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Resistance to water uptake : a comparison of ponderosa pine and Douglas-fir seedlings and an investigation… Standish, J. T. 1983

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RESISTANCE TO WATER UPTAKE: A COMPARISON OF PONDEROSA PINE AND DOUGLAS-FIR  SEEDLINGS  and AN INVESTIGATION INTO THE EFFECTS OF PLANTING  by 3.T. S t a n d i s h B.S.F., The U n i v e r s i t y of B r i t i s h C o l u m b i a ,  1977  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES Department o f S o i l S c i e n c e  We a c c e p t t h i s t h e s i s as c o n f o r m i n g to t h e r e q u i r e d s t a n d a r d  THE UNIVERSITY OF BRITISH COLUMBIA September, 1983 C)  -John Thomas S t a n d i s h , 1983  E-6  In p r e s e n t i n g  this thesis i n p a r t i a l  f u l f i l m e n t o f the  requirements f o r an advanced degree a t the U n i v e r s i t y o f B r i t i s h Columbia, I agree t h a t t h e L i b r a r y s h a l l make it  f r e e l y a v a i l a b l e f o r reference  and study.  I further  agree t h a t p e r m i s s i o n f o r e x t e n s i v e copying o f t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by t h e head o f my department o r by h i s o r h e r r e p r e s e n t a t i v e s .  It is  understood t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l gain  s h a l l n o t be allowed without my  permission.  J.T.  Department  of  S o i l Science  The U n i v e r s i t y o f B r i t i s h 2075 Wesbrook P l a c e Vancouver, Canada V6T 1W5 Date  (2/79)  Standish  29 S e p t e m b e r 1983  Columbia  written  ABSTRACT  Resistance and  to  water  coastal  menziesii)  ponderosa pine (Pinus  Douglas-fir (Pseudotsuga  seedlings  Ohm's Law  in  uptake  was  studied  menziesii  ponderosa  (Mirb.)  Laws.)  Franco  var.  in a controlled environment using  an  analogy to l i q u i d phase water flow for calculating resistance  from measurements of s o i l  and  needle water potential and, uptake rate.  This study compares resistance in ponderosa pine to s i m i l a r l y measured resistance  i n Douglas-fir  from  another  study.  uptake in planted Douglas-fir seedlings was  Ponderosa pine the  range of  Weibull  shows v i r t u a l l y soil  water  function was  constant  Resistance  to  water  also investigated.  needle water potential through  potential where uptake occurs.  A  modified  found to give good o v e r a l l results for predicting  uptake as a function of s o i l water potential using nonlinear  regression  techniques.  Generally,  uptake rates are high compared to Douglas-fir at s o i l water  potentials greater than about -0.5 soil  dries.  constant  and  greater  than  increases  -1.5  The  but decrease more rapidly as the  to water uptake appears to in  much less than MPa.  rapidly; at  about equal. potential  Resistance  MPa  At -2.0  increase  in ponderosa pine  Douglas-fir  lower  MPa,  soil  at  water  resistances  soil  be  be  potentials  potentials  resistance  for the  two  largely a result of  plant and s o i l - r o o t contact resistance.  less  water  in resistance below - 1 . 5 MPa may  more or  species  are  of s o i l water increases  in  - i i i-  Resistance  in  experimental which had  planted  Douglas-fir  treatments.  The  lifting  seedlings.  and  H a l f of the p l a n t e d  seedlings  showed  statistically in  soil  physical  decreased totally  identical  soil account  seedlings.  in  their it  resistance i s responsible  is  using  consisted  of  P l a n t i n g was  randomly  three  seedlings  simulated  s e e d l i n g s were randomly a l l o c a t e d to  the  of s u b j e c t i n g  and  among  seedlings  greater  (by  to v i b r a t i o n .  about  vibrated seedlings,  treatments,  planted lower inferred  f o r the  selected  by of  to the c o n t r o l s e e d l i n g s .  resistance  Therefore  a  studied  group  Planted  properties  for  15 months.  considerably  r e s i s t a n c e than c o n t r o l s .  was  treatment  re-potting  t h i r d treatment which c o n s i s t e d  Planted  control  grown i n pots f o r about  carefully  seedlings  and  vibrated  resistance that  lower t o t a l  is  reduced  similarity  doubtful  seedlings  compared  times)  however, were  Because of it  six  that could  to  planted  soil-root  contact  resistance.  - iv -  TABLE OF CONTENTS  Page ABSTRACT  i i  TABLE OF CONTENTS  iv  LIST OF TABLES  vi  LIST OF FIGURES ACKNOWLEDGEMENTS  vii '  ix  INTRODUCTION  1  CHAPTER 1: RESISTANCE TO WATER UPTAKE IN PONDEROSA PINE COMPARED TO DOUGLAS-FIR  5  Introduction  6  Methods and Materials  7  Seedlings and Soil  8  Experimental Environment  10  Soil Water Potential  11  Needle Water Potential  13  Uptake  15  Resistance  17  Comparison of Resistance  18  Results  19  Discussion  34  Summary and Conclusions  41  -  V  -  Page CHAPTER 2: RESISTANCE TO WATER UPTAKE IN PLANTED DOUGLAS-FIR SEEDLINGS  43  Introduction  44  Methods and Materials  45  Results  51  Discussion  64  Summary and Conclusions  68  CONCLUSIONS  70  LITERATURE CITED  76  APPENDIX I:  Spectral Irradiance in the Experimental Environment  86  APPENDIX II:  Statistical Summary for Regression of Needle Water Potential as a Function of Soil Water Potential for Ponderosa Pine  89  APPENDIX III:  Calculations for Uptake and Resistance for Ponderosa Pine  91  APPENDIX IV:  Observed Uptake Rates in Ponderosa Pine Compared to Average Uptake Curves  100  APPENDIX V:  Uptake and Needle Water Potential Data for Ponderosa Pine  103  APPENDIX VI:  Data for Douglas-Fir Experiment  107  APPENDIX VII:  Statistical Summaries for Resistance in Douglas-Fir Seedlings  112  APPENDIX VIII:  Stomatal Resistance Measurements and Estimated Maximum Transpiration Rates  116  APPENDIX IX:  Seedling and Water Pathway Dimensions  119  APPENDIX X:  Soil Physical Properties  123  APPENDIX XI:  Estimates of Soil Resistance  126  - vi -  L I S T OF TABLES  Page Table 1:  Soil Properties  Table 2:  Summary of Average Seedling and Water Flow Pathway Dimensions for Ponderosa Pine Seedlings  Table 3:  Average Coefficients for Uptake Equations  Table ki  Kruskal-Wallis Test for Differences in Resistance to Water Uptake for Douglas-Fir (F) and Ponderosa Pine (Py) Soil Characteristics and Water Pathway Dimensions for Ponderosa Pine and Douglas-Fir  Table 5: Table 6:  Seedling and Water Pathway Dimensions for Douglas-Fir Seedlings  18 23  34 39 51  - vii-  LIST  OF F I G U R E S  Page  Figure 1  Needle Water P o t e n t i a l vs S o i l P o t e n t i a l f o r Ponderosa Pine  Water  Figure 2  Water P o t e n t i a l D i f f e r e n c e vs S o i l Water P o t e n t i a l f o r Ponderosa Pine  22  Figure 3  Uptake i n Ponderosa P i n e per U n i t Root Area as a F u n c t i o n of S o i l Water P o t e n t i a l - R e g r e s s i o n Model I  24  Figure 4  Uptake i n Ponderosa Pine per U n i t Root Area as a F u n c t i o n of S o i l Water P o t e n t i a l - R e g r e s s i o n Model II (Modified Weibull Function)  25  Figure 5  Average R e s i d u a l s Equation Model I  f o r Uptake  27  Figure 6  Average R e s i d u a l s f o r Uptake Equation Model I I  28  Figure 7  R e s i s t a n c e t o Uptake per Unit Root Area f o r Ponderosa Pine (Uptake Model I)  30  Figure 8  R e s i s t a n c e t o Uptake per U n i t Root Area f o r Ponderosa Pine (Uptake Model I I )  31  Figure 9  R e s i s t a n c e t o Uptake per U n i t Root Area f o r Ponderosa Pine (Uptake Model I I ) , D o u g l a s - f i r on S i l t Loam and D o u g l a s - f i r on Loamy Sand  33  F i g u r e 10  Needle Water P o t e n t i a l vs S o i l Water Potential f o r Control Douglas-fir Seedlings  52  F i g u r e 11  Needle Water P o t e n t i a l vs S o i l Water Potential f o r Planted Douglas-fir Seedlings  53  20  - viii -  Page  Figure 12  Needle Water Potential vs Soil Water Potential for Planted and Vibrated Douglas-fir Seedlings  54  Figure 13  Uptake Rate for Control Douglas-fir Seedlings  56  Figure 14  Uptake Rate for Planted Douglas-fir Seedlings  57  Figure 15  Uptake Rate for Planted and Virbrated Douglas-fir Seedlings  58  Figure 16  Resistance to Uptake for Control Douglas-fir Seedlings  60  Figure 17  Resistance to Uptake for Planted Douglas-fir Seedlings  61  Figure 18  Resistance to Uptake for Planted and Vibrated Douglas-fir Seedlings  62  - ix -  ACKNOWLEDGEMENT I owe a debt o f g r a t i t u d e t o Dr. T.M. B a l l a r d ( P r o f e s s o r , Department o f S o i l S c i e n c e and F a c u l t y o f F o r e s t r y ) f o r h i s guidance and encouragement during  my g r a d u a t e s t u d i e s  producing  this  thesis.  and f o r h i s p a t i e n c e  Dr. T.A. B l a c k  (Professor,  S c i e n c e ) d e s e r v e s s p e c i a l t h a n k s , not o n l y editorial sions.  and h e l p f u l a d v i c e i n Department o f S o i l  f o r h i s sound t e c h n i c a l and  a d v i c e , but a l s o f o r many e n t h u s i a s t i c and s t i m u l a t i n g d i s c u s -  The s u p p o r t and encouragement o f t h e o t h e r members o f my commit-  tee,  Dr. L.M. L a v k u l i c h  Soil  Science)  (Professor  and Dr. C A .  Rowles  and Department (Professor  Head, Department o f  E m e r i t u s , Department o f  S o i l S c i e n c e ) , i s a l s o g r a t e f u l l y acknowledged.  The b r e v i t y r e q u i r e d  i n t h i s acknowledgement p r o h i b i t s s i n g l i n g out a l l  of t h e f a c u l t y members and s t u d e n t s who p r o v i d e d  encouragement, h e l p f u l  a d v i c e and many s t i m u l a t i n g i d e a s d u r i n g my graduate program; however, a few i n d i v i d u l a s need t o be mentioned.  F.M. K e i l l i h e r  Department  invaluable  of S o i l  assistance.  Science)  Dr. M.W.  provided  Sondheim  (now w i t h  ment), Dr. J.P. Demaerschalk ( P r o f e s s o r ,  (Ph.D.  advice  candidate,  and t e c h n i c a l  t h e B.C. M i n i s t r y o f E n v i r o n Faculty  o f F o r e s t r y ) , Dr. A.  Y  Kozak ( P r o f e s s o r Greig cal  and A s s o c i a t e  Dean, F a c u l t y  o f F o r e s t r y ) , and Dr. M.  (UBC Computing C e n t r e ) gave e x t r e m e l y h e l p f u l a d v i c e  matters.  on s t a t i s t i -  -  X  Thanks are extended to Dr. W. Herman (Pacific Soil Analysis Inc.) and M.  Goldstein  (Soilcon  Laboratories Ltd.)  for  their  assistance  and  provision of laboratory f a c i l i t i e s .  I also must thank Westwords for very prompt and efficient word processing services and N. Cukor, V. Kwong and N. Smith of Talisman Graphics for producing the final figures.  The B.C. Ministry of Forests provided tree seed and seedlings and their Research Branch provided assistance and advice in the early stages of my studies.  In particular,  I express  my  appreciation to R.L.  Schmidt  (former Director, Research Branch) and Dr. T.E. Baker (Manager, Ecology and Earth Sciences, Research Branch).  Finally, I offer many thanks to my  family, Sheila, Matthew, Jack and  Emily for thier patience, encouragement and moral support.  Research for this thesis was  partially supported by a grant from the  Natural Sciences and Engineering Research Council of Canada.  - 1 -  INTRODUCTION  The importance of water to plants has been long recognized and ic  enquiry into plant water relations  years ago (Hales,  1727).  began over two hundred and  of protoplasm, as a solvent  substances,  and as  a chemical  stomata,  for  cell  in  leaf  and  1969).  Ecologically,  sensible  and latent  temperatures  (Gates,  distribution  of  affected  for gases, minerals and growth  reagent (e.g., in photosynthesis  hydrolysis of starch to sugar); important  it  enlargement  also maintains c e l l and growth,  (Bidwell,  1974;  water  important  in  heat,  is  thus  1980;  affecting  Oke, 1978).  vegetation  over  t u r g i d i t y which  Salisbury  partioning  soil,  and in  opening and closure  movement  plant  and  geographic  areas  or  energy  into  and ambient  regions  is  of  Ross,  At a much broader scale,  air the also  by the a v a i l a b i l i t y of water, as i l l u s t r a t e d in various vegeta-  tion and ecological mapping and c l a s s i f i c a t i o n 1972;  fifty  Water is important physiologically as a major  constituent  is  scientif-  Damman,  1979;  Franklin  and  schemes (e.g. see Burger,  Dyrness,  1973;  Holdridge,  1947;  Krajina, 1969; Merriam, 1898; P f i s t e r et al., 1977; Pojar, 1983; Walter, 1973).  Moisture affecting and of  has  been widely  seedling  Cieary,  1978).  recognized as a major environmental  s u r v i v a l and growth (Waring, Hinckley et al. (1978),  water status in forest  1970;  variable  Greaves, Hermann  in their l i t e r a t u r e  trees, point out that the growth is  limited by either an excess or a scarcity of water.  review  generally  - 2 -  The  water  from  the s o i l  water the  balance  flow  leaf,  model  and l o s s  from under  based  current.  of a plant  the s o i l  This  an  to the r o o t and through  Hiilel  and S l a t y e r  Liquid  widely  was  proposed  used  by many  by  authors  of  simple  electrical  van den  Honert  including  Cowan (1975)  water  (Kramer,  1969;  1979) which i s p r o p o r t i o n a l to a d e c r e a s i n g  water  flow  i s mainly  and i n v e r s e l y  Rose (1966),  At steady  soil-plant  system and the t o t a l  be e x p r e s s e d ,  state,  a  passive  through  the r o o t s , i n t o the xylem and  flow  i s equal  (soil,  through  r e s i s t a n c e from  a g a i n u s i n g an e l e c t r i c a l  resistances  process  p r o p o r t i o n a l to r e s i s t a n c e along  pathway.  component  f o r the flow  by a  Rutter  g r a d i e n t from the s o i l ,  the l e a v e s  can  flow  Law  (1971 and 1980) J a r v i s (1975),  Kramer and K o z l o w s k i ,  to  the p l a n t t i s s u e s t o  (1967).  