate. The crown i s dense and consists o f well-developed, proportionally distributed branches•i Cone: High-alpine type: dark, slightly black with resin flows. Axis i s 4 cm. long. The tip of bract short with unformed/ shoulder; seed wing shiny and purplish. Sub-alpine type: dark purple, length of axis i s 6 cm., the t i p of bract i s long,slender,pointed, seed wing yellowish brown. The high-alpine type variation grows five to six meters in height with slender form and spire-like crown, but scarcely distributed branches, Figure 10. They occur i n small pure groups dispersed along the timber l i n e . The sub-alpine type variation develops normally, reaching 25-30 meters in height and mingle with other forest trees forming continuous stands on well-drained mountain-sides, Figure 11. Trees were found with similar habit to the high- alpine type variation on dry sterile sites on the lava rock o f Belknape Crater (61) and McKenzie Pass \( 30) Oregon and on Hurrican H i l l , Washington (2 ) » The trees have similar forms, colors and habits to the high-alpine type trees, but they are higher in growth, and their stomatal indices ( Y ) on the lower surface of the needle are 607, 504 and 457 respectively. The needle collected at Logan Pass, Montana ( 3) show an index of 461* The habitat at Belknape Crater is very dry and nothing else, but lava outcrop covers the ground without any plant association. The alpine f i r i s pioneer here, where wind blown material has / begun to accumulate and may exhibit an ecotypic variation different from the others. The high stomatal index suggests a dry ecological, condition and presents an essential distinction between the two variations located along Belknape Crater and at Logan Pass. The observations on the sampled trees and the analysis of the measured data give a picture of the diversity of quality differences in the stands scattered on the West Coast. It i s obvious, that the greatest diversity i s expected between the extremes located in northern and southern limits of the natural range of alpine f i r . If the traits in these ex- tremes inherited and became aquired the finding of some geno- typic variations in i t s natural range can be expected. 66 - • SUMMARY. This is a morphological-anatomical study of natural stands of Alpine f i r . Its objectives were ( a ) to determine the va r i a b i l i t y of needle and cone characteristics for two morphological t r a i t s : distance between stomata in a row line on the abaxial surface of the needle and difference between cone scale and bract length; (b ) to become familiar with other t r a i t s ; and i f possible ( c) to ascertain whether alpine f i r i s essentially one population and i t s phenotypic variations follow certain environmental gradients. ' Sixty five stands, located in the Yukon Territory, Bri t i s h Columbia, Washington, Montana and Oregon,were sampled. laboratory procedures involved: preservation of needles and young cones in PAA, needle and cone cross-sections by microtome, several sets of observations and measurements on needles,seed wings, cone axes, cone scales and bracts. The following tree mean data were used as the basic units for the study of variation: needle length, width'and apex form; presence of groove and ridge; color of foliage; stomatal distance on the abaxial needle surface; stomatal distribution on the adaxial needle surface; stomatal structure; seed wing area; seed weight; length of scale; length of bract; difference between scale and bract lengths; length of cone axis; histology of needle, scale and bract; and resin canal position. Mean, - 67 - standard deviation and range, were calculated from tree means, and stand means were computed. Correlation analysis and 'V test were used to show the association between several traits and environmental factors .and prove significant differences between the stand means. The correlation was the highest between stomatal distances and precipitation. The stomatal distance in a row line i s inversely related to the increase of precipitation,and so stomatal frequency increases with higher precipitation. A l l traits show higher variab i l i t y between stands than between trees sampled. The v a r i a b i l i t y of traits i s appar- ently largely due to the environmental factors. Nevertheless they sometimes suggest casual associations. Eastern and western varieties, and high-alpine and sub-alpine f i r have been described. The major conclusion i s that care must be exercised i n describing t r a i t v a r i a b i l i t y . The particular environmental gradient must be specified because the same t r a i t can show c l i n a l , ecotypic and random relationship patterns, depending on the component with which the t r a i t i s compared. -. 68 - LITERATURE' CITED. 1. Anonymous. 1961. Native trees of Canada. Dept. of Northern Affairs and National Resources.Forestry Branch, 6th Edition, Bull. 61. 2. Boivin, Bernard. 1959. Abies balsamea fLinne) Miller et ses variations. Le Naturalists Canadien 86(10):219-223. 3. Busgen,M. 1929. The structure and l i f e of forest trees. 3rd edition. John Wiley and Sons, Inc., New York. 4. Chiasson, L.P. I960. Report on tree breeding research. Proc. 7th Meeting Comm. For. Tree Breed. Canada.Part II. pp. " C 1-4. 5. Chiasson, L.P. 1962. Report on tree breeding. Proc. 8th Meeting Comm. For.Tree Breed.Canada. Part II.pp.C 1-2. 6. : Collingwood, G.H. and Warren D. Brush. 1955. Knowing your trees. 15th Edition.Pub: The American Forestry Association,Washington. 7. Coulter, John M. and Charles J. Chamberlain. 1910.Morphology of Gymnosperms. 458 p. The University o f Chicago Press, Chicago, I l l i n o i s . 8. Dallimore, W., and A.B. Jackson. 1961. A handbook o f Coniferae including Ginkgoaceae. Reprint.682 pp. Edward Arnold Ltd. London. 9. Duffield, John V/. 1962. Personal communication. 10. Duncan, D.B., 1955. Multiple Range and Multiple F Test. Biometrics 11:1 - 42. - 69 - 11. Esau, Eatherine. I960. Anatomy of seed plants. Ed. John Wiley and Sons,Inc. 12. Forest Genetics Research and Related Projects in the west- ern United States and British Columbia. 1958. Report of a Survey by the For.Gen. Research Foundation, Berkeley, California. 13• Foster, A.S. and E.M. Gifford. 1959. Comparative morphology of vascular plants. Ed: W.H. Freeman and Co. San Francisco, California. 14. Fulling,E.H. 1934. Identification, by leaf structure, of the species of Abies cultivated in the United States. Bull.. Torrey Bot.Club 61:497-522. 15. Gardner, E.J. I960. Principles of Genetics. John Wiley & Sons, Inc. N.Y. pp. 366. 16. Harlow, W.M. and E.S. Harrar. 1950. Textbook of Dendrology. 3rd edition. McGraw-Hill Book Co.Inc.,Toronto,pp.555. 17. Hutchinson, A.H. 1924. Embryogeny of Abies. The Botanical Gazette. 77(3):280-289. 18. Jeffrey, E.C. 1922. The Anatomy of Woody Plants. Ed.' The University of Chicago Press. Chicago,111., pp.460. 19• Johansen, D.A. 1949. Plant Microtechnique.Ed: McGraw-Hill Book Co. Inc.,N.Y.,pp.523o 20. ELaehn, F.U. and J.A. Winieski. 1962. Interspecific hybridiz- ation in the genus Abies. Silvae Genetica 11:130-142. 21. Ehuchel, H. 1954. Das Holz. Verlag H.R. Sauerlander and Co., Haran and Frankfurt am Main, pp.472.. - 70 - 22. Laing, E.V. 1956. The genus Abies and recognition of species. Scottish Forestry. 10 (1) :20-25. 23. lanner, M. Ronald and Stanely Li Krusman. 1963. Abies - A Bibliography of Literature for tree improvement workers. U.S. Forest Service, Res.Paper PSW-10. 24. L i , Jerome C.R. 1959. Introduction to s t a t i s t i c a l inference. Edwards Brothers, Inc. Ann Arbor,Michigan,pp.553. 25. Masters, Maxwell T. 1891. Comparative morphology, anatomy and l i f e history of the Coniferae. Jour.Linn. Soc. London Bot. 27:226-332. 