phase  potential  t o Ohm's  f o r water  (1948) and has s i n c e been (1965),  The r a t e o f  s t a t e c o n d i t i o n s , can be expressed  analogy  model  r a t e s o f uptake  to the atmosphere by e v a p o r a t i o n .  steady  on  depends on t h e r e l a t i v e  root,  xylem  the same  a l l segments  the s o i l  o f the  to the l e a v e s  analogue, as t h e sum o f the and  leaf)  tissue  from  linked  in  series  ( H i n c k l e y et al., 1978).  Resistance  to water  leaves  i s apparently  1977a;  Tinklin  that  flow  through  relatively  and Weather ley,  1976).  xylem  resistance  may  small 1966).  r e s i s t a n c e i n c r e a s e s with  rates,  xylem  (Boyer,  flow  significant  to the  1971; H e r k e l r a t h et al.,  However, t h e r e  decreasing  be  the r o o t s  i s some  and, at very  (Oarvis,  evidence low flow  1975; R i c h t e r ,  - 3 -  Although resistance to flow in the bulk soil can become large as the soil dries, many authors indicate that the major source of resistance to liquid phase water flow is in the root, probably at the endodermis, or at the contact zone between the soil and the roots (Andrews and Newman, 1969; al.,  Herkelrath et al., 1977a, 1977b; Newman, 1969a, 1969b; Nnyamah et 1978).  Large plant resistance in the root has been suggested in several studies of plants growing in solutions 1971;  Tinklin  (Boyer, 1969; Stoker and Weatherley,  and Weatherley, 1966; Tomar and O'Toole, 1982).  plants growing  in soil  at normal rooting densities, soil  resistance  appears to be relatively small compared to root or soil-root resistance  (Dosskey,  Nnyamah et al., 1978; (Dosskey  contact  1978; Herkelrath et al., 1977a; Newman, 1969b; Reicosky  and Ritchie, 1976).  and Ballard, 1980; Faiz,  Oarvis, 1975;  For  Several  authors  1973; Faiz and Weatherley, 1978;  Nnyamah et al., 1978) indicate that soil-root contact is a  major source of total resistance.  This thesis arose from a keen interest in the relation between various forest site classifications and prescriptions for tree species selection with regard  to seedling  water requirements.  Given the problems in  studying water relations from a holistic ecological viewpoint (Elston and Monteith, 1975), i t seems inevitable to shift to what Gates (1980), in  the introductory  approach".  chapter  of his text,  called  "a reductionist  More specifically, differences in resistance to water uptake  arising from cultural practices, such as planting, and between contrast-  _ 4 _  ing  species  offered  presented a l o g i c a l  The of  objectives resistance  of  tractable  extension  this  subjects  research.  This  of work conducted by Dosskey  (1978).  t h e s i s are  for  to compare the  magnitude and  to water uptake i n ponderosa pine  to  Douglas-fir  and  to  (Pseudotsuga  i n v e s t i g a t e the  menziesii  e f f e c t of  (Mirb.)  planting  on  (Pinus  pattern  ponderosa  Franco  var.  resistance  in  also  Laws.)  menzxesii) Douglas-fir  seedlings.  A  study  obtained  of  with  an  in  ponderosa  pine  by Dosskey (1978) f o r D o u g l a s - f i r  resistance subject  resistance  to  water  uptake  of Chapter 2. overview  in  planted  A summary of the  of  their  in  comparison  to  results  i s presented i n Chapter 1 and Douglas-fir  seedlings  is  f i n d i n g s from both s t u d i e s  significance  is  given  at  the  end  the along  of  the  thesis.  Both of the ance was  studies  c a l c u l a t e d using  measurements uptake. the  employ many s i m i l a r methods.  of  soil  Seedlings  experiments  the  water  Ohm's Law potential,  Chapter 2 are  to water flow water  resistbased  on  potential  and  under  s i m i l a r greenhouse  conditions  and  were conducted  in a  similar controlled  environment  in  In the  i n t e r e s t of b r e v i t y , s i m i l a r d e t a i l s of  e x p e r i m e n t a l s t r a t e g y , methods and 1);  needle  example,  were grown  the same growth chamber.  Chapter  analogy  For  however,  departures  explicitly  stated.  materials made  in  are presented only once ( i n  the  experiments  described  in  - 5 -  CHAPTER  RESISTANCE  TO  1  WATER U P T A K E I N  COMPARED  TO  PONDEROSA  DOUGLAS-FIR  PINE  - 6 -  INTRODUCTION  Ponderosa pine (Pinus cially  ponderosa  Laws.) i s a widely d i s t r i b u t e d commer-  important tree species i n western North America occurring from  southern B r i t i s h Columbia east t o North Dakota, south to Trans-Pecos, Texas and west to southern C a l i f o r n i a and Mexico (Fowells, 1965). British  Columbia, i t i s confined  to the southern i n t e r i o r ,  In  south of  about C l i n t o n , at elevations mostly below 900 metres above sea l e v e l (Krajina et al., 1982).  Although i t s commercial importance i s r e l a t i v e -  l y small on a province wide b a s i s , i t assumes p a r t i c u l a r importance as one of a few species, or sometimes the only native species, which can survive and produce timber on d r i e r s i t e s i n the i n t e r i o r regions.  Much has been w r i t t e n on the morphological adaptations to drought and silvics  of the species  ( f o r example,  see B a r r e t t ,  1979; C u r t i s and  Lynch, 1957; Fowells, 1965; K r a j i n a , 1969; Mirov, 1967; Waring, 1970) and some aspects of water r e l a t i o n s have been studied by Cleary (1971), Lopushinsky (1969) and Lopushinsky and Klock (1974).  Because  of i t s  widely recognized a b i l i t y to survive and grow under dry conditions, i t i s i n t e r e s t i n g to speculate on what c h a r a c t e r i s t i c s might be responsible for maintaining a favourable water balance under adverse c o n d i t i o n s .  Lopushinsky (1969) and Lopushinsky and Klock (1974) i d e n t i f i e d r e l a t i v e l y rapid stomatal closure as an important adaptation. of  As an extension  these sort of s t u d i e s , i n v e s t i g a t i o n of resistance to l i q u i d  phase  water flow from the s o i l to the f o l i a g e and comparison with r e s u l t s from  - 7-  other species seem logical. reported  in part by  (Pseudotsuga  menziesii  Studies conducted by Dosskey (1978) and  Dosskey and  Ballard  (Mirb.) Franco var.  (1980) on Douglas-fir  menziesii)  provide a basis  for comparison.  METHODS AND  MATERIALS  Resistance to water uptake was calculated based on an Ohm's Law analogy to water flow proposed by van den Honert (1948):  [1]  °1  =  Af soil-root R soil-root  =  A¥root-leaf R root-leaf  =  where q is the rate of water flow and  Ay leaf-air R leaf-air  AY and R are, respectively, the  water potential difference and the resistance to flow along a particular pathway. In this study, only liquid phase water flow from the bulk soil to  the needle  was  measured.  Also, flow was  measured as the  transpiration rate at steady state in a controlled environment so it is equated to the uptake rate, U.  Therefore, equation 1 can be simplified  and solved for R:  ^s - * N [2]  R=  TJ  Ys is the bulk soil water potential and ^  is needle water potential.  Equation 2 provided the basis for calculating resistance in this study.  - 8-  Seedlings and Soil  Thirty grams of ponderosa pine seed, seedlot registration number Py 82E 5/B3/3002/0.50, were obtained from Forests Seed Centre at Duncan, B.C.  the British  Columbia  Ministry of  The seeds had been collected from  natural stands in the southern interior region of British Columbia (in the general vicinity of Penticton, B.C.) at an elevation of about 500 m above sea level.  Seeds were sown in coarse sand and gravel flats in a heated greenhouse on February 10, 1980 and young seedlings were transplanted to 11.4 cm (4-1/2") square, tapered pots on March 11 and 12, 1980.  The seedlings  were grown for over a year under natural illumination supplemented by a bank of fluorescent lights until the time of experimentation.  Periodi-  cally monitored daytime greenhouse temperatures generally ranged between 20° and 25°C and relative humidity was 80% plus or minus about 5%. Daytime irradiance was estimated to be about 20 to 25% of f u l l (but at  was of course greater on clear, sunny days). 8 hours  photoperiod. that  1981.  Photoperiod was kept  or greater, when necessary to correspond to the natural This was gradually increased prior to experimentation so  i t equalled the 15 hour photoperiod used  environment  sunlight  in the experimental  a week before the experimental runs were conducted  in May  Seedlings were watered daily, or less, as required to keep soils  near field capacity and were fertilized  regularly with f u l l strength  modified Hoagland's solution (see Dosskey, 1978, Appendix II).  - 9 -  Soil was obtained from the B horizon of a Sunshine soil series on the University of B.C. Endowment Lands, dried and sieved to remove particles greater than 2 mm. soil  Seedlings were potted with a pre-weighed mass of  packed to equal volumes.  Average total  360 cm^ and soil bulk density was 998 kg*m"3.  soil  volume was  about  A summary of some soil  properties is presented in Table 1.  Table 1 Soil Properties Texture Loamy Sand  % Sand* % S i l t * % Clay* 79  17  4  % 0Mb  pHC  Ksd  5.6  6.3  96 cm day~1  ^Based on 2 mm and smaller fraction, by hydrometer method Walkley-Black method 1:5 soil:water S a t u r a t e d hydraulic conductivity constant head permeameter method b  c  By December of 1980,  182 seedlings were growing and about 70% of them  had developed secondary needles. with  abnormally  high or  low  Over the next several months seedlings  leaf  area or soil  volume were culled.  Seedlings which had not shed their primary leaves prior to experimentation were also culled, leaving 128 seedlings.  Seventy-eight of these  were randomly selected for use in the experiments.  Soil  psychrometers  were then inserted into each of the pots at a point 1 cm from the root collar in the manner described by Dosskey (1978).  The lead wires were  coiled several times around inside the pot to avoid development of large thermal gradients in the f i r s t one-half metre.  - 10 -  Drainage holes in the pots were taped over and the seedlings sealed in plastic bags as described by Dosskey (1978). for  ten  minutes  every  day  to  allow  for  Seedlings were ventilated soil  atmosphere  exchange.  Seedlings were then watered from below and lightly sprinkled from above to bring them to field capacity and were placed in the growth chamber three days prior to experimentation to allow them to become acclimatized to the experimental environment.  Soils  were  again watered  experiment  and subsequent  to  field capacity at the beginning of  irrigation was withheld.  the  Measurements of  needle water potential, uptake and soil water potential were taken over a period of eleven days during May 1981.  Experimental Environment  The  experiment  was  conducted  in  a Percival  P-C-78 growth chamber  (#67-26) with lighting provided by a bank of fluorescent  and incande-  scent lights.  The spectral irradiance at mid-crown level is shown in  Appendix  Photon  I.  mid-crown  level  to about 25% of  flux  density  ranged from 490 sunlight at f u l l  to  within 540  the  visible  Mmol-m~2«s ^, -  spectrum  at  corresponding  solar noon during mid-summer.  The  maximum variation observed was about 20% and values were lowest near the front and near the corners of the growth chamber. avoided  during  the  experiment  and seedling  These locations were  positions  were randomly  rotated three times daily to eliminate bias associated with location.  - 11 -  In order to reduce boundary layer resistance and variation in temperature, humidity and carbon dioxide concentrations, the growth chamber was ventilated at an average velocity of 0.33 m«s"^ (measured with a hot wire  anemometer  chamber). m-s"^  at  mid-crown  level  in the  centre  of  the  growth  Average velocity at mid-crown level varied from 0.33 to 1.3  and was greatest near the ends; turbulence varied from 0.35 to  0.76 and was greater at the ends and at the front centre.  Temperature was controlled at 20 ±1°C during light periods and 15 ±1°C during dark periods.  Relatively  high soil  temperatures, similar to  those observed by Dosskey (1978), were measured in this study, probably due to the greenhouse effect of the plastic bags enclosing the seedlings (Dosskey, 1978).  Soil temperatures of 25°C were common and occasionally  reached 29°C in very dry soil (soil water potentials of -2.7 MPa and lower).  Relative humidity was controlled at 65 ±5% during light period and 85 ±5% for the dark period. monitored  with  a  Both temperature and relative humidity were  hygrothermograph  which  was  regularly  checked  and  calibrated by measurements with an Assmann psychrometer.  Soil Water Potential  Soil water potential was measured with Wescor PCT-55-05 thermocouple  - 12 -  psychrometers.  The psychrometric mode of measurment was used because of  its speed and r e l i a b i l i t y .  Psychrometers were calibrated in distilled  water and in sealed  vials  suspended over a 0.5 molal sodium chloride solution and were checked by immersion in distilled  water after the experiment was completed.  The  two point calibration is adequate since some preliminary measurements confirmed  the  linearity  potentials studied.  of  output  over  the  range  of  soil  water  After allowing ample time for equilibration, six  readings were taken for each psychrometer over the 0.5 molal solution. Psychrometers showing variation of more than ±0.5 UV, amounting to ±0.1 MPa or less in soil water potential at a potential of about -2.2  MPa,  were not used.  Measurements were taken three times daily beginning four hours after the start of the light period at about four hour intervals in conjunction with uptake measurements.  Measurements for determining the soil water  potential-needle water potential relationship were taken at 7-1/2 to 10 hours after the beginning of the light period.  Actual times of measure-  ments within this period were systematically varied.  Variability in soil water potential within a pot was investigated in a manner similar to that of Dosskey  (1978).  In addition, measurements  were compared with those from soil samples which were rapidly transferred  to Wescor C-51  and C-52  sample chambers.  Results were also  similar; up to ±0.3 MPa difference was found within a pot.  The greatest  - 13 -  variation seemed to be at lower soil water potentials (less than about -2.0 MPa) suggesting that variable contact between the psychrometer cup and thermal gradients within the pot are important factors contributing to variability.  Needle Water Potential  Needle water potential was measured with Wescor C-51 and C-52 thermocouple psychrometer sample chambers.  Sample chambers were calibrated  using 0, 0.05, 0.2, 0.5514 and 0.9 molal sodium chloride solutions and exhibited strong linearity in the range of water potential from -0.23 to -4.16 MPa.  Accuracy of measurements was within ±0.025 MPa at equilib-  rium.  Sample chamber holders were coated with paraffin adsorption to the holder surface (Boyer, 1967).  to minimize water  Large (0.95 x 0.48 cm)  sample chamber holders were used to accommodate the large, bulk needle samples used in the experiment.  One needle from  each  of three  randomly  selected  fasicles  (from a  randomly selected seedling) was removed, brushed with xylene, blotted dry, cut into sections and inserted in the sample chamber.  Since a  complete sample of even one needle was much too large to f i t in the sample chamber holder, middle, distal and proximal subsamples of each needle were used.  Several segments, about 0.5 cm in length, for each  - 14 -  subsample location and each needle were included in the bulk sample. The xylene treatment was used to speed the equilibration time in the sample chamber (Dosskey,  1978; Dosskey and Ballard, 1980; Neumann and  Thurtell, 1972).  Several different techniques were tried and practiced in order to minimize the time between xylene treatment and cutting and insertion in the sample chamber.  Using a pair of sharp surgical scissors for cutting the  needle sections speeded the sampling process considerably. Comparisons of razor cut and scissor cut needles showed no detectable difference in water potential.  After a considerable amount of practice the sampling  process could be completed within 60 seconds.  Sample chambers were checked using  procedures  recalibrated  given  by  with d i s t i l l e d  solution when necessary.  for contamination after each measurement Wescor water  (1979)  and  were  and a 0.5 molal  cleaned and  sodium  chloride  When no contamination was apparent, the sample  chambers were cleaned and recalibrated every other day.  Several different sample equilibration times, varying from 15 minutes to 5 hours were investigated and 20 minutes was found to be a suitable period.  Water potential at 20 minutes was found to be within less than  0.2 MPa of that at periods of 2 hours or more.  The relationship between soil and needle water potential was determined from  linear  regression  analysis.  A  grand  total  of 47  paired  measurements of soil and needle water potential were made; four or more individual seedlings were measured on each day and were fitted  to a  simple linear regression model:  [3]  y  where a  Q  = a  N  0  + b!  V  s  is the intercept and b-| the regression coefficient.  Uptake Seedlings were weighed two times daily at four hour intervals using a beam balance with an accuracy of ±0.01 g and uptake was calculated as the  average of the two four hour periods.  Because of the destructive  sampling of seedlings for needle water potential, uptake was determined from a separate group of 14 seedlings.  Measurements of soil  water  potential were taken with each weighing and the three readings were averaged  for determining the relationship between the average daily  uptake rate and soil water potential for each seedling.  Water uptake rate as a function of soil water potential was then determined  using  nonlinear  regression  analysis  to  provide a  range  of  continuous uptake values over the range of soil water potential where appreciable  uptake  expressed as:  occurs.  A general model for uptake  (U) can be  - 16 -  [4]  U = a +  where  F(f ) s  "a" represents  an asymptote  which  should a p p r o x i m a t e l y equal t h e  e v a p o r a t i o n r a t e from the bag s u r r o u n d i n g each pot (Dosskey,  The  v a l u e of " a " was e s t i m a t e d from  seven  pots  small.  containing  This  no  the average  seedlings  v a l u e was s u b t r a c t e d  and was from  1978).  of measurements  found  t o be  t h e measured  from  relatively  weight  loss to  a r r i v e at a t r u e uptake v a l u e ( e x c l u s i v e of e v a p o r a t i o n ) .  Uptake  f o r each o f t h e 14 s e e d l i n g s was then computed u s i n g the BMD-PAR  nonlinear Centre.  regression  program  (Ralston,  1983) a t t h e U.B.C.  Computing  Two r e g r e s s i o n models were used:  ( V > c  [5]  U = ab  [6]  U = a  2  MODEL I -b(*s-c)d 1-e  MODEL I I  Where a, b, c and d a r e c o e f f i c i e n t s whose v a l u e v a r i e s from s e e d l i n g t o seedling used  and "e" i s t h e base  by Dosskey  (Tsuga  (1978) i n h i s s t u d i e s  heterophylla  mertensiana (Dosskey,  (Raf.)  (Bong.) C a r r . ) 1978; Dosskey  W e i b u l l f u n c t i o n which engineering  of the n a t u r a l  and, more  Sarg.)  seedlings  and B a l l a r d ,  logarithms.  of Douglas-fir, and  1980).  Model  i n forest  hemlock  data  (Tsuga well  I I i s a modified  and l i f e  mensuration  i n c l u d i n g growth c u r v e s ( W e i b u l l , 1951; Yang e t a l . ,  I was  hemlock  h i s uptake  has been used i n r e l i a b i l i t y recently,  western  mountain  and f i t t e d  Model  1978).  testing i n  applications  - 17 -  Coefficients for each seedling were computed for each regression model and an average curve for each model was coefficients.  derived  by  averaging the  Uptake values for each seedling for both models and the  average for each model were generated for fixed (0.1 MPa) intervals of soil water potential.  Resistance  Resistance to water uptake was calculated from equation 2 for soil water potential values of -0.1 to -2.4 MPa at 0.1 values of needle water potential equations 3, 5 and 6.  MPa  intervals using the  and uptake derived  from regression  Because resistance values can be d i f f i c u l t to  interpret and compare for seedlings with different water pathway dimensions, these were estimated and a l l analyses were conducted on unit root area basis.  Water Pathway Dimensions  Needle surface area, root surface area and the average pathlength for water movement from the bulk soil to a root surface were estimated and calculated according to the procedures used by Dosskey (1978) (also see Dosskey and Ballard, 1980) and are summarized in Table 2. was that root density was determined in a dry state.  One exception  Dry densities were  then converted to fresh densities based on fresh:dry density ratios later determined from several seedlings from the population.  - 18 -  Table 2 Summary o f Average S e e d l i n g and Water Flow Pathway Dimensions for Ponderosa Pine S e e d l i n g s  Root Area Needle A r e a (cm2) (cm2)  3  Needle:Root Oven-Dry Root: Area R a t i o Shoot R a t i o Z d  Average  116  44.9  0.40  1.8  Coefficient of Variation  23%  23%  23%  24%  0 n e s i d e d area assuming a f l a t needle; t h i s underestimate of a c t u a l one s i d e d needle area ratio a  i s the c a l c u l a t e d average d i s t a n c e s o i l to a r o o t s u r f a c e  b  (cm)  0.19  13%  assumption r e s u l t s i n an and the n e e d l e : r o o t area  t h a t water must move through the  Comparison o f R e s i s t a n c e  Although  the experimental  were n e a r l y i d e n t i c a l to  rigorous  environment  and methods  used  in this  to those used by Dosskey (1978), s e v e r a l  comparison  of  the r e s u l t s  sampling and c a l c u l a t i o n t e c h n i q u e s  of  the s t u d i e s  study  obstacles  exist.  The  i n both s t u d i e s , n e c e s s i t a t e d by t h e  d e s t r u c t i v e sampling o f f o l i a g e f o r needle water p o t e n t i a l measurements, precludes on  loamy  (although  any p u r e l y sand  rigorous  textured  the average  s e e d l i n g s on s i l t  soil curve  loam a r e ) .  analysis.  Generated data  i s not p u b l i s h e d and generated  for Douglas-fir  on a u n i t r o o t  data  points  area  basis  for individual  - 19 -  The generated data violate assumptions of normality and homogeneity of variance which must be at least approximated  in order to use parametric  statistical tests such as analysis of variance or covariance.  These  shortcomings were not rectifiable by any commonly used data transformations.  In order to surmount these problems, the Kruskal-Wallis test, a nonparametric test suggested by Sokal and Rohlf (1981) in lieu of analysis of variance, was resistance  employed to test for significant differences between  for ponderosa pine and  Douglas-fir on  silt-loam. textured  soil.  RESULTS  Needle Water Potential  A plot of needle water potential against soil water potential shown in Figure 1 shows a linear, almost horizontal trend. of the data failed  to produce a statistically  Regression analysis  significant regression  coefficient indicating that needle water potential can be regarded  as  constant under the experimental conditions and over the range of soil water potential studied.  Mean needle water potential i s -2.2  the 95% confidence interval is from -2.3 to -2.1  MPa  and  MPa.  It should be noted that the linear relationship shown in Figure 1 i s not  - 20 -  Soil Water Potential Ok) (MPa)  -3.0  -2.0  i  i  ^  •  /  /  /  /  /  • a  m  /  /  /  /  /  /  /  /  /  /  /  /  /  /  /  /  • /  / J *  • •  •• • •  /  •  •  • •  o  -1.0  =  s  •  •  • • •  • •  Figure 1 NEEDLE WATER POTENTIAL vs SOIL WATER POTENTIAL FOR PONDEROSA PINE  (Error Bars Represent the 9 5 % Confidence interval)  - 21 -  d i r e c t l y comparable and S l a t y e r  to r e l a t i o n s h i p s such as those shown by Cowan (1965)  (1967) which show the d a i l y course of needle water  over s e v e r a l days of a d r y i n g purpose of such comparisons, regarded  as a s e r i e s  period  under  needle water  of needle water  potential  natural  conditions.  potential  in Figure  potential  For the 1 can  measurements taken  be  near  mid-day over a d r y i n g p e r i o d of s e v e r a l days.  The  data shown i n F i g u r e  needle  water  relatively This  potential  high  soil  h y p o t h e s i s was  1 aLso  and  soil  water not  suggest t h a t water  potentials  verifiable  by  the r e l a t i o n s h i p  potential (greater  may than  regression  not  be  about  between  linear -0.1  analysis.  at  MPa).  However,  f u r t h e r s t u d i e s with a g r e a t e r number of o b s e r v a t i o n s at high s o i l  water  potentials  small  might,  in fact,  show a n o n l i n e a r  range near the upper extreme  S i n c e needle water the  water  potential  potential  f u n c t i o n of s o i l  of s o i l  water  is virtually  relationship  potential.  a  potential.  constant i n t h i s  d i f f e r e n c e between the s o i l  water  over  experiment,  and needles i s a l i n e a r  This r e l a t i o n s h i p  i s shown i n F i g u r e  2.  Uptake  Uptake  r a t e s were p r e d i c t e d  equation  was  derived  using  Values f o r the average  f o r each s e e d l i n g both r e g r e s s i o n  coefficients  and an average  models  regression  ( e q u a t i o n s 5 and 6 ) .  f o r both models are shown i n T a b l e  -  22  -  Soil Water Potential (0s)  (MPa)  Figure 2 WATER POTENTIAL DIFFERENCE vs SOIL WATER POTENTIAL FOR PONDEROSA PINE  - 23  3.  No  statistics  residual  sums  determinable generated  such  of  for  uptake  Models I and  as  standard  squares an  values  deviations  are  average for  -  presented  equation. each  for  the  c o e f f i c i e n t s or  because  these  Figures  3  and  average  seedling  the  and  are  not  show  the  curves  for  4  II.  Table 3 Average C o e f f i c i e n t s f o r Uptake E q u a t i o n s  Model  a  b  I  3.99  II  8.52  Both models g i v e of  generally  e r r o r s ) and  on  iterative halvings  smaller standard  the  some of  root  magnitude  consistently  One  of  1.49  -2.11  n/a  1.65  2.07  the  during the  reliable  residuals  the  and,  for few  to  several  of  the  For  the  to  soil  be  convergence  of  increment  these  reasons,  c o e f f i c i e n t s i s not  some p h y s i c a l l y For  example,  the maximum asymptotic uptake r a t e and  water  potential  where  uptake  a  1983).  i t i s e a s i e r to a t t a c h  meaningful i n t e r p r e t a t i o n to some of the c o e f f i c i e n t s .  related  to  seedlings,  iterations.  deviations  advantage of Model I I i s t h a t  "a" r e p r e s e n t s  basis  standard  l i m i t s had  permit  i n d i c a t o r of p r e c i s i o n ( R a l s t o n ,  coefficient  the  (analogous to  However, i n many cases,  last  standard  On  f o r the c o e f f i c i e n t s , Model I appears t o  c o e f f i c i e n t s i n order process  -2.20  individual seedlings.  mean square  deviations  calculation occurred  d  good f i t f o r most  g i v e more p r e c i s e p r e d i c t i o n s . imposed  c  becomes  zero  the  "c" i s (i.e.  UPTAKE IN PONDEROSA PINE PER UNIT ROOT AREA AS A FUNCTION OF SOIL WATER POTENTIAL - REGRESSION MODEL I  -2.0  -1.0  Soil Water Potential  (MPa)  Figure 4 UPTAKE IN PONDEROSA PINE PER UNIT ROOT AREA AS A FUNCTION OF SOIL WATER POTENTIAL - REGRESSION MODEL 11  0  - 26  where s o i l features  water p o t e n t i a l  are  regression  also  helpful  require  coefficients  "b"  respectively,  and  the  Average curves  an  "d"  average r e s i d u a l s  are  and  both  soil  water  tently  The  and  so  bias  6.  