26. Mergen F., Jeffery,B. and B.A. Simpson. 1964. A r t i f i c i a l hybridization in Abies. Der Zuchter 34 (6/7) :242-251. 27. Myers, Oval, Jr. and F.RV Bormann. 1963. Phenotypic variation in Abies balsamea i n response to altitudinal and geo- graphical gradients. Ecology 4 4 ( 3 ) : 428-435• 28. Roller, K.J. 1966a. Resin canal position in the needles of balsam, alpine and Fraser f i r . For.Sci. 12:2. 29. 1 1966b. Report on a putative natural hybrid between alpine f i r and balsam fir.Dept.Publication.Mimeo. 30. Schmidt, R.L. 1957. The s i l v i c s and plant geography of the genus Abies in the coastal forest of British Columbia. Dept.Lands and Forests, British Columbia For. Serv. Tech. Pub. T-46,pp.31. 31. Schoute, J.C. 1913. Beitrage zur Blattsteuungslehre. Extrait du Rec. d. travaux bot. Neerlandair.. 10 (3/4) • 32. Snedecor, G.W. 1956. Statistical Methods. 5th Ed. Iowa State College Press,Ames,Iowa, pp.534. 33. Stebbins, Jr., G. Ledyard. .1957. Variation and Evolution in plants. 3rd Ed. Columbia University Press N.Y., pp.643. . ' 34. Stover, Ernest L. 1944. Varying structure of conifer leaves in different habitats. Bot. Gaz. 106 (1) : 12-25. 35. Sudworth, G.B. 1908. Forest trees of the Pacific slope. Forest Service,U.S.Dept. of Agriculture.Government Printing Office,Washington, pp.441. 36. Sweet, H. I960. Notes, preparation of conifer needles for good sections. Unpublished. 37. Takeda, H. 1913. A theory of 'transfusion-tissue'. Ann. Bot. 27: 359-363. 38. Zucker, Milton. 1963. Experimental morphology of stomata. Papers and discussions on the Physiology and Bio- chemistry of leaf stomata. Ed: The Connecticut Agricultural Experiment Station, New Haven. Bulletin 664, pp. 80. Table: 1. DISTRIBUTION OP DATA OP ABIES LASIOCARPA ON THE WEST. COAST. X,:Stomatal frequency/ Xj. :Precipitation,inch X3:Needle width,millimeter X„ :Mean,distance in units between two stomata in one row line, 1 unit equal to 0.02 millimeter X s:Elevation,feet Xg :Latitude,degrees X7:Longitude,degrees Y :Combined data of stomatal • distances on twenty ran- domly selected needles. Explanation on page 19.. No Location X, x 3 X« xb- X, Y 1 Garibaldi Lake Trail,BC. 95 120. 1.74 4 4800 50.0 123.3 451 2 Hurricane Hill,Olympic N.P.,Wash. Logan Pass,Glacier N.P.,, 148 80 1.96 4 5464 47.5 123.8 457 3 Mont. 103 40 1.65 4 7000 48.5 113.8 461 4 Black Tusk,Garibaldi Prov.Park,BC. 118 140 1.69 5 5600 50.0 123.3 466 5 Arrowsmith Mt•,Peak,V.I. 127 130 1.87 5 5300 49.2 124.5 466 6 Scott Lake,Ore. 95 70 1.75 5 5000 44,.4 121.9 468 7 Arrowsmith Mt.,V.I. 127 100 1.88 4 2500' 49.2 124.5 468 8 Manning Park,BC. ' V 104 70 1.84 5 5200 49.1 120.5 475 9 Wood River,BC. 70 35 1.41 5 3000 52.3 118.5 480 10 Siskiyou -Mt s., Ore. 125 43 1.87 5 7433 42.0 123.6 481 11 DowniBCr. ,BC. 86 35 1.61 5 2500 51.2 118.3 481 12 Azouzetta Lake,BC. 78 35 1.96 5 4000 55.4 122.7 482 13 Ottauwau River,Alta. 88 15 1.52 5 2500 55.0 116.6 483 14 Boulder L.,Olympic N.P., Wash. 107 100 2.20 5 4500 47.5 123.9 485 15 Giscome Rd.,Willow R.,BC« , 67 30 1.40 5 2300 54.1 122.5 485 16 Naver Cr.,BC. 75 30 1.57 5 3000 53.2 122.2 486 17 Arrowsmith Mt., H i l l , V, .1. 96 110 1.87 5 4500 49.2 124.5 487 18 Parsnip River,BC. 71 25 1.37 5 3500 55.0 123.0 487 19 Mt.Rainier,Timberline,Wash91 120 1.62 5 8000 46.3 121.8 488 20 Porbidden Plat. , V. I. Pine River Valley,Alta. 92 140 1.64 5 5100 49.4 126.0 488 21 87 20 1.67 5 300a 55.5 122.5 490 22 Alaska Hwy.,Mi.500,BC. 143 17 1.66 5 2300 59.2 126.0 491 23 Bear Lake,BC. 88 25 1.58 5 2500 54.5 122.0 492 24 Hurricane Ridge,Wash. 105 80 1.71 5 5000 47.5 123.5 495 25 Mt.Rainier,Wash. 100 80 1.75 5 6500 46.3 121.9 495 26 Bear Lake,BC. 88 25 1.58 5 2000 54.5 122.0 495 27 Alaska Hwy.,Mi.356,BC. 74 20 1.60 5 3500 58.5 122.3 498 28 Aleza Lake,BC. 82 40 1.95 5 2500 53.5 121.7 499 29 Mt. Hood, Ore. 81 73 1.84 5 5200 45.3 121.4 501 30 McZenzie Pass,Ore. 