between  -2.8  MPa;  -1.2  and  plotted  in  Figures  3 and  variability  which  variability  seedlings,  environmental  chamber  and  seedlings.  variability No  the  experiment  too  low  that  to  account  physiological  significant  in  the  ectomycorrhizal and  the  each  for  nonlinear  coefficient.  interpretable  as  These  but  shown by  effect,  1978).  a  the  The  plot  of  the  residuals  for  t r e n d s shown f o r the  plotted  overpredicts  between  Model -1.8  II  MPa  uptake tends  but  to  consis-  underpredicts  MPa.  show c o n s i d e r a b l e among  programs  Examination of  c o n f i r m s the  and  potential).  curve (Yang et al.,  consistently  -1.5  uptake  -2.2  uptake v a l u e s  of  I  for  easily  models e x h i b i t  Model  potentials  overpredict  between -2.0  not  i n F i g u r e s 5 and  residuals.  needle water  estimate  shape of the  curves of i n d i v i d u a l s e e d l i n g s average  to  because most computer  initial  scale  for  i s equal  -  degree  of  for  individual  attributed  to  variability mycorrhizal  variability.  in  i n uptake.  the  growth  infection  variation  of  appears to  be  Therefore,  the  among end  between i n d i v i d u a l s e e d l i n g s  source of v a r i a b i l i t y  seedlings  physiological  o b s e r v a b l e at  environmental  observed  differences  be  development was  degree of  f o r the  can  4  i t appears i s the  most  - 27 -  - 4  -  3  'E |  CO  •  - 2  -  1  -  0  - - 1  - - 2  -3.0  -2.0  -1.0  Soil Water Potential (MPa)  Figure 5 AVERAGE RESIDUALS FOR UPTAKE EQUATION MODEL I  ( +=underprediction, — = overprediction )  CO  E _W CO  •o CO  cu DC d)  - - 3  O) CC  - - 4  >  0  V.  Q  <  - 28 -  —|  -3.0  i  i  i  i  |  i  i  i  -2.0  i  |  i  r  -1.0  Soil Water Potential (MPa)  Figure  6  AVERAGE RESIDUALS FOR UPTAKE EQUATION MODEL 11  ( + = underprediction, — = overprediction)  - 29 -  Resistance  t o Water Uptake  Average s e e d l i n g  r e s i s t a n c e , on a u n i t r o o t  e q u a t i o n 2 i s shown f o r both 8.  Resistance  increase then  based  as s o i l  decreases  water  until This  for  shown  artifact  as  Model  potential  zero  Resistance  i n Figure  tude and becomes i n f i n i t e l y shown at h i g h e r  of b i a s i n t h e uptake  ever,  Model  5,  result  of b i a s  estimated resistance.  in  rise  and  7)  -0.1  therefore  shows  a  t o -1.2  water  7 and slight  MPa  and  p o t e n t i a l of  appears  indicated to  be  an  process.  8) shows a much  rapidly.  soil  different  water  an order  The i n i t i a l  poten-  of magni-  decrease i n  water p o t e n t i a l s appears t o be a r e s u l t (see F i g u r e water  i n resistance  range,  At a s o i l  by n e a r l y  l a r g e a t -2.2 MPa.  at s o i l  i n predicted that  from  I I (Figure  relatively  equation  overestimated  the steep  (Figure  at a s o i l  o f -2.0 MPa, r e s i s t a n c e has i n c r e a s e d  slightly  models i n F i g u r e s  from  d e c r e a s e s between s o i l water p o t e n t i a l s of -0.1 and  -1.1 MPa and then i n c r e a s e s  resistance  basis, calculated  behaviour i s i n agreement with the b i a s  based on uptake  pattern.  I  decreases  o f the r e s i s t a n c e c a l c u l a t i o n  Resistance  tial  uptake  regression  i t approaches  about -2.4 MPa. uptake,  on  uptake  area  6); similarly  potentials  between  would  below -1.9 MPa.  -1.1 and -1.8 MPa  uptake; i n f a c t , which  resistance i s  uptake  result  in  How-  i s not a  i s generally  over-  underestimates  of  - 30 -  h 3.0 a> o  c ^  Soil Water Potential  (MPa)  Figure 7 RESISTANCE TO UPTAKE PER UNIT ROOT AREA FOR PONDEROSA PINE ( Uptake Model I) (Error Bars are Approximate)  -  31  -2.0  -  -1.0 Soil Water Potential  0.0 (MPa)  Figure 8 RESISTANCE TO UPTAKE PER UNIT ROOT AREA FOR PONDEROSA PINE ( Uptake Model 11) (Error Bars are Approximate)  - 32 -  Comparison of Ponderosa Pine to Douglas-fir  The  results of this study compared to Dosskey's study of Douglas-fir  (Dosskey, 1978), conducted under similar environmental conditions and using  similar  Douglas-fir  techniques,  regulate  indicate  both  ponderosa  needle water potential equally  needle water potential against show a nearly  that  well;  pine  and  plots of  soil water potential for both species  horizontal line.  Uptake rates in ponderosa pine are  greater than in Douglas-fir at high soil water potentials, but rates decrease more rapidly in ponderosa pine as the soil dries.  A comparison of resistance to water uptake for ponderosa pine (based on uptake Model II) and Douglas-fir on s i l t loam and loamy sand soil (from Dosskey, 1978) i s shown in Figure 9. sand i s an approximation derived  The curve for Douglas-fir on loamy  from dividing resistance values per  seedling by the average seedling root area.  Because very few observa-  tions could be made at soil water potentials greater than -0.5 MPa in both studies, values  in that range are not included  in the species  comparison.  Resistance  for Douglas-fir  on s i l t  loam was compared  to that for  ponderosa pine at soil water potential values of -0.6, -1.0 and -2.0 MPa using the Kruskal-Wallis test.  A summary of the statistical analysis is  presented in Table 4.  The results show a very highly significant difference between the two  - 33 -  -2.0  -10  Soil Water Potential  0.0  (MPa)  Figure 9 RESISTANCE TO UPTAKE PER UNIT ROOT AREA FOR PONDEROSA PINE ( Uptake Model 11) DOUGLAS-FIR ON SILT LOAM AND DOUGLAS-FIR ON LOAMY SAND (Curves for Douglas-Fir from Dosskey, 1978)  - 34 -  species  at s o i l  indicated  at  significant  water -2.0  because  potentials MPa  is  of -0.6 and -1.0 MPa.  not  considered  to  be  The d i f f e r e n c e physiologically  e s t i m a t e s o f r e s i s t a n c e based on uptake Model  I for  ponderosa p i n e a r e a r t i f i c i a l l y low.  Table 4 K r u s k a l - W a l l i s Test f o r D i f f e r e n c e s i n Resistance to.Water Uptake f o r D o u g l a s - f i r (F) and Ponderosa Pine (Py)  S o i l Water P o t e n t i a l (MPa)  Chi-Square F vs Py Model I  Chi-Square F vs Py Model I I  -0.6  8.06*  12.00**  -1.0  8.76*  17.02**  -2.0  15.75**  2.29 n.s.  n.s. Not s i g n i f i c a n t l y d i f f e r e n t * S i g n i f i c a n t l y d i f f e r e n t at = 0.005 * * S i g n i f i c a n t i y d i f f e r e n t at = 0.001  Comparison results  of r e s i s t a n c e  of  the  to water uptake, as shown i n F i g u r e 9, and the  Kruskal-Wallis  test  indicate  ponderosa pine on loamy sand i s s i g n i f i c a n t l y on  silt  loam  at h i g h e r s o i l  water  potentials.  that  resistance  for  lower than f o r D o u g l a s - f i r As s o i l  approaches -2.0 MPa,' r e s i s t a n c e i n c r e a s e s more r a p i d l y  water  potential  i n ponderosa p i n e  and becomes a p p r o x i m a t e l y equal f o r both s p e c i e s .  DISCUSSION  The  observed  constancy  of needle water  potential  and r e l a t i v e l y  rapid  - 35 -  decrease in uptake as soil water potential decreases generally agree with the findings of Lopushinsky (1969) and Lopushinsky and Klock (1974) that ponderosa  pine is relatively sensitive to decreasing soil  water  potential compared to Douglas-fir and that leaf water potential should decrease slowly  as soil water potential decreases.  The shape of the  uptake curves from this study compared to the curve for Douglas-fir on loamy sand soil from Dosskey (1978) bear the same general relationship as the transpiration curves for the same species shown by Lopushinsky and Klock (1974); that i s , transpiration (or, in this study, uptake) f a l l s rapidly in ponderosa pine compared to Douglas-fir.  Transpiration  in ponderosa pine was shown to be about 3% of the maximum rate at a soil water  potential of -2.0  MPa  compared  to about  20%  of maximum for  Douglas-fir (Lopushinsky and Klock, 1974).  This agrees closely with uptake predicted with Model II at a soil water potential of -2.0 MPa in this study, which is about 3% of the maximum uptake rate.  Model I predicts an uptake rate of about 30% of the maxi-  mum rate at the same soil water potential. However, both models predict greater uptake rates than would be expected from Lopushinsky and Klock (1974) at higher soil water potentials.  The  estimated  based  maximum  transpiration  rate  (1.7  on measurement of the stomatal resistance  x  10~^mg*s~ -cnr^)  of a well  watered  seedling, agrees closely with the maximum rate predicted by the average uptake  curve  for  Model  II  (about  2.0  x  10~3mg's"^•cm~2).  The  maximum rate predicted by the average curve for Model I (about 5.0 x  - 36  10"3mg*s~ 'cm-^) i s n e a r l y  Based  on  these comparisons,  observed  uptake  for  poor  model.  relatively low  three  soil  water  as  times  well  individual In  potentials  make  questionable.  a r e s u l t of u n d e r e s t i m a t e s of  same  This  Dosskey  regression  the  low  probably soil  There  the  the  However,  source  are  probably  several  i n the  at  resistance  to  rate  this  at (the  problem  relative  least  in  intercept in  using  errors  that  uptake r a t e approach  zero  in predicting resistance  by  why  uptake  Lopushinsky  were made at s o i l  and  falls  less rapidly  Klock  (1974).  at  upper  extreme are  in  experimental  temperatures  probably  only  conditions,  in this  rough  such  study, may  as  account  slightly  (for  properties exploitation  example  such by  as  from  -0.1  to  conductivity,  roots  d i f f e r e n c e s between the  seedlings  0.2  1967).  i n the two  few than  r e s i s t a n c e at Differences  the  high  relatively  some d i s c r e p a n c i e s .  soil  volume  (Rutter,  Very  approximations.  for  r a t e of decrease i n t r a n s p i r a t i o n as  than  water p o t e n t i a l s g r e a t e r  i n t h i s study; t h e r e f o r e , e s t i m a t e s of uptake and  the  the  reasons  study  (only f i v e ) o b s e r v a t i o n s MPa  difficulty  predictions  water p o t e n t i a l s .  transpiration  -0.3  of  and  a  be,  to  to  be  of  may  large  predicted  seems  high  evaporation  the  of  I  estimates  (1978) encountered  model.  major  Model  overprediction  occur as both the water p o t e n t i a l g r a d i e n t is  comparison  resulting  part,  the  as  p a r t i c u l a r , i t s overly  uptake  i n equation 4 ) .  greater.  seedlings,  water  "a"  -  Also,  water p o t e n t i a l decreases  MPa) of  soil  will  soil  and  depend  on  its  degree  Finally, s t u d i e s may  soil of  physiological be  significant.  - 37 -  Predictions soil  o f uptake  water  realistic fitting  potentials than  Model  based  on Model  below  about  -0.5 MPa,  from  Model  predictions  I I to the data  o b s e r v a t i o n s a t t h e upper  extreme  and the uptake  the water  potential  rate  and are judged  I.  i s probably  a v o i d s the problem o f o b t a i n i n g gradient  I I seem r e a s o n a b l e , at l e a s t f o r  of s o i l  The  major  limitation  the d i f f i c u l t y  water  t o be more  of  potential.  difference  getting  T h i s model  p r e c i s e e s t i m a t e s of the water  by f o r c i n g  to  potential  the uptake curve t o z e r o when  approaches z e r o .  This strategy  probably  produces some d i s t o r t i o n s a t other p a r t s of the curve which are apparent in  the p l o t  this it  model  o f average  residuals  seems j u s t i f i a b l e  shown  at l e a s t  seem to produce more r e a l i s t i c  i n Figure  6.  However, use o f  from a pragmatic v i e w p o i n t s i n c e  p r e d i c t o n s of average uptake  rates.  S i n c e the d i f f e r e n c e i n r e s i s t a n c e shown between the c u r v e s i n F i g u r e s 7 and 8 a r e a f u n c t i o n resistance increase then  t o water  as s o i l  of t h e i r uptake  water  respective  based  on Model  potential  d e c r e a s e s to n e a r l y  uptake models i t f o l l o w s I (Figure  7) shows a  decreases to intermediate  z e r o a t about  ^  s  = -2.2 MPa.  With  regard t o  subsequent Resistance decrease  drop  i n resistance  based  on  i n resistance  Model  II  as s o i l  -1.1 MPa i s a l s o an a r t i f a c t  below  artifact ^  s  =  of the c a l c u l a t i o n s , the  -1.8 MPa  i s reasonable water  Even i f  decreases from -1.2 t o -1.8 MPa  s  can be e x p l a i n e d as a r e l a t i v e l y minor  slight  v a l u e s and  the e x i s t i n g knowledge of the s p e c i e s t h i s does not make sense. the s l i g h t decrease i n r e s i s t a n c e a s V  that  potential  seems u n r e a s o n a b l e .  in this  respect.  The  decreases down to about  of c a l c u l a t i o n s ( r e s u l t i n g from o v e r p r e d i c -  t i o n s o f uptake) but the subsequent  r i s e i s not.  - 38  Statistically covariance)  of  Douglas-fir the  rigorous  underlying  However,  the  differences together  p a r a m e t r i c t e s t s (such  differences  cannot be  in  properly  assumptions differences  in  resistance  as a n a l y s i s of v a r i a n c e  between  ponderosa  pine  or and  a p p l i e d to t h i s study because too many of  of  in  resistance  suggest  -  such  the  and  tests  are  patterns  the  of  results  a meaningful d i f f e r e n c e  too  severely  uptake,  of  the  the  violated.  magnitude  Kruskal-Wallis  in resistance  of  test  between the  two  species.  The  experimental c o n d i t i o n s  were p r a c t i c a l l y  identical  dimensions f o r the  two  for this  study and  the  (see Dosskey, 1978).  species  on  study  of  S o i l s and  loamy sand s o i l  are  Douglas-fir  water pathway  compared  i n Table  5.  Table this which  5  indicates  study has might  5%  that  soil  l e s s sand, 5%  result  in  Therefore,  comparable. root  The  surface  differences  in  soil  differences  between  Since  i s not  the  two  species  between  the  water  the  two  uptake  are  3.2%  The  times  retention  be  much  and  a  major  higher  i t s lower bulk  to  say  the  it  for  that  ponderosa  soils  that  factor  are  soil  to a  pine.  If  for  the  responsible  would  greater  i t i s suggested not  matter  o f f s e t by  largely  species,  in  be  greater  were  soil  more o r g a n i c  which water must move i n the  would  case,  similar.  water  reasonable  resistance  to  this  seems  three  resistance  and  However, t h i s may  it  about  very  increased  average d i s t a n c e  is  are  more s i l t  slightly  unsaturated c o n d u c t i v i t y . density.  conditions  be  expected  that  for  ponderosa  pine.  differences contributing  in to  soils the  - 39 -  observed d i f f e r e n c e s of the d i f f e r e n c e s on s i l t  i n r e s i s t a n c e to water uptake  (particularly  shown when ponderosa pine i s compared t o  i n view  Douglas-fir  loam).  Table 5 S o i l C h a r a c t e r i s t i c s and Water Pathway Dimensions for Ponderosa Pine and D o u g l a s - f i r d  Parameter  Ponderosa P i n e  Douglas-fir  loamy sand 79 17 4  loamy sand 84 12 4  S o i l Texture % sand % silt % clay  0  0  0  S o i l Bulk D e n s i t y (kg.m-3) % Organic M a t t e r Average  Z (cm)  998  0  e  a  1,120  5.6  2.4  0.19  0.064  F r o m Dosskey (1978) °Based on 2 mm and s m a l l e r f r a c t i o n , by hydrometer method W a l k l e y - B l a c k method ^1:5 s o i i : w a t e r C a l c u l a t e d average d i s t a n c e t h a t water must move from the root surface a  c  A  complete  and  and  thorough  non-sampling  errors  measurement o f s o i l tion  of  sampling the  a  "worst  errors  error  analysis  associated  incorporating  with  prediction  water p o t e n t i a l i s not p o s s i b l e . case"  upper  limit  of about ± 5 0 % i s s t i l l  d i f f e r e n c e s found.  for  combined  soil  a l l the  of  uptake  A rough sampling  to  a  sampling and  the  approximaand  non-  not l a r g e enough to account f o r  - 40 -  The observed differences in uptake and resistance may  reflect different  adaptive strategies of drought avoidance and tolerance.  Low  resistance  to water uptake at high soil water potentials facilitates the high rates of uptake observed in this and other studies of ponderosa pine and other pine species.  The relatively rapid decrease in uptake as the soil dries  is probably at least partly a result of the  greater  stomata to decreasing  In contrast, uptake in  soil  water potential.  sensitivity  of  Douglas-fir f a l l s much more gradually as the soil dries.  Rapid  stomatal  closure  is  probably  an  important  drought  avoidance  mechanism for ponderosa pine, which often grows on dry sites and in relatively dry climatic regions where the major source of growing season soil  water  is usually  from  snowmelt and  spring  rainfall.  Coastal  Douglas-fir is mainly a successional species; an avoidance strategy such as rapid stomatal closure in response to decreasing soil water potential would probably not be useful because any soil water conserved might be depleted by competing vegetation and maintenance of transpiration may necessary to avoid  excessive  leaf temperatures  (Bunce et a l . ,  be  1977;  Duhme, 1974; Passioura, 1976).  The  rapid increase  potentials of -1.5 and  in resistance to water uptake between soil water and -2.0  MPa  is a result of increasing plant, soil  soil-root contact resistance.  Comparison with Douglas-fir studied  by Dosskey (1978) under similar conditions and growing in similar soil suggests that responsible.  some physiological and  morphological factors might  It might be that differences in root morphology and  be the  - 41 -  degree  of m y c o r r h i z a l  development  (1978) noted t h a t  the r o o t s  of  root  fine  lateral  mycorrhizal were  present  hairs  i n ponderosa  responsible.  Dosskey  of D o u g l a s - f i r had developed a l a r g e and  that  there  was  development on some r o o t s ; n e i t h e r  responsible in  are i n d i r e c t l y  pine.  f o r a more gradual  These  increase  root  a  slight  number  degree  of  of these c h a r a c t e r i s t i c s c h a r a c t e r i s t i c s could  in soil-root  contact  be  resistance  Douglas-fir.  SUMMARY AND CONCLUSIONS  Resistance based  on  to water  uptake  independent  estimates  determined  as  controlled  environment.  observations found  a  was  function  used  to be v i r t u a l l y  i n ponderosa  of  of  needle  decreasing  Linear  to  needle  over  seedlings  water soil  regression  estimate  constant  pine  potential  calculated and  water  potential  analysis  based  water  the range  was  uptake in a on  47  p o t e n t i a l , which  was  of observed  soil  water  potential.  Uptake r a t e s were e s t i m a t e d two  different  satisfactory  models. overall  using nonlinear  A modified predictions.  based on a t o t a l o f 92 o b s e r v a t i o n s where uptake plotted for  occurs.  for fixed  Predicted  values  of s o i l  the s p e c i e s was determined.  Weibull  r e g r e s s i o n a n a l y s i s based on function  provided  the most  f o r 14  seedlings  Curves were f i t t e d i n the range of s o i l  uptake water  values  were  water p o t e n t i a l  then  generated  p o t e n t i a l and an average  and  curve  - 42 -  Average seedling resistance to water uptake was calculated and plotted based on predicted needle water potential and generated average uptake values for regular,  fixed  intervals  of soil  water  potential.  The  results were then compared to results from a similar study of Douglasf i r conducted by Dosskey (1978).  Compared to Douglas-fir, resistance to water uptake in ponderosa pine seedlings differs, both in magnitude soil water potential decreases. water  potential  equally  well,  and in i t s pattern of change as  Both species seem to regulate needle maintaining  nearly  constant  potential over a relatively wide range of soil water potentials.  water Uptake  rates are comparatively large for ponderosa pine at higher soil water potentials but decrease much more rapidly as the soil dries. Resistance to uptake is lower for ponderosa pine until soil water potential reaches about -2.0 MPa.  Resistance starts increasing at a soil water potential  of about -1.0 MPa and rises rapidly between -1.5 and -2.0 MPa.  The relatively  low resistance to water uptake at higher soil  water  potentials agrees with the observed high transpiration rates at high soil water potentials reported in the literature for ponderosa pine and other pine species.  The rapid  increase in resistance as soil water potential drops from  about -1.5 to -2.0 MPa probably reflects increasing soil and soil-root contact resistance, as well as plant resistance.  - 43 -  CHAPTER 2  RESISTANCE TO WATER UPTAKE IN PLANTED DOUGLAS-FIR SEEDLINGS  ¥f  -  -  INTRODUCTION  Coastal Douglas-fir has  long  been  British  and c o n t i n u e s  Columbia  Douglas-fir physiology  (Pseudotsuga  has  and  the  probably  Pacific  requires  received  as  on s i t e  t o flow  along  s o r t of i n f o r m a t i o n  of  the  menziesii) species i n  United  Many past  States.  study  of i t s  s t u d i e s of i t s  c h a r a c t e r i s t i c s or p l a n t  seedling  water  relations  and p l a n t water s t a t u s , but o f  segments  is difficult  producing  ( o r more)  and s o i l  of s o i l  various  much  of  i n B.C.  understanding  some knowledge, not o n l y  resistance This  however,  timber  Northwest  and ecology as any s p e c i e s  status;  (Mirb.) Franco var  t o be a major  water r e l a t i o n s have c e n t r e d water  menziesii  o f the water  to gather  flow  pathway.  because many p l a n t and  environmental v a r i a b l e s can confound i n t e r p r e t a t i o n s .  Dosskey  (1978) r e v e a l e d  some s i g n i f i c a n t  c h a r a c t e r i s t i c s r e l a t e d to r e s i s t a n c e soil-plant  system.  was  soil-root  that  resistance also  been  (1977a, as  contact  by  resistance  pathway.  a  number  may  phase water arising be  a  from major  S i g n i f i c a n t contact o f authors  such  source  of r e s i s t a n c e  significance  of c o n t a c t  could  important  be an  Douglas-fir  resistance factor  seedlings.  to water  flow.  i t i s logical  influencing  and s o i l  flow  Given  source  of  r e s i s t a n c e has  by F a i z  e t al. (1973)  the p o t e n t i a l  to speculate  the s u r v i v a l  i n the  h i s studies  as H e r k e l r a t h  1977b), Nnyamah et al. (1978) and was i d e n t i f i e d  a major  planted  to l i q u i d  An i n t e r e s t i n g i n f e r e n c e  i n t h e water inferred  insights into species  that i t  and growth o f  - 45 -  The objective of this experiment i s to investigate the relative magnitude of resistance to water uptake in planted Douglas-fir seedlings and to see i f soil-root contact effects could be contributing to increased total resistance.  METHODS AND MATERIALS  Many of the methods employed in this experiment have been described and discussed in the previous chapter reporting the results of a study on ponderosa  pine and will  not be repeated here.  The greenhouse and  experimental environments, general properties of soils, instruments used for measuring uptake and soil and needle water potential and watering and fertilization procedures are the same in both experiments.  Six-hundred,  one year  old bareroot  Douglas-fir  seedlings,  92G/B2/2890/0.145, were obtained from the B.C. Ministry  seedlot  of Forests'  Green Timbers Nursery in Surrey, B.C. on February 8, 1980 and stored in a sealed ambient  vinyl outdoor  lined  paper  seedling bag in a sheltered location at  temperatures.  On February  11, the seedlings were  planted in 11.4 cm square, tapered pots using preweighed volumes of soil packed  to equal volumes.  Seedling roots were thoroughly washed to  remove the nursery soil and roots in excess of about 10 cm in length were pruned prior to potting.  Seedlings were grown in the greenhouse  for 15 months prior to experimentation.  - 46 -  Average soil bulk density was 995 kg-nr^ and average soil volume about 370 cm^.  Other general soil properties are described in Table 1 in the  previous chapter.  Effect of Needle Removal! on Uptake  Seedlings were of sufficient needle area to suppose that removing a few needles would  not  appreciably  affect  water  balance.  For  example,  removal of 10 needles would constitute a needle area reduction of only about one or two percent. A nested factorial experiment to investigate the effects of removing 3 or 10 needles on water uptake rates was run in the growth chamber during January of 1981. selected and assigned to treatments.  Nine seedlings were randomly  Treatment factors were defined as  three levels for the number of needles removed (0, 3 and 10) combined with two qualitative levels of soil water potential (-0.3 to -0.6 MPa and -1.4 to -2.2 MPa).  Soil water potential was monitored continually  throughout the experiment using Wescor PCT-55-05 soil psychrometers. The results of analysis of variance conducted showed no effect of removal of 3 or 10 needles on uptake rates (at  a  significant  <0.25).  Measurement of Needle Water Potential  Since up to 10 needles could be removed without producing a detectable change in water uptake, this opened the possibility of making repeated  - 47 -  measurements of needle water potential on an individual seedling.  Fur-  thermore, the same seedling could be used for measuring uptake so that the awkward statistical procedures for calculating resistance used in the previously described experiment on ponderosa pine could be avoided.  Exploratory  experiments  were conducted  during February  of 1981 to  determine a suitable sample holder size, sampling procedure and equilibration time for a single needle sample in the Wescor C-51 and C-52 thermocouple psychrometer sample chambers.  Using the .64 x .08 cm (1/4  x 1/32 inch) sample holder, a 30 minute equilibration time resulted in water potential measurements that were usually within ±0.1 MPa of the value  at two hours.  Occasionally  the difference  was as much as  ±0.2 MPa.  Paired measurements of needle water potential from the same seedling at the same time were then taken for single needle samples and bulk samples of 10 needles. observations. 0.078 MPa  The results were analyzed using a t-test for paired Water potential measured for single  lower  than  for bulk samples  needles averaged  but the difference  was not  statistically significant (at <*<0.20).  Investigations into variability in water potential among needles on the same seedling sampled at the same time revealed that needle water potential  estimates could vary up to about 20% (about ±0.2 to ±0.3 MPa).  However  this  variability  was  reduced  to ±0.15 MPa  by restricting  sampling to needles of the same age and comparable crown position.  - 48 -  In the e n s u i n g e x p e r i m e n t , n e e d l e s were s e l e c t e d from the m i d - p o i n t of a l a t e r a l t w i g near mid-crown l e v e l .  The needle was  treated with xylene,  b l o t t e d d r y , c u t i n t o t h i r d s u s i n g a r a z o r b l a d e , i n s e r t e d i n the sample chamber and a l l o w e d t o e q u i l i b r a t e f o r 30 minutes b e f o r e water measurements were t a k e n .  potential  A needle c o u l d be sampled, t r e a t e d and s e a l e d  i n the sample chamber w i t h i n 30 seconds.  