95 60 2.00 5 5320 44.3 121.8 504 Table 1. cont. - 73 - No Location X, X, X, X, X, x r Y 31 V/illson Cr.,New Denver,BC. 47 40 1.87 5 3000 50.6 •118.5 505 32 Champion Lakes, BC. 99 35 1.65 5 3600 49.2 118.5 507 33 Andimaul Nash,BC. 34 Eagle Cap,Slope, 91 40 1.60 5 1600 55.3 127.5 510 Wallowa Mts.,Ore. 90 30 1.66 5 5500 45.3 116.9 514 35 New Haz©lton,BC. 36 Pendleton Bay,BC. 81 40 1.84 5 3600 55*0 127.5 514 62 20 1.90 5 2400 54:?.5 125.7 514 37 Kicking Horse Pass, Yoho N.P..BC. 73 30 1.79 5 4300 51.4 116.2 515 38 Alaska Hwy.,Mi.305,BC. 93 17 1.66 5 1100 59.0 123.5 523 39 Alaska Hwy.,Mi.l60,BC. 103 17 1.97 5 3500 58.0 122.5 524 40 Pine Lake,Yukon 110 8 2.15 5 2700 60.4 133.8 525 41 Alaska Hwy.,Mi.480,BC. 88 25 1.88 5 2800 59.0 126.8 527 42 Manning Park,BC. 77 50 1.60 5 4000 49.1 121.0 529 43 Wallowa Lake,Ore. 90 20 1.81 5 3000 45.3 116.9 529 44 Alaska Hwy.,Mi.852,Yukon 99 12 1.67 5 2500 60 o 2 133.0 530 45 Mt.Rainier,Mash. 106 79 1.62 5 2800 46.3 121.8 532 46 Crater Lake,Ore. 67 69 1.86 5 7500 43.0 122.3 539 47 Slave Lake,Alta. 99 15 1.73 5 2000 55;3 115 oO 539 48 Cirrus Mt.,Banff N.P.,Alta.65 35 1.71 5 6200 51*5 115.3 539 49 Mt.Bonaparte,Wash. 74 20 1.62 5 4500 48.7 119.2 540 50 Divide Cr.,Glacier N.P., Mont. 68 17 1.81 5 4500 48^5 113.5 543 51 Dixie Pass,Ore. 78 30 1.61 6 5000 44.3 117.8 544 52 Lava Camp Lake,Ore. 87 40 1.75 5 5700 44.3 122.0 546 53 Alaska Hwy.,Mi.720,Yukon 83 25 1.89 5 3200 60.0 129.1 558 54 Graves Mt.,Wash. 75 17 1.80 5 5200 48.2 116.0 561 55 Lake Louis,Alta. 65 40 1.86 5 5680 51.3 116.3 563 56 Alder Trail,Glacier N.P* Mont. 70 40 1.76 5 4200 48^3 114.0 574 57 Mt.Idaho,New Denver,BC. 57 50 2.02 5 6500 50.5 118.4 574 58 Alaska Hwy.,Mi.735,BC. 59 Mt .Edith Cavell,Jasper 106 20 2.01 5 3000 60.0 128.0 580 N.P., Alta. 67 40 1.86 6 6000 53.0 112.0 587 60 Upper Pine River Valley, Alta. 65 25 1.88 5 3200 55.5 122.8 598 61 Belknap Crater,Ore. 90 50 1.95 6 5300 44.3 122.0 607 62 Nagle Cr.,North Bend Hwy. * BC. 67 40 1.35 6 4000 52.0 118.5 610 63 Nisqually,Wash. 75 45 1.82 6 1000 46.5 123.2 620 64 Beaverdell Range,BC. 63 25 1.89 7 2500 49.3 118.3 644 65 wallowa Mt.,East Peaks,Ore.67 35 1.86 6 6000 45.3 116.9 646 Table 2. BASIC ELEMENTS OP STATISTICAL ANALYSIS ON STOMATAL DISTANCE . X SD Range Max: Min: X l * 2 h X4 X5 X6 Y l T 2 8 .767692E+01 4.690769E+O1 1.761076E+02 5.053846E+01 4.095338E+03 5.1098460E+02 5.221999E+02 5.179230E+02 2.015882E+01 3.330996B+O1 1.746707B+00 3.657513B-00 1.624364E+03 4.8534750E+01 4.6846203+01 - 4.526736E+01 sq.tam 148 47 inch 140 8 2 . 2 0 1.35 unit 6.5 4 .0 feet 8000 1000 degrees 60.30 42.00 CD. 643 448 CD. 649 449 V T2 5.200615E+O2 4.551996E+01 CD. 646 448 l l 2 i A Stomatal frequency per sq.millimeter Precipitation Needle width Mean of stomatal distances Elevation Latitude X : Mean of variables SD: Standard deviation. 1 unit 0 , 0 2 millimeter CD.rcombined items,ex- planation on page 2 0 . T.: Distance between two stomata in a row line,first measurement Tj: Second measurement TABLE 3 CORRELATION COEFFICIENTS OF STOMATAL DISTANCE. Row X j ^ X̂  X^ X^ x 6 T I - 0 - 0.3759 x O.I698 -0.4447 x 0.0444 -0 .0153 x II 0.3759 x - 0 - 0.1121 - 0 . 2 0 7 5 0.4533 x - 0 . 4 6 3 7 x - 0 . 3 9 3 3 x I H O.I698 0.1122 - 0 - - 0 . 0 0 0 9 0.1825 -0.0715 0.1939 IV -O.bktil x - 0 . 2 0 7 3 - 0 . 0 0 0 9 - 0 - - 0 . 0 9 9 8 -O.069U 0.7550 x v o.o444 0 .4533 x 0.1825 - 0 . 0 6 9 4 . - 0 - - 0 . 6 1 9 1 x -0.0I136 VI - 0 . 0 1 5 2 -0.4637 x -0.7152 -0.7152 -0 .6191 x -0= -0.0127 VII -0.5051* x -0.3933 x 0.1939 0.755C x - 0 . 0 4 3 1 - 0 . 0 1 2 7 - 0 - The correlation coefficients' values in the Table: r » .244 . 0 5 indicates a negative slope r » .318 tt+" Indicates positive slope .01 -..