E x p e r i m e n t a l C o n d i t i o n s and P r o c e d u r e s  S e e d l i n g s were weighed tial  and c o n c u r r e n t measurements of s o i l water poten-  were t a k e n t h r e e t i m e s d a i l y  at four  hour  intervals  f o u r hours a f t e r t h e s t a r t of the l i g h t p e r i o d . measurements were made between 7-1/2 the  light  altered. their  period  and  the  time  S e e d l i n g s were s e l e c t e d  location  i n t h e growth  beginning at  Needle water p o t e n t i a l  and 10 hours a f t e r the b e g i n n i n g o f of  measurement  was  systematicaliy  i n random o r d e r f o r measurements and  chamber was  changed  randomly  three times  d a i l y a f t e r each w e i g h i n g .  Uptake was c a l c u l a t e d as the average f o r two  f o u r - h o u r p e r i o d s and  water  daily  readings.  soil  R e s i s t a n c e was  potential  was  calculated  averaged  from  water  f o r the t h r e e potential  and  uptake v a l u e s u s i n g e q u a t i o n 2 (Chapter 1 ) .  Variability  in soil  water p o t e n t i a l was  found t o be comparable  to that  i n the ponderosa p i n e e x p e r i m e n t s (Chapter 1 ) .  Soil  temperatures  were  less  than  i n the  ponderosa  pine  experiment,  - 49 -  probably  because  surface  was  farther  Temperatures dry  the D o u g l a s - f i r below  seedlings  the bank  of  were  lights  taller  the  i n the growth  were g e n e r a l l y 25 t o 26°C but o c c a s i o n a l l y  s o i l s with s o i l  and  soil  chamber.  reached 27°C i n  water p o t e n t i a l s of l e s s than -2.0 MPa.  Treatments  Fifteen  s e e d l i n g s were randomly  treatments: seedlings  control,  were  original  soil  kept  a  to  misting.  planted  carefully  repacked  minimum  selected and  lifted  and a l l o c a t e d  planted from  and  their  vibrated.  pots,  to i t s p r e v i o u s volume.  (about  Immediately  one  after  minute) planting  and  Root  roots  soils  to one of t h r e e of the  replanted  and  the  exposure  time  was  moist  by  were  were  Ten  kept  watered  to  field  capacity.  Five  of the p l a n t e d  vibration high  bulk  984  while  densities  planted,  were  randomly  selected  f o r 30 minutes on a v o r t e x mixer.  ( a t about  minimal  seedlings  and  kg'm"^,  field  still  capacity)  allowing  sampled planted  at and  respectively.  apparently  vibration the  end  of  Keeping helped  t o move the  and s u b j e c t e d soil  treatments  Allowing  f o r sampling  compaction  particles.  experiment  vibrated  water c o n t e n t  to keep  soil  were  to  for  995,  error,  Mean  control, 1020  they  can  and be  regarded as e q u a l .  Soil  psychrometers  were  then  inserted  in  each  pot  in  the  manner  - 50 -  described i n the previous chapter and seedlings were watered and placed in the growth chamber for a three day equilibration experiment began.  period before the  Measurements were made over a period of s i x days i n  A p r i l 1981.  Experimental Design and Data Analysis  The  experiment  effects model.  was set up as a nested  analysis of variance, mixed  Five seedlings were allocated to each treatment  r e p l i c a t e measurements were taken for each seedling.  and s i x  Treatment effects  were considered fixed and both seedlings and replicates were treated as random e f f e c t s .  Because individual seedlings depleted s o i l water at d i f f e r e n t rates and depletion was especially slow for planted seedlings, covariance analysis was used to analyze the data. analyzed  Before the f i n a l analysis, the data were  for normality and homogeneity of variance by plotting  B a r t l e t t ' s test  (Sokal and Rohlf, 1981).  In order to meet the assump-  tions of analysis of variance and covariance data were transformed base-10 logarithms.  and by  using  Data were then analyzed using the UBC ANOVAR (Greig  and O s t e r l i n , 1978) program at the University of B.C. Computing Centre.  - 51 -  Seedling and Water Pathway Dimensions  Seedling  and  water  pathway dimensions were determined using methods  described by Dosskey (1978) and are summarized in Table 6. Table 6 Seedling and Water Pathway Dimensions for Douglas-fir Seedlings (Measured at the end of the Experiment) 3  Root Area Needle Area Needle:Root (cm2) (cm2) Area Ratio 0  Treatment  Oven Dry Root: Shoot Ratio  (cm)  Control  246 (21%)  111 (17%)  .46 (15%)  .60 (28%)  0.14 (19%)  Planted  290 (23%)  101 (13%)  .37 (32%)  .62 (34%)  0.12 (20%)  Planted and Vibrated  301 (28%)  130 (30%)  .44 (18%)  .57 (70%)  0.14 (28%)  Numbers in parentheses are coefficients of variation °0n e-sided needle area Z is the calculated average distance that water must move through the soil to a root surface d  C  RESULTS  Needle Water Potential  Plots of needle water potential against soil water potential for each of the three treatments are shown in Figures 10, 11 and 12.  In these  figures, different symbols represent five individual seedlings and the  Figure 10 NEEDLE WATER POTENTIAL vs SOIL WATER POTENTIAL FOR CONTROL DOUGLAS-FIR SEEDLINGS  ( The different symbols respresent individual seedlings. Line is from Dosskey, 1978 )  - 53 -  -3.0 _i  Soil Water Potential (i/'s) in MPa -1.0 L_  /  /  /  /  /  /  /  /  /  /  /  ca  CL  2 L-1.0 ca  +-« c CD  o  CL i_ CD  -2.0  /  o  •o co o  z  -3.0  Figure 11 N E E D L E WATER POTENTIAL vs SOIL WATER POTENTIAL FOR PLANTED DOUGLAS-FIR SEEDLINGS  ( The different symbols represent individual seedlings. Line is from Dosskey, 1,978 )  - 54 -  Soil Water Potential OAs) in MPa -1.0  Figure 12 N E E D L E WATER POTENTIAL vs SOIL WATER FOR P L A N T E D AND VIBRATED DOUGLAS-FIR  POTENTIAL SEEDLINGS  ( The different symbols represent individual seedlings. Line is from Dosskey, 1978 )  - 55  solid  line  shown  is  (1978) i n h i s study ences  in  the  rigorous all data the  of  this  control  squares  methods  on  study  between  (Figure  points,  and  the  fitted loamy  of r e s u l t s i s not  i n d i v i d u a l data  from  least  Douglas-fir  sampling  comparison  the  the  -  line  sand.  this  study  appropriate.  there  i s general  regression  lines  and  the  planted  and  potential  for  the  planted  treatment  by  Dosskey  Because  of  differ-  and  Dosskey's,  However, l o o k i n g agreement between  from  10)  derived  vibrated  Dosskey  a at the  (1978) f o r  (Figure  11) t r e a t -  ments.  Needle  water  compare,  p a r t l y because  treatment fore,  a  Looking  to reduce  trend,  at  data  increasing  needle  suggested, at l e a s t increased browning  soil  linear the  stomatal and  i t was water  even points  water  not  possible  for  i s more the  p o t e n t i a l below about  i f i t did for  exist,  cannot  seedlings -1.0 be  i n d i v i d u a l seedlings,  p o t e n t i a l as  f o r some of the  soil  of  needle  seedlings.  tips  was  when s o i l water p o t e n t i a l s were near -0.5  in  to this  MPa.  There-  clearly  shown.  a  trend  toward  water p o t e n t i a l d e c r e a s e s T h i s may  c l o s u r e i n response to water s t r e s s .  hooking  difficult  observed  on  be a r e s u l t  is of  Some y e l l o w i n g ,  several  seedlings  MPa.  Uptake  Figures  13,  potentials  14  and  15  for  the  three  by  Dosskey  curve developed  show the  uptake  treatments  values  along  at  with  (1978) f o r D o u g l a s - f i r  different the on  uptake loamy  soil  water  regression sand.  Con-  - 56 -  (The different symbols represent individual seedlings. Curve is from Dosskey, 1978) - 0.3  6) •  Soil Water Potential  (MPa)  Figure 13 U P T A K E R A T E FOR CONTROL DOUGLAS-FIR SEEDLINGS  E  - 57  (The different symbols represent individual seedlings. Curve is from Dosskey, 1978)  Soil Water Potential (MPa)  Figure 14 U P T A K E R A T E FOR PLANTED DOUGLAS-FIR SEEDLINGS  8  - 58 -  0.4  (The different symbols represent individual seedlings. Curve is from Dosskey, 1978) 0.3  co  »E ^ cn  h0.2 .A  a  CD *-< CO  CC CD J£ CO  a 13  h0.1  •  © e •3.0  -2.0  -1.0  Soil Water Potential (MPa)  Figure 1 5 U P T A K E R A T E FOR PLANTED AND VIRBRATED DOUGLAS-FIR SEEDLINGS  - 59  sidering  differences in seediing  -  size,  there  agreement between observed uptake r a t e s and (1978) f o r the seedlings -2.0  range  i s obviously  other  dicted with  the  of  13)  and water  the  planted  of  p r e d i c t e d r a t e s from Dosskey  planted  soil  degree  and  potential  vibrated from  (Figure  about  15)  -0.5  to  than  in  MPa.  Uptake the  in  c o n t r o l (Figure  i s a reasonable  two  from  the  less  treatments  regression  in and  i s considerably  (Dosskey,  above o b s e r v a t i o n s  on  1978).  needle  f i r m t h a t the s e e d l i n g s were i n f a c t  Resistance  less  These  water  low  (Figure than  14)  would  uptake  p o t e n t i a l and  be  rates  preagree  seem to  con-  s u f f e r i n g from water s t r e s s .  to Water Uptake  Observed r e s i s t a n c e curves  seedlings  f o r the  for Douglas-fir  Figures  16,  17 and  planted  and  vibrated  ments show g e n e r a l  18.  on  (Figure  i s probably  partly a result  Resistance  for  steeply  rising  18)  agreement with  and  than  loamy  treatments sand  Resistance  the p l a n t e d  greater  three  along  from Dosskey  f o r the  control  seedlings  with  the  resistance  (1978) are (Figure  i s s i m i l a r and  the r e s i s t a n c e c u r v e s .  shown i n  16)  and  both  treat-  Resistance  v i b r a t e d s e e d l i n g s appears to be somewhat lower but  the  for  of d i f f e r e n c e s i n s e e d l i n g  planted the  seedlings  other  resistance  for  shown  treatments. one  or  two  in  Also  for this  dimensions.  Figure a  the  trend  17 of  is  clearly  much  i n d i v i d u a l seedlings  as  more soil  - 60 -  h  o  100  o  o» •  r-10  h i I  •3.0  I I  l  l l i  l  I  "i  1 1 1 r  I  I  |  I  I  I  l  l  I  l  -1.0  2:0  Soil Water Potential  I l 0  (MPa)  Figure 16 R E S I S T A N C E TO UPTAKE FOR C O N T R O L DOUGLAS-FIR SEEDLINGS.  (Thie different symbols represent individual seedlings. Curve is from Dosskey,1978) (Error bar represents 95% confidence interval for resistance, shown at mean value fortes)  - 61 -  -3.0  -2'.0 Soil Water Potential  -1.0 (MPa)  0  Figure 17 RESISTANCE TO UPTAKE FOR PLANTED DOUGLAS-FIR SEEDLINGS  (The different symbols represent individual seedlings. Curve is from Dosskey ,1978) (Error bar represents 95% confidence interval for resistance, shown at main value for^s)  - 62 -  ~  MOO  'co CO  CO  0. r«^  O r-10  CU  o c  CO CO CO CU  CC |  I  -3.0  I  I I  I  I  I I  l  |  I  l  I  l  I  I  l  I  -2.0  l  |  -1.0  Soil Water Potential  (MPa)  Figure 18 RESISTANCE TO U P T A K E FOR PLANTED AND VIBRATED DOUGLAS-FIR SEEDLINGS  (The different symbols represent individual seedlings. Curve is from Dosskey, 1978) (Error bar represents 9 5 % confidence interval for resistance, shown at mean value forV's)  - 63 -  water p o t e n t i a l d e c r e a s e s For at  the other least  gets  planted  water p o t e n t i a l s of about - 1 . 0 MPa.  resistance  appears  unit  effect.  a n a l y s i s of r e s i s t a n c e data root  area  Treatment  basis  showed  means were  s e e d l i n g s to be s i g n i f i c a n t l y vibrated seedlings  seedlings  (cc =  similar regardless  unit  needle  analyses  area  a  more  slowly  As the s o i l than  f o r the  different  area  needle  significant  area  treatment  resistance  i n planted  from c o n t r o l and the p l a n t e d and  However,  of whether root  highly  and showed  different  0.05).  or u n i t  on a s e e d l i n g , u n i t  very  tested  are not s i g n i f i c a n t l y  were  The  to i n c r e a s e  constant,  seedlings.  Covariance and  i s apparent.  two t r e a t m e n t s , r e s i s t a n c e appears to be n e a r l y  down t o s o i l  drier,  - 0 . 1 and - 1 . 0 MPa  between about  data  the from  were  basis.  planted  and  vibrated  the c o n t r o l .  Results  analyzed  on  a  D e t a i l s of the  seedling,  statistical  are i n Appendix V I I .  analyses  attributable coefficient  also to of  showed  a very  variation  among  intraclass  highly  seedlings  correlation  s e e d l i n g s w i t h i n a treatment  significant within  (Sokal  variance a  and  component  treatment. Rohlf,  The  1981) f o r  i s 0 . 9 2 which can be i n t e r p r e t e d as showing  t h a t 9 2 % o f the v a r i a t i o n w i t h i n a treatment i s accounted f o r by d i f f e r ences  between  variability  Significant  individual  seedlings  and  only  8%  i s attributable to  w i t h i n an i n d i v i d u a l s e e d l i n g .  variability  between  i n d i v i d u a l seedlings  physiological differences, variability  may  result  from  i n the growth chamber environment  - 6k -  or d i f f e r e n c e s i n extent ferences  i n the growth  of m y c o r r h i z a l  development.  Environmental d i f -  chamber appear to be r e l a t i v e l y  minor; a l s o , t h e  e f f e c t s of any d i f f e r e n c e s were accounted by randomly changing the l o c a tion  of s e e d l i n g s  during  for  ectomycorrhizal  development  appeared  individual  seedlings.  in  the  extent  seedlings. probably  the experiment.  development to  be  at  minor  the  and  However, even  of development  might  Seedling  roots  end  the  experiment  and  between  treatments  and  of  similar  some a p p a r e n t l y be  a  source  P h y s i o l o g i c a l d i f f e r e n c e s between  of  were  minor  examined  differences  variation  seedlings,  between  however,  are  the major source of v a r i a b i l i t y .  DISCUSSION  Analysis higher  of  resistance  data  shows  that  planted  seedlings  i n the s o i l .  I t i s speculated  that  roots  growing s e e d l i n g s can e s t a b l i s h i n t i m a t e c o n t a c t reduced s o i l - r o o t  The  results  hypothesis.  of  contact  vibrating  possible  seedlings alters  planted  seedlings  of such n a t u r a l l y the s o i l  to p l a n t e d seem  seedlings  to  resulting seedlings.  confirm  i s about  this  s i x times  and v i b r a t e d s e e d l i n g s .  explanations  are  that  vibration  themselves so as to lower t h e i r  soil  with  r e s i s t a n c e compared  Mean r e s i s t a n c e f o r p l a n t e d  g r e a t e r than f o r p l a n t e d  Other  a much  r e s i s t a n c e to water uptake than s e e d l i n g s which have been growing  naturally  in  have  d e n s i t y and pore geometry  somehow  affects  the  r e s i s t a n c e or t h a t v i b r a t i o n  resulting  i n reduced s o i l  resist-  - 65 -  ance.  The former  resistance ated  independently  seedlings  the  from e f f e c t s on the s o i l ,  lower s o i l  of  the  constant soil  or may  dries. is  potentials  plant  cannot be s u b s t a n t i density  and pore  r e s i s t a n c e which might account f o r the reduced  data  resistance  Calculations small  greater  than  -0.5  potentials  of  to MPa  MPa  density.  Nnyamah e t al. (1978)  sandy  loam  soil  about  -1.0 MPa  Douglas-fir Inasmuch  remained  resistance  these  would  total  (1960)  17  sandy  conditions  found  that  findings seem  to  small  and B a l l a r d  silt  loam  too s m a l l  to  to  and  Newman at s o i l rooting  in a  gravelly  (1980)  silty  account  small  normal  water  this  soil  loam  of  and  that  as  water  resistance  at s o i l  are a p p l i c a b l e  be  i s more o r  soil  account f o r no more than 3% o f t o t a l in  planted  at  be r e l a t i v e l y  under  Dosskey  growing  for  suggest  resistance  i n Pachappa  relatively  or g r e a t e r .  seedlings  as  Gardner  r e s i s t a n c e should  -0.7  resistance could  i n Figure  f o r one or two o f the s e e d l i n g s  by  compared  shown  f o r i n d i v i d u a l seedlings  be i n c r e a s i n g  (1969) concluded t h a t s o i l  soil  a c t to lower  However changes i n s o i l  resistance  suggests t h a t  resistance  water  v i b r a t i o n could  resistance.  Examination  less  that  by any e x i s t i n g e v i d e n c e .  geometry c o u l d total  possibility,  p o t e n t i a l s of inferred  that  resistance i n clay  soils.  experiment, f o r the  soil  observed  d i f f e r e n c e s between t r e a t m e n t s .  Laboratory soils  to  analyses be  characteristic  conducted  virtually and  at the end o f the experiment  identical  saturated  in  bulk  hydraulic  density,  water  conductivity.  show the retention  Unsaturated  - 66  hydraulic Jackson would  conductivity (1972).  increase  potential Figures  These  by  falls  16,  estimated  estimates  about t h r e e  from  17  was  and  -0.06 18  -  according  indicate  to four  to  -0.5  show no  to  that  the  method  bulk  soil  given  resistance  o r d e r s of magnitude as s o i l  MPa.  The  resistance  correspondingly  sharp  by  data  water  shown  increase  in  in  total  resistance.  Even though the they to  may  provide  total  soil  and  to the  l e s s of t o t a l  The  some i n s i g h t  resistance.  calculations the  c a l c u l a t e d c o n d u c t i v i t i e s are only  the  Soil  resistance  calculated  root  surface  contribution based  of  on  average p a t h l e n g t h  could  r e s i s t a n c e at a s o i l  soil  the  for  control,  0.14,  0.14  and  these  calculations,  cannot e x p l a i n  view  of  differences  the  the  water p o t e n t i a l of -1.5  vibrated  pathlengths  flow  from  f o r a maximum of about 2%  account  respectively.  above  resistance  It  i s therefore of  cm,  and  Given  should  evidence,  between treatments can  soil  contact  0.12  and  conductivity  be  planted the  MPa.  soil  to a  root  seedlings  are  errors  regarded  or  inherent  in  equal  and  as  d i f f e r e n c e s i n r e s i s t a n c e between t r e a t m e n t s .  in  result  planted  resistance  f o r water  c a l c u l a t e d average p a t h l e n g t h f o r water flow from the  surface  In  i n t o the  rough a p p r o x i m a t i o n s ,  or  by  inferred  differences  resistance  in  in  it be  e f f e c t s of that  appears  Douglas-fir  be  accounted  f o r only  vibration  on  differences  soil-root  to  between  contact and  other  unlikely  that  by  differences  seedling  physiology.  treatments  resistance. plant  may  be  a  Significant  species  has  been  - 67 -  inferred al.,  i n several  1977a,  studies  (Dosskey  and B a l l a r d ,  1977b; Nnyamah e t al., 1978) and F a i z  as a major component o f t o t a l  1980; H e r k e l r a t h e t (1973) i d e n t i f i e d i t  r e s i s t a n c e i n sunflower (Helianthus  annuus  L.).  The  significant  several  variance  reasons.  component  I t confirms  among  seedlings  observations  made  Dosskey and B a l l a r d (1980) t h a t c o n s i d e r a b l e apparently  similar seedlings  mental  conditions.  future  experiments.  With  reference  seedlings of  of s i n g l e  relatively  Although  information  also  be u s e f u l  in  designing  experimental e r r o r s  with  needles  from  associated  and sampling  errors  f o r detemination  the c o n t r o l l e d  i s problematic,  silvicultural  bareroot  seedlings  may  measurements  associated  of water  to the c a r e f u l p l a n t i n g  soil  resistance  findings High  practice, considerably  Therefore  high  experimental  significantly  compared  study.  the  implications.  In f i e l d  increased  may  environ-  component between  growth.  this  v a r i a t i o n can occur between  experiment, the l a r g e v a r i a n c e  extrapolation  important  (1978) and  with the  potential are  insignificant.  conditions  planted  that  by Dosskey  growing under s i m i l a r , c o n t r o l l e d  p o t e n t i a l and uptake  sampling  field  to t h i s  implies  water  Such  i s noteworthy f o r  study  to  have  resistance  in  a f f e c t s u r v i v a l and e a r l y i s taken  of the s o i l  resistance  can be expected  this  contact  l e s s care  and packing  contact  of  conditions  i n planting practiced i n  and, i n some  in field  planted  cases,  seedlings.  - 68  In a d d i t i o n promote  -  to c a r e f u l s t o c k h a n d l i n g and  rapid  root  growth  and  potential practical solutions  planting,  encourage  to the  c u l t u r a l measures to  mycorrhizal  development  are  problem.  SUMMARY AND CONCLUSIONS  R e s i s t a n c e to water uptake i n p l a n t e d , Douglas-fir ance  was  needle  seedlings  calculated  water  trolled  from  potential  and  for differences treatments and  the  Uptake r a t e s ,  direct  and  The  i n the  on  vibrated,  s o i l was soil  individual  and  control  compared. water  Resist-  potential,  seedlings  e x p e r i m e n t a l design used was  in  a nested  a n a l y z e d with c o v a r i a n c e a n a l y s i s  a  con-  analysis to  allow  range of s o i l water p o t e n t i a l encountered between  individual seedlings. u n i t root  area  Data were analyzed on  were c l o s e l y  a  seedling,  basis.  needle water p o t e n t i a l and  seedlings  and  measurements of  uptake  data was  u n i t needle area and  trol  i n loamy sand t e x t u r e d  environment.  of v a r i a n c e  planted  comparable  resistance with  those  values f o r the presented  by  con-  Dosskey  (1978) i n a s i m i l a r study i n which d i f f e r e n t t e c h n i q u e s were used.  Results  of  the  covariance  analysis  show  difference  between t r e a t m e n t s ; r e s i s t a n c e  six  times  greater  The  latter  also  two  indicates  than  for  treatments that  highly s i g n i f i c a n t .  control are  or  among  very  for planted  planted  statistically  variability  a  and  highly  seedlings vibrated  identical.  individual  significant  The  seedlings  i s about seedlings. analysis is  very  - 69 -  Increased resistance to water uptake in planted  seedlings  a result of greater  resistance.  the d i f f i c u l t y ity  s o i l - r o o t contact  in obtaining  for the s o i l s ,  and s o i l  appears to be  precise estimates of unsaturated  the r e l a t i v e contributions  resistance  cannot  be precisely  determined.  similarity  of s o i l  properties, such as texture,  of contact  Because of conductivand  However, because of the bulk density  and water  retention c h a r a c t e r i s t i c , among treatments i t seems unlikeiy that resistance resistance.  alone could  account  soil  for the observed  difference  soil  in t o t a l  - 70 -  CONCLUSIONS  R e s i s t a n c e to L i q u i d in  planted  growing soil  Douglas-fir  potential  C-51  and  PCT-55-05 s o i l ly  and  used  equal.  were  C-52  taken  R e s i s t a n c e was  of  studied  using  potted  steady  state  psychrometer  conditions  sample  uptake  then c a l c u l a t e d  since,  at  steady  and  using  chambers  T r a n s p i r a t i o n was measured  and  seedlings  Measurements of needle water  under  thermocouple  a measure  i n ponderosa pine s e e d l i n g s  was  environment.  psychrometers.  as  uptake  seedlings  in a controlled  water  Wescor  phase water  and  gravimetrical-  state,  they  based on the Ohm's Law  are  relation-  s h i p shown i n e q u a t i o n 2 (Chapter 1 ) .  Because  sampling  s e e d l i n g water estimates water  of  and  uptake  and  needle water  i s destructive  potential  f o r the study of ponderosa  resistance  relationships.  potential  b a l a n c e , i n d i r e c t methods were used to o b t a i n  potential  uptake  f o r needle water  pine.  were then generated from  In the study of p l a n t e d  as  and  upsets  independent  a function  of  soil  Continuous v a l u e s o f  the observed  Douglas-fir,  i t was  regression found  that  removal of a s m a l l number of n e e d l e s (up t o 10), c o m p r i s i n g about 1% o f the  total  balance.  needle  area,  Therefore,  had  in  no  this  detectable  experiment,  effect  on  seedling  measurements  of  water  uptake  and  water p o t e n t i a l were taken from the same s e e d l i n g s .  S e e d l i n g s i n both experiments were grown i n a loamy in  a  heated  experimental  greenhouse  for a  runs i n the growth  period  of  chamber.  about On  15  sand  months  textured  soil  prior  to the  completion of the  experi-  ments, s e e d l i n g and water pathway dimensions were determined.  - 71 -  R e s i s t a n c e t o water uptake i n ponderosa p i n e , on a u n i t r o o t area b a s i s , was  estimated using  f i r s t model was Douglas-fir, points  two  western  However,  at  soil  uptake  potentials,  nonlinear  regression  i d e n t i c a l t o t h a t used by Dosskey  for individual  resistance  different  hemlock  mountain  seedlings well water  was  and  potentials  consistently  resulting  in  of  about  low,  It  fitted  data  r e a s o n a b l e p r e d i c t i o n s of  overpredicted  artificially  The  (1978) i n h i s study of  hemlock.  and gave  models.  -1.5 at  MPa  or  lower  decreasing  higher.  soil  water  estimates  of  resistance.  The  second  model,  results  although  slightly  low  The  uptake  at s o i l  Weibull  water  of t h i s  function,  resistance potentials  model  between  i s that  This  limit  the  i s useful  because  a c c u r a c y and  gave  estimates  be  overall  artificially  -1.1  t o -0.2  constrained  MPa.  within  water p o t e n t i a l range where  limitations  number  better  are  about  i t can  l i m i t s at the extremes of the s o i l  occurs.  techniques  modified  resulting  main advantage  realistic  a  of  of the e x p e r i m e n t a l  observations  that  can  be  made at these extremes.  R e s u l t s of the study of ponderosa for  Douglas-fir  by Dosskey  (1978).  pine appears to remain v i r t u a l l y uptake  rates  are  relatively  decrease more r a p i d l y about -1.5 MPa,  p i n e were compared  potential  c o n s t a n t as the s o i l  high  as the s o i l  Needle water  at  dries.  high  soil  to those  dries.  water  reported  i n ponderosa Generally,  potentials  but  At s o i l water p o t e n t i a l s above  r e s i s t a n c e to water uptake i n ponderosa p i n e seems to be  l e s s than i n D o u g l a s - f i r .  However, r e s i s t a n c e  i n ponderosa pine (based  - 72 -  on  the second  uptake  below  -1.5  These  observations  species  are  in  (Lopushinsky and  uptake  stomata  rapidly  at s o i l  water  potentials  MPa.  agreement  concerning t r a n s p i r a t i o n  potential in  model) i n c r e a s e s  f o r ponderosa  to  decreasing  with  as a f u n c t i o n  K l o c k , 1974). pine  other  may  The  reflect  availability  of d e c r e a s i n g relatively  the  of  comparisons  these  soil  water  rapid  greater  water  of  decline  sensitivity  (Lopushinsky,  of  1969;  Lopushinsky and K l o c k , 1974).  The  source of  rapidly  increasing  resistance  from the second uptake model at s o i l difficult at  the  to i s o l a t e . soil-root  Differences  water  to water  uptake  p o t e n t i a l s below  -1.5  I n c r e a s i n g p l a n t r e s i s t a n c e or c o n t a c t  interface  observed  appear  in resistance  to  be  to water  the  most  uptake  predicted is  resistance  likely  between  MPa  sources.  Douglas-fir  and ponderosa pine do not seem to be e x p l a i n a b l e by d i f f e r e n c e s  in  soil  resistance.  The  differences  ences  in ecological  pioneer  tree  transpiration excessive early be  observed between the s p e c i e s may  species as  the  needle  advantageous  in soil  Coastal  southwestern dries  temperatures  successional  competing  strategies.  may  Douglas-fir  British be  reflect  by  any  water  conserved would  shrubs or other v e g e t a t i o n .  Ponderosa  in  Maintaining  order  to  avoid  Duhme (1974) f o r other  s p e c i e s ) ; furthermore, conserving  since  differ-  grows mainly as a  Columbia.  