-'^0- the factors to which the other factors are correlated x significantly correlated at .01$ level TABLE 4. REGRESSION COEFFICIENTS OF CLIMATIC FACTORS Elimination of T, stomatal distance The variable to be omitted Constant term ( a ) 6.686ll|2E+01 1.059740E+02 6.185729E+01 1.5U535E+02 9.328281E+01 4.5l5440E+01 Residual Variance (SEE) % 6.577782E+02 .71231080 71.2J 6.578876E+02 .70730190 70.73 6.5652U8E+02 .70295760 70.29 7.773202E+02 .61.2iih2ii0 64.21* 7.943285E+02: .62867110 62.87 9.09Ui79E+02 .57008730 57.01 SEE b, RES.V. TV-5 9.40 *5 45.15 + 9.4o x, EXPLANATION OP TABLE 5. ANATOMICAL CHARACTERISTICS OP NEEDLE Epidermal tissues are categorized as follows: A: well-developed hypodermal cells,two or more layers B: developed, "but thin,broken layers C: ill-developed layers 0 : no hypodermal cells under the epidermis 1 : hypodermal cells, are only under the ab- axial epidermis 2 : hypodermal cells are at the angles of needle wings . 3 : hypodermal cells are at the angles and under the abaxial epidermis 4 : continuous or alternately broken hypodermal layers line the mesophyll in two or more rows Mesophyll tissue can be palisade-like formed quadrangular cells and closely attached to each other, and spongy parenchyma involved numerous intercellular .spaces between them.. Size of resin canal is inticated as: I : 150-200 micron large cavity II : 100-149 do III : 70-99 do Endodermis: Inner sheath means the perycicle, which i s i l l -defined in most cases Transfusion tissue: Percentage means the ratio of parenchyma and tracheid elements in the abaxial part or half of the stele 1/2 - phloem and xylem band occupies the upper half of the stele l/3 - the band occupies the upper one/third space of the stele - 78 - Table 5. ANATOMICAL CHARACTERISTICS IN THE NEEDLE CROSS-SECTION OF SOME SELECTED ALPINE FIR SAMPLES. Samples are ranked as for the stomatal distance No. of sample LOCATION Yj = stomatal distance Yg = bract and scale length difference (hundredth of mm.) EPIDERMAL TISSUE MESOPHYLL TISSUE ' RESIN CANAL ENDO- DERMIS TRANS- :USION TISSUE VASC verti- cal ULAR BUNDLE ROWS horizontal NOTES xylem phloerr 1 Garibaldi Lake Trail, BC Y s . 451 Y B - 50 A - 4 3 layers Rounded palisade spongy parenehl- ma 1 some times conti- nuous 70% T I/O® 13 5 5 Xylem and phi located in the upper half of the stele. 3 Logan Pass, Glacier NP. Montana Ys - 461 YB - 690 B-3 Rounded palisade spongy parenchi- ma 1 different cavity size one cont. ill-de- veloped Inner sh 60% T 1/3 @ 17 5 7 11 Downie Creek, B.C. Ys =481 YB- - C - 3 Palisade like layers dense tissue wi thou t ai r spaces II Broken 70% T 1/3 16 5 6 20 Forbidden Plat., V.I. Ys =488 YB = 650 B-3 Palisade like layers dense tissue II cont. ill-d. inner sheath 70% T 1/2 > 20 5 5 29 Mt. Hood, Ore. Y S = 500 Y B . - A - 4 Palisade like layers abundant chloro- phyll II 3 eplth. layers cont. ill-d. Inner sheath 70% T 1/2 13 4 6 37 Yoho NP., B.C. Ys " 515 YB-480 C - 3 rounded palisade spongy paronchi- ma II differen cavity size , Broken 70% T ,1/3 15 4 8 Conspicuous!) separated bundles 38 Alaska Hwy., Mi. 305, B.C. Y S = 523 Y r - C - 3 Palisade like layers dense tissue II Cont. 70% Par. 1/2 11 5 7 41 Radium Hot Spring, Alaska Hwy., Mi .480 BC Y S = 527 YB= - C - 0 Palisade like layers spongy parenchi- ma Il-lll 0 - 4 canals Broken 80% Par. 1/2 12 4 8 Medial and marginal can. Sometimes one epithelial cell sheath only 54 Graves Mt.,.Wash. Y s „ 561 YB-340 B -3 Palisade like layers spongy parenchi- ma II outer sheath * Broken large uneven cells 60% Par 1/2 17 5 6 * large 58 Alaska Hwy., MI.735BC Ys = 580 YB= 170 A - 4 Rounded palisade spongy parenchima II Cont. 80% T ,1/2 17 6 7 59 Mt. Edith Cavel Jasper NP., Alta. Y S = 587 Y B - - C - 2 Palisade like layers spongy parenchi- ma III one sheath • only Broken 80% T 1/3 13 5 6 64 Beaverdell, B.C. Y S =644 YB = 550 B-3 Palisade like layers spongy parenchi- ma III * • Broken 50% Par. 1/4 16 5 7 •Sometimes one additional epithelial sheath 65 Wallowa Mt., Ore. Y S = 646 YB = - A - 4 2-3 alter- nated lay- ers Rounded palisade cells spongy parenchi- ma II • Cont. 80% Par. 1/2 13 5 6 *3 epithelial sheaths MEAN OF THE COLUMNS OF BUNDLE; . 15 MEAN OF HORIZONTAL XYLEM ROWS; 5 MEAN OF HORIZONTAL PHLOEM ROWS; 7 - TABLE: 6. GEOGRAPHICAL DISTRIBUTION OF ALPINE FIR SAMPLED, AND SOME CHARACTERISTICS OF RESIN CANALS IN THEIR NEEDLES No. of sample Location Lat. and - Long. (DejsTees) Elevation (feet) Resin canal Micron Diameter Height Micron 22 Crater Lake, Ore. 43°02,N - 122°04,W 7500 170.0 170.0 2.14 20 Siskiyou Mt., Ore. 42°02,N - 123°03,¥ 7400 235.0 I63.O 2.66 53 Mt. Edith C a v e l l , A l t a . 53°07,N - ll5°24,W 6000 211.3 237.2 2.34 12 Mt. Tusk, Garibaldi, B.C. 50°O5,N - 12U°10,W 5600 . 188.3 195.0 1.43 7 Forbidden P i t . , B.C. . U9°2U,N - 126°00,¥ 5ioo 187.0 151.3 1.53 13 Scot Lake Mt.jWash,,Ore. 44°12,N. = ll5°U8.,w.' 45oo 177.8 ; 196.1 1.27 4 8 Liard Hot Spring, B.C. 59°05,N - 126°54,W 2800 96.3 186.3 3.87 60 Bear Lake, B.C. 54°42,N - 122°30,w 2500 1 2 8 . 8 181.3 0.93 4 7 Alaska Hwy. Mi.5oo. B.C. 59°12,N - 127°06,W 2300 112.5 97.5 1.99 2U Hazelton, B.C. 55°12,N - 125°l8,W 1600 106.4 U 4 . 4 1.52 Note: Each sample represents 60 needles from a sample unit of three t r e e s . Total: 600 needles f o r entire t a b l e . Sx « Standard deviation of the resin canal position with regard t o the centre of stele and the edge of needle. - 80 - Table: 7. MEASURED DATA OF THE SCALE AND BRACT OF ABIES LASIOCARPA. X̂  :. Length of scale, aver age of twenty samples,tenth of mm. X 5: Length of bract,average of twenty samples,tenth of mm. Xg: Meandifference of 20 scale and bract length,millimeter. Y : Sum of the differences in length of the scale and bract . of ten randomly selected samples.-.Hundredth of millimeter. NOg. : Code number of stomatal specimens from Table 1. No Nog Location X5 x 6 Y 1 1 Garibaldi Lake Trail,BC. 109 102 1 5 5 2 2 Hurricane Hill,Olympic N.P., Wash. 115 106 1 90 3 2 Hurricane Hill,Olympic N.P.,-Wash. 123 112 1 100 4 — Hurricane Peak,Olympic N.P. Wash. 105 90 2 150 5 58 Alaska Hwy.,Mi.735,BC. 81 64 2 170 6 20 Forbidden Plat. V.I. 128 111 2 175 7 18 Parsnip River,BC. 106 87 2 185 8 45 Mt.Rainier,Wash. 104 83 2 195 9 25 Mt.Rainier,Wash. : 105 85 2 195 10 57 Mt.Idaho,New Denver,BC. 102 82 2 210 11 5 Arrowsmith Mt.,Peak,V.I. 136 113 2 240 12 28 Aleza Lake,Lower Elevation,BC132 107 2 245 13 4 Black Tusk,Garibaldi Prov.P. BC. 139 108 4 300 14 4 Black Tusk,Timberline,BC. Mt.Bonaparte,Wash. 142 110 3 315 15 49 143 110 3 320 16 54 Graves Mt.,Wash. 130 96 3 340 17 55 Lake Louis,Alta. 135 101 4 340 18 39 Alaska Hwy.,Mi.l60,BC. 157 121 3 360 19 19 Mt.Rainier,Timberline,Wash• 147 109- 380 20 28 Aleza Lake,Higher Elevation, BC. 135 93 2 380 21 17 Arrowsmith Mt.,Hill,V.I. 223 183 4 400 22 24 Hurricane Ridge,Olypic N.P. Wash. 154 115 5 435 23 — Arrowsmith Mt.,V.I. .158 115 4 4 4 3 24 5 7 Mt.Idaho,New Denver,Lower Elevation,BC. 147 9 9 5 480 25 37 Kicking Horse Pass, Yoho N.P. BC. 1 4 5 9 5 5 4 8 5 Table 7.Cont. - 31' - No . location x 5 x 6 Y 26 36 Pendelton Bay,BC. 132 77 5 500 27 64 Beaverdell Range,BC. 152 103 5 550 28 - Hurricane Ridge„Olypic . N. P. t ,Wash. 170 115 6 555 29 — Jackson Glacier,Glacier N.P.,Mont. 154 92 6 620 30 23 Bear lake,BC. 173 109 6 625 31 25 Mt.Rainier,Wash. 177 113 9 650 32 20 Forbidden Plat.,BC. 201 136 7 655 33 4 Black Tusk,BC. Hurricane lodge,Olypic N.P. 187 121 7 660 34 • •- *162 Wash. 103 6 665 35 — Kokanee Galcier Prov.P.,BC. 177 110 7 670 36 — Jackson Glacier,Glacier N.P.,Mont. 163 93 7 695 37 39 Alaska Hwy.,Mi.l60,BC. 193 118 8 750 38 31 Willson Cr.,New Denver,BC. 180 99 7 805 39 50 Divide Cr.,Glacier N.P., Mont. 190 108 8 815 40 56 Alder Trail,Glacier N.P., Mont. 218 130 10 890 Result of s t a t i s t i c a l analysis: Mean of Y : 428 hundredth of millimeter Standard deviation: 32 hundredth of millimeter F = 131.576, greater than the F in the Table at 5 7° and 39 DF. The differences between the value of Y are highly significant in several cases which are expressed in the diagram of the Duncan's test. Figure 7. ' TABLE 8. MEASUREMENT DATA ON SEEDS AND CONES OF ALPINE FIR. ra 's E E D AXIS 8 V.o Location WEIGHT • WING . SIZE. LENGTH GR/1000 (cm'2-) mm. 5 Alaska Hwy.,Mi.735,Yukon — • O.lSxxx 7 Parsnip River,BC. — 0.82 - 9 Mt.Rainier,Washington - 0.15xxx - 12 Aleza Lake,3C.Lower Elevation. . 5.94 0.64 51 18 Alaska Hwy.,Mi.l60,BC. -' 1.40xxx - 20 Aleza Lake,BC.Higher Elevation. 5-Q2 0.75 22 Hurricane Ridge,Olympic UP.,Wash. • - 0.5.0 - 25 Kicking Horse Pass,Yoho NP.,BC. . -. 0.82 - 26 Pendleton Bay,BC. • 6.21 0.40 43x 35 Kokanee Glacier Prov.P.,BC. . • 0.75 76x 38 Willson Cr.,New Denver,BC. 8.93 l.o5xxx 32xx LEGEND: ' x PROBABLY; SIGNIFICANT does not exceed 1.0% probability l e v e l , xx SIGNIFICANT f a l l s ..between 1.0.'and 0.1% probabil ity l eve l , .xxx HIGHLY SIGNIFICANT exceeds 0 .1$ probabil ity l e ve l . Fisher's Table was.used from SNEDECOR (1956) 1 3D Range Max: Min: . . TABLE 9. BASIC ELEMENTS OP STATISTICAL ANALISIS ON SCALE AND BRACT h x 2 h h h x s T i 6.605000E+01 5.032000E+01 4.659100B+O3 1.4825O0E+O2 1.056000E-HD2 4.350000E-O0 4.260000E+01 3.995763E+01 3.306113E+00 1.367355E+03 3.291617E+01 1.904757E+01 2.402456E-00 2.229039E+O1 inch 140 17 degrees 60.00 47.50 feet 8000 2500 mm 22.3 8.1 . mm 18.3 6.4 nun • 10 1 1/100 mm '890. 7 0 Y i + r2 4.292500E+01. 4.276250E+01 2.300210E+O1 2.256187E+01 1/100 mm 890 40 1/100 mm 890 55 X : X 2 : Precipitation Latitude Altitude Length of scale Length of bract Average of the differences be-tween X^and X̂ X : Mean values of the variables 3D: Standard deviation T, Total of difference between the length of scale and bract firs t measurement The same as second measurement TABLE 10. CORRELATION COEFFICIENTS OF SCALE AND BRACT X X ,51238391. -.5123839IX .52091683* .05976233 .40767108x -.17006502 -.24360658 -.66971426 x -.09742801 -.19494580 -.06546816 -.00383624 .52091683 x «.66971426 x ~Q« -.05264150. .08963397 -.02668998 '-.13810740. .05976233 -.09742801 -.05264150 «0« .75863036'x .82373938 x .83568909" .40767108x -.19494580 .08963397 .75863036- :-0« .30683411 .28103459 ».17006502 -.06546810 -.02668998 .82373938 * .30683411 «0« .96162277 x •.24360658 -.00383624 -.13810740 .835 68909 x .28103459 ,96162277 X -0- "-" indicates a negative slope -0- the factors to which the other factors are correlated x significantly correlated at .01 level Correlation coefficients: R = .312 .05 R = ,403 .ol The variable to be omitted TABLE 11. REGRESSION COEFFICIENTS OF THE VARIABLES Elimination of Y,difference of the length between scale and bract. Constant term Residual variance (SEE) " & X X X X X X 9.389189E400 1.581377E401 -3.536110E-01 -5.480186E+00 -5.731713E400 3.478677E^00 Res.Var. REMARKS: SEE = n-4 4.126255E+00 3.066287E+01 3.024501E4ol 3.078240E-&01 3.7225163+01 3.932972E401 •99314110 .94748580 .94667800 .94418000 ,93062180 .92471830 X, - Precipitation,inches X2-_ Latitude, degrees X 3 -Alt itude,feet Length of scale,mm X5-Length of bract,mm X&* Individual mean of sample,millimeter 99,3 94,7 94,6 94,4 93.1 92.5 - 36 - FIGURE: 1. DISTRIBUTION OF ALPINE FIR STANDS SAMPLED ON THE WEST.COAST OF NORTH AMERICA FIGURE 2. REGRESSION LINE OF PRECIPITATION ON STOMATAL DISTA NCE (Figures beside the dots indicate the elevation in hundreds. Yon the line represents the correlation equation.) STOMATAL DISTANCE Y 6 5 0 - 6 0 0 - 55 0 - 5 0 0 - 4 50 Low frequency 0.12 millimeter 6o° 10 3o° 52 o 32 o 3o2 6<P 56 o CO 55 o High frequency, 1 1 i 0.09 millimeter I r 4 8 o TO 20 30 4 0 50 100 150 INCH P R E C I P I T A T I O N Figure 3. Comparison between means of the distances of; stomata on the lower surface of needles as Duncan's multiple range t e s t , (LSD). i i i • i i i r ~ i i i i i i i — i i — i — i — i — i — i — i — i — i — i — i — i — i - 4 5 4 461 4 6 6 4 6 8 4 8 0 481 4 8 3 4 8 5 4 8 7 4 8 7 4 9 0 4 9 2 4 9 4 4 9 8 5 0 0 5 0 6 510 514 515 5 2 4 5 2 6 5 2 9 5 3 2 5 3 9 5 4 0 5 4 4 5 5 8 5 6 4 5 7 4 5 8 7 6 0 7 619 6 4 6 4 5 7 4 6 6 4 6 8 4 7 5 4 8 0 4 8 2 4 8 5 4 8 6 4 8 7 4 8 8 4 9 1 4 9 4 4 9 5 4 9 9 5 0 4 5 0 7 514 514 5 2 3 5 2 5 5 2 8 5 3 0 5 3 8 5 3 9 5 4 2 5 4 6 561 5 7 4 5 8 0 5 9 8 6 0 9 6 4 4 "> i i i i i i i i i — i — i — i — i — i — i — i — i — i — i — i — i — i — r T — i — r — r - 391 411 4 2 2 4 3 0 4 3 6 4 4 1 4 4 5 4 4 8 4 5 2 4 5 4 4 5 4 4 5 4 4 5 5 4 5 5 4 5 6 4 5 8 4 5 9 4 6 0 461 4 6 2 4 6 3 4 6 4 4 6 6 3 7 5 4 0 2 4 1 6 4 2 6 4 3 3 4 3 9 4 4 4 4 4 7 4 5 0 4 5 3 4 5 4 4 5 4 4 5 4 4 5 5 4 5 6 4 5 7 4 5 9 4 5 9 461 4 6 2 4 6 3 4 6 3 4 6 5 4 6 6 - — 2 7 2 9 2 9 3 0 3 0 31 3 1 3 1 3 2 - - 3 2 3 3 - - 3 3 3 4 - - 4 6 6 - 3 4 J I I I I I J 1_ J 1_ J I ' » ' J—I I I I I 1 1 I I I i i i - I l _ 55 - J — i — i — i — i i ' i ' 60 65 Legend:Sample Y :t(r.oi) SSR 40 10 15 20 25 30 35 -No. of specimens SAMPLE NO. - V a r i e t a l means ranked i n order; stomatal distances - Shortest si g n i f i c a n t ranges from Table r- •— - tfr.oi). x standard error of a v a r i e t a l mean, . S / f 45 50 - 39 - FIGURE; U . DISTRIBUTION PATTERNS OF ECTODERMAL CELLS UNDER THE NEEDLE EPIDERMIS'' OF THE ALPINE FIR IN. ITS CROSS-SECTION. Each l i n e i n the needle cross-section represents a single hypodermal layer. There are discontinuous ( l ) , continuous (2) and mul t i p l i e d (3) l a y e r s . No. Aspect Elevation No'. Aspect Elevation . Ft. Ft. Figures at the l e f t of the cross-sections refer to the code number i n Table 1 . - 90 - Figure '..5. SHAPE AND SIZE OF SEEDWINGS SAMPLED FROM VARIOUS STANDS' OF ALPINE FIR., (life size ) 61. Q 24 Q 45 Q 23 37 - Q 62 Q 58 & 36 Q 28 Q 28 Q 31 <? Figures at the above left indicate the sample number as coded in Table 7. Figure? 6. DISTRIBUTION OF VASCULAR STRANDS IN THE SCALE OF ALPINE FIR . (front view,tangential section) Figure 7. Comparison between means of the difference i n the length of scale and bract. Duncan's multiple range test (LSD). tr.oi S S R z < U_ LLJ o £ CO Q D- U J 3 »-O < °*" n ° o to C O < T 1 1 — r i — i — i — n r ~i—r T — r T i—r i — r i — r T ~ \ — i — i — r ~i—r T* " i — r 55 100 170 185 195 240 300 320 340 380 400 443 485 550 620 650 660 670 750 815 90 150 175 195 210 245 315 340 360 380 435 480 500 555 625 655 665 695 805 890 — 399 417 430 437 443 449 452 455 458 459 460 460 461 462 462 463 464 464 465 382 410 425 434 441 446 451 454 457 459 459 460 461 461 462 463 462 464 465 465 7.9 8.2 8.5 8.6 8.7 8.8 7.5 8.1 8.4 8.5 8.7 8.8 I— 8 . 9 - -9.0- •9.1- H 9.2- _i i i_ - i i i i i i _ - i — i — i 1 i i i i _ 10 15 Legend: 20 25 SAMPLE NO. - i — i — i i i • 30 I I u 35 40 Samples - No. of specimens Y - V a r i e t a l means ranked i n orderj •" '• ; distance between'scale and bract length tfri0l) - Shortest significant ranges from Table SSR u(r.OI) x standard error of a v a r i e t a l mean Figure 9. - 93 - Flgur* 11,  Figure 14. Code No. from Table 7. 5,37,60, 7,26, 20,12,25,17,38,24 24,38, 3, 8,22, 2, 31,31,19, 9,31 Legendt £ epidermis 8 subsidiary cells 0 guard cells CH substoaaatal chamber Figure 17, x 1500 Figaro 19, x 140 - 97 - Figure 21. (Abaxial sarf&ce) x 50 E epidermis eel]s B bundle Rc r e s i n crnal Figure 22. { Adaxial surface ) x 100 Figure 23. ( old needle ) x 50 well—. developed • nd well- separated bundles Figure 25. ( old needlo ) x 80 Figure 24. ( old needle ) x 80 Figure 26. ( young needle ) x 60 99 - L: m I .1,1 ^^a^4H- s Figure 27. X 400 Figure 28. Legend: AL Alburdnous c e l l Xy X/leia Ph Phloem x 350 refusion tissue En Etxoc'emis Pa Parenchyma cells Ta Tracheids Ex Bundle sheath extension,through broken endoderais Figure 30. x 400 - 101 - Figure 32. 350 - 102 - • 0 **» 6 Figure 3 3. x 0,5 Legendt L Lenticel B Resin blister


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