necessary  (as suggested  broad  soil  likely  water be  may  depleted  not by  pine o c c u r s mainly i n  - 73  dry  interior  these the  regions  conditions,  or  in  moist  dry  soil  -  sites  in  conditions,  s p r i n g , are e x p l o i t e d by m a i n t a i n i n g  rates  of  t r a n s p i r a t i o n and  depleted  as  Several pine  the  other  including:  (Lopushinsky,  establishment  of  a  of  the  study  seedlings,  of  three  planted,  and  1969, deep  1973;  increase  in  and  resistance  were  therefore  appears t h a t  a  of  result  basis  of  (such  as  the  (Mirov,  to  the  similarity bulk  in high  rapidly  conserved.  for  ponderosa  avoidance  strategy  Klock,  in  were  uptake  of  1974),  1957),  and  to  the  soil rapid  needles  and  Douglas-fir  compared:  control,  show  planted control  resistance  treatments  planted  seedlings  while  soil-root  density  i s then noted  Lynch,  Planted  identical  among  is  availability  and  uptake  treatments  and  mainly  1967).  increased  soil  decreasing  water  water  statistically  texture,  to  vibrated. to  increased  water  Under  therefore  moisture  drought  ( C u r t i s and  experimental  seedlings  and  Lopushinsky  taproot  resistance  planted  Soil  this  stomata  with a t h i c k c u t i c u l a r l a y e r  In  open stomata and  progresses  facilitate  climates.  occurring  p h y s i o l o g i c a l adaptations  pines  sensitivity  moist  probably  photosynthesis.  season  m o r p h o l o g i c a l and  and  water  growing  more  a  and  vibrated  seedlings.  i n planted  resistance.  in  physical  water  retention  It  seedlings  contact soil  marked  On  resistance  B a l l a r d , 1980; 1977a;  and  other  F a i z , 1973;  Newman, 1969b;  Ritchie,  evidence F a i z and  3arvis,  1976), i t i s concluded  i n the  literature  Weatherley, 1978;  1975; that  Nnyamah et soil-root  al.,  characteristic),  (see  Dosskey  Herkelrath 1978;  contact  the  properties  rough approximations of u n s a t u r a t e d h y d r a u l i c c o n d u c t i v i t y , e s t i m a t e s soil  is  et  and al.,  Reicosky  resistance  of  and  i s an  - 74  important source of the  Comparing  the  data  on  increased  needle  -  total  resistance.  water p o t e n t i a l , uptake r a t e s  ance t o uptake from c o n t r o l and  the  planted  and  and  vibrated seedlings  t h i s experiment to the c u r v e s generated by Dosskey (1978) u s i n g techniques The  for estimation,  d i f f e r e n c e s that  ences and  i n the  seedling  i s a remarkable  apparent  could  populations,  be  degree  of  in  indirect  similarity.  l a r g e l y explained  differences  from  by  seedling  differ-  dimensions  sampling e r r o r .  Statistical ance  a n a l y s i s of the  component  statistical unit  needle  pathway  among  area to  observed  variability  or  unit  also  do  variability.  (1978)  and  data  individual seedlings; the  root  differences  dimensions  Dosskey  experimental  s i g n i f i c a n c e whether  attributable  the  are  there  resist-  in not  appear  Dosskey  seems  and  variance  i s analyzed  basis  seedling  This  the  data  area  shows a s i g n i f i c a n t  and  different to  Ballard  among D o u g l a s - f i r s e e d l i n g s was  on  a  seedling,  the  (1980)  is  Calculated  enough  confirm  maintains i t s  therefore  dimensions.  vari-  to  not water  account  for  observations  that  of  physiological  the major source  variability  i n p r e d i c t e d uptake r a t e s .  The  increased  important the  field,  to  ensure  seedlings.  r e s i s t a n c e to  silvicultural although  water uptake  implications.  i t s importance  satisfactory survival The  seedlings  in this  i n planted  C a r e f u l handling  i s recognized, or  seedlings  early  growth  experiment  may of  of  not  has  seedlings be  in  sufficient  planted  were p l a n t e d  some  with  bareroot a  great  - 75 -  deal more care than would be feasible i n operational  f i e l d planting but  they s t i l l show a s i g n i f i c a n t increase in resistance.  Nursery practices  which promote early and vigorous root growth and other c u l t u r a l measures such as inoculation with mycorrhizal  fungi may be important,  particu-  l a r l y when seedlings are planted i n dry s i t e s or they encounter warm dry weather during or immediately following planting.  - 76 -  LITERATURE  Andrews, R.E. and E . I . Newman. and  plant:  I I I . Evidence  CITED  1969. 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Water t r a n s p o r t i n p l a n t s as a c a t e n a r y  D i s c u s s i o n s of the Faraday S o c i e t y No.  1973.  Vegetation  S p r i n g e r - V e r l a g N.Y.  Waring, R.H.  22(1):5-10.  1970.  Inc.  of the e a r t h . 237  E n g l i s h U n i v e r s i t y Press,  p.  Matching s p e c i e s to s i t e .  r e g e n e r a t i o n of ponderosa p i n e . Sen. F o r . Oregon  State Univ.  3:146-153.  R.K.  pp. 54-61  In:  Proc.  Hermann (ed.) F o r . Res.  C o r v a l l i s , Ore. 97331.  Lab.,  - 85 -  W e i b u l l , W.  1951.  A statistical  applicability.  Wescor.  1979.  3. Appl. Mech.  18:293-296.  I n s t r u c t i o n Manual.  Wescor, Inc. Logan, Utah.  Yang, R.C.,  d i s t r i b u t i o n f u n c t i o n of wide  C-51  f u n c t i o n s as f l e x i b l e  Res. 8(4):424-431.  psychrometer.  15 p.  A. Kozak and 3.H.G. Smith.  Weibull-type  sample chamber  1978.  The p o t e n t i a l o f  growth c u r v e s .  Can. 3. F o r .  - 86 -  APPENDIX I S p e c t r a l I r r a d i a n c e i n the E x p e r i m e n t a l  Environment  -  87  -  1020 -,  ~ - 823  A  .30  .41  .52  .63  .74  W a v e l e n g t h (ftm)  Figure IA SPECIAL IRRADIANCE IN THE GROWTH  CHAMBER  (Source: F.M. Kelliher, University of B.C., Dept. Soil Science) (Measured at 54cm below lights, sampled at 1nm intervals)  - 88 -  .30  .41  .52  .63  .74  Wavelength (^im)  Figure IB S P E C T R A L IRRADIANCE ON A CLEAR DAY (NOVEMBER 11, 1981) AT LOGAN , UTAH  (Sampled at 1 nm Intervals) (Source: LI-COR .INC.)  .85  - 89 -  APPENDIX I I S t a t i s t i c a l Summary f o r R e g r e s s i o n o f Needle Water P o t e n t i a l as a F u n c t i o n of S o i l Water P o t e n t i a l f o r Ponderosa Pine  - 90 -  ANOVA Source Regression  SS  MS  0.028617  0.028617 0.098025  d.f. 1  Residual  45  4.4111  Total  46  4.3973  Regression i s not s i g n i f i c a n t at  11  F 0.29194 n.s.  = 0.05  Standard error of the regression c o e f f i c i e n t = 0.038534 95% confidence i n t e r v a l f o r the regression c o e f f i c i e n t = -0.05675 t o 0.09839 Standard error of the sample mean = 0.04567 95% confidence i n t e r v a l f o r the sample mean = -2.31 t o -2.13 MPa  - 91 -  APPENDIX I I I C a l c u l a t i o n s f o r Uptake and R e s i s t a n c e f o r Ponderosa P i n e  >  ui & u M  ooooo ^  oo  o ^  ) m > ) > -n <  t-i  nor  > -1 m O  co > o ^ ^— co _L J - — > Ul Ul Ul 73 w w w ir II II n c/l O CO > c CO CO C0 2 J> J> > O 73 73 73  ca ca J> 73 n t/l c 2 CO W  > cn > 73 II t/l c 3 > \  £. ^  IO  n o Z —I  i/i c 3 O  to i/i o c c o 2 3 CO J> CJ II II II O Z l/l l/l i/i o c c c c m s 3 3 rn co > II + + + — o ca > -  tH -  r~ r  * CO z 1 H O : — i/i o H Ul m i w > -n  r~ u  > r~ o c r > H -  Z cn  < 73 > cn m i/i  x *~o  *-*-~C  u o oo ooo-  -*  oo  Ul *  oooo  8 8  o o r- r 73 0 73 TJ O O C 73 O O m TJ TJ TJ TJ o o o o l/l H 1/1 1/1 l/l o o to TJ TJ m C >-H >-H O H CO > Z Z M c c c rm s e c w C_ II II II Z O TJ II I C. • ll 13 U * c -> O O 1/1 "D H H TJ > l / l • it II -H Z Z C/l « -«• I - - O H — > G3 H - - m Z ^ •—- C_ fO • ro — z a 7; (/> C/l M j i m m C_ * ^ 01 -1 m —• CO O > O - - 73 X > Z > • z + _» —. > m - Z o r —i i / i ui o o rn -n C *fc m 73 l/> Ul »-H > i/i < J I H m rn > J J -n Z > m O O 73 m m  00 *1 01  -  o •n  Z j> i/i  o o m -n -n  C TJ M »  o z H l/l  %  > Z — o ui m  •n O 73 2 >  73 m > O  l/l C 2 > -  > ca i> 73  Ul l/l - C CO U — 2 CO •n w CO > - 73 O J> l/l Ul ~ C > X 2 Z w o a CO —* X w • o X w X  II  — — •fc.  > Ci Ci CO O > C 33 2 c » J3 > m H m > < O m O 73 m > n Q -n m o o <-< a m m Z -n — 1 TI l/l »-< Ci -n *-* O m 73 Z •H O l/l J> r O c -  I  -  > -1 Z o  § 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o 0o  oooooooooooooooooooooooooooo  -  X J> - 73 >-<73 CD Ci m z m > • > H2 73 — — I m m Ul o 2 O O O O O O — l/l rn CO - TJ 73 m J> l/l 73 CO ' i—i > ^v- H 73 N) C O • 03 w X O CO - - m n 73 TJ - r— - d 2 > O *-> - 73 CO -s z m l/l c 2 O  l/l c 2 ca  i/i c 3 >  O CO > 73  CO CO J> 73  TJ 73 O tn 73 >  3 -n TJ O 73 m l/l m m a  TJ Z 73 O >- l/l  >zr >  IO  l/l C 2 >  • O -0 l/l  73 *w  l/l C 2 CO l/i .C s Ci  ^  IO  A w  c TJ -< X Ul (O  73 m l/l Ul ro  Z H m O m 73 irt 2  V  z TI  o 73 -rt O 73 2 J> H —1 m O  7] m m 73 i/i >  > O Z O r> m  O C —1 T) rz -i  ) O O O O Q O O O O O O O O O O O O O O O O O O O O O O O O O ) O O O O O O O O O O O O O O O O O O O O O O O O O O O O Q O ) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0  26 -  r  C C  c  C O U T P U T : W R I T I N G RES AND U P T K FOR 14 S E E D L I N G S AND FOR AVERAGE C AVERAGE IS » 15 C W R I T E S RES AND U P T K FOR 3 S E E D L I N G S (OR AVERAGE FOR # 1 5 ) PER P A G E C 32 DO 4 0 0 M = 1 , 1 5 , 3 33 WRITE(6,40S)M,M+1.M+2 •EXTENS ION* OTHER C O M P I L E R S MAY NOT ALLOW E X P R E S S I O N S IN OUTPUT L I S T S •EXTENS ION* OTHER C O M P I L E R S MAY NOT ALLOW E X P R E S S I O N S IN OUTPUT L I S T S 34 4 0 5 F D R M A T ( 2 X , ' S E E D L I N G * " , 1 2 , BX . ' S E E D L I N G # ' . I 2 , 8 X , ' S E E D L I N G * ' . I 2 ) C THANK YOU FOR THE E X T E N S I O N S ON FORMAT WRITE S T A T E M E N T # 4 0 5 C C WRITE(6,415) 35 415 F O R M A T ( 3 X , ' R E S ' , 7 X , ' U P T K ' 6 X , ' R E S ' , 7 X , ' U P T K ' , 6 X , ' R E S ' . 7 X . ' U P T K ' ) 36  37 38 39  41 42 43  DO 5 0 0  N=1,24,1  W R I T E ( 6 , 5 0 5 ) R E S ( M . N ) , U P T K ( M , N ) ,RES(M+1 , N ) , U P T K ( M + 1 , N) , RES ( M + 2 , N) , U XPTK(M+2,N) 5 0 5 F 0 R M A T ( 3 ( F 1 0 . 4 , - F IO. 4 ) )  5O0  CONTINUE  400  CONTINUE STOP END  /E XECUTE O.1493301E O . 3 9 8 7 8 4 2E 01 SEEDLING # SEEDLING 1 RES RES UPTK O.1573 0.2673 7.8355 .3247 . 1654 1492 .3868 . 1730 . 9086 .4519 . 1798 . 9856 .5174 . 1856 .2916 .5805 . 1904 . 765 1 .6379 . 1939 .3627 .6863 . 1960 .0535 . 7224 . 1964 1.8154 . 7433 . 1951 .1 . 6 3 2 4 0.7471 . 1918 1.4930 0.7326 . 1865 1.3890 0.6997 . 1791 1.3144 0.6495 . 1694 1.2652 O. 5844 . 1574 1.2387 0.5074 . 1431 1.2335 0.4226 . 1265 1.2495 O. 3341 . 1076 1.2873 O. 2462 0.0864 1.3491 O. 1629 0.0632 1.4381 0.0874 0.038O 1.5593 0.0223 0 . 0 1 10 1.7197 -O.O308 -0.0175 1.9291 -0.0715 -0.0474 2.2012  64.000 65.000 66.000 67.000 68.000 69.000 70.000 71 . 0 0 0 72.000  73.000 74.000 75.000 76.000 77.000 78.000 79.000 80.000 81 . 0 0 0 82.000 83 . 0 0 0 84.000 85.000 86.000 87.000 88 . 0 0 0 89 . 0 0 0 90.000 91 . 0 0 0 92.000  2 UPTK 13.3196 12.0712 10.9777 10.0178 9.1734 8.4293 7 .7723 1913 .6768 2205 8154 4556 1357 8512 5984 3738 1746 9983 3.8426 3.7058 3 . 5862 3.4825 3 . 3935 3.318 1  ! 1 1 1 9 3 3 E 01 SEEDLING # RES O.1524 . 1710 . 1902 . 2095 .2285 . 2467 .2636 .2786 . 2910 .3002 . 3056 .3067 . 3028 . 2935 .2786 . 2579 .2313 . 1989 .1611 . 1 184 . 0 7 13 .0206 -0.0326 -0.0875  3 UPTK 13.7456 11.6756 9.9851 8.5976 7.4534 6.5056 5.7171 5.0584 4.5062 4.0416 3.6497 3.3183 3.0375 2.7995 2.5978 2.4270 2.2829 2.1621 2.0616 1.9792 1.9131 1.8617 1.8242 1.7996  SEEDLING  RES 0 .0317 O .0393 0 .0482 O .OS84 O .0699 0 .0827 0 .0965 0 .1111 0 . 1262 0 .1411 0 . 1554 0 . 1683 O . 1788 O . 1861 O . 1892 0 . 1871 0 . 1789 O . 1636 0 . 1405 O . 1092 0 .0694 O .0212 -0 . 0 3 5 1 -o .0987  SEEDLING  4  UPTK  66 . 1764 50 .8324 39 . 4 0 9 2 30 . 8 3 7 1 24 . 3 5 4 0 19 . 4 1 2 8 15 . 6 1 7 9 12 . 6 8 1 7 10 . 3 9 3 3 8 .5971 7 . 1774 6 .0478 5 . 1434 4 .4150 3 . 8249 3 .3445 2 .9517 2 .6292 2 . 3637 2 . 1448 1.9643 1.8157 1 .6939 1 . 5950 H 7  UPTK RES O .. 0 9 9 3 21 . 0 8 8 6 O . 1091 18 . 2 9 9 3 0 . 1 190 15 . 9 5 8 3 13 . 9 8 6 1 0.. 1 2 8 8 12 . 3 1 8 9 0 . 1382 0 . 1472 10.. 9 0 4 5 . 1554 9 .7007 O. 8 . 6729 0 . 1625 7 .7926 O . 1683 O . 1724 7 .0 3 6 6 6 . 3857 O . 1747 • 5. 8 2 3 9 0. 1747 O . 1723 5 . 3381 4 .9 1 7 2 0. 1671 4 . 5520 0. 1 5 9 0 4 . 2351 0. 1478 3 .9598 O . 1333 o . 1 156 3 .7209 3 .5139 0. 0 9 4 5 o . 0702 3 .3350 3 . 1809 0 . 0429 3 .0 4 9 1 0.0 1 2 6 - o . 0203 2 .9374 2. 8 4 3 9 - 0 . 0554 SEEDLING #10 UPTK RES 0 . O680 3 0 . 8154 0. 0767 26 . 0 3 9 9 22 . 1349 0. 0858 18 . 9 2 6 9 0.0 9 5 2 16 . 2 7 9 8 0 . 1046 0. 1 140 14 . 0 8 5 9  SEEDLING RES  0 . 2953 0 . 3559 0 .4206 0 .4873 0 .5534 0 .6156 0 .6706 0 .7150 0 . 7458 0 .7604 0 . 7570 0 .7352 0 . 6953 0 .6390 0 .5691 0 . 4890 0 . 4030 0 .3152 0 . 2297 0 , 1503 0 .0798 0 ..0201 - 0 .. 0 2 7 5 0630  -o.  SEEDLING RES  0 . 1339 0 . 1880 0. 2562 0 . 3391 0. 4355 0. 5427 0. 6558 0. 7682 0. 8717 0. 9575 10.1 7 1 10.4 3 6 10.3 2 7 0. 9835 0. 8988 0. 7849 0. 6509 0 . 5074 0. 3650 0. 2334 0 . 1 199 0. 0290 -0. 0376 - 0 . 08 10  SEEDLING RES 0. 0. 0. 0.' 0. 0.  2219 2052 1923 1827 1759 1715  5 UPTK  SEEDLING # 6 RES UPTK  8  SEEDLING RES  7 .0932 5 .6102 4 .5143 3 .6955 3 .0776 2 .6075 2 . 2476 1.9709 1. 7 5 8 3 1. 5 9 5 8 1. 4 7 3 5 1.3841 1. 3 2 2 8 1. 2860 1.2720 1. 2800 1 . 3 103 1. 3 6 4 6 1. 4459 1. 5 5 8 5 1.7091 1. 9 0 6 7 2 . 1641 2 .4989  UPTK  15 . 6 4 2 3 10 . 6 2 2 8 7 .41 10 5 .3115 3 .9106 2 .9579 2 . 2983 1. 8 . 346 1 . 5044 1. 2673 1.0968 0 .. 9 7 5 1 0 8905 0.. 8 3 5 6 0 .. 8 0 5 4 0. 7975 0. 8112 0. 8477 0. 9 1 0 0 10.0 3 6 1 1. 3 7 0 1 3.2 3 3 1 5. 8 2 2 1 9.4 3 4 1  UPTK 9. 4 3 7 7  Q 73 18 9 . 8738 9 . 8568 9 .6 8 1 6 9 .3 5 6 6  0 . 1799 0 . 1677 0 . 1578 0 . 1499 0 . 1436 0 . 1389 0 . 1354 0 .1331 0 . 1317 0 .1312 0 . 1315 0 . 1323 0 . 1335 0 . . 1347 0 . 1357 0 . . 1358 0 . . 1342 0 . . 1296 0 . 1202 0 . 1029 0. 0736 0. 0258 -0. 0504 1699  -o.  0 . 1343 0 . 1476 O. 1 6 1 0 0 . 1743 0 . 1872 0 . 1994 0 . 2105 0 . 2202 0 . 2282 0 . 2339 0 . 2370 0 . 2371 0 . 2338 0. 2269 0 . 2159 0 . 2007 0 . 181 1 0 . 1570 0 . 1284 0. 0954 0 . 0582 0 . 0171 0275 0753  -o. -o.  SEEDLING RES 0. 0. 0. 0. 0. 0.  1824 1655 1513 . 1392 1290 1202  11 . 6 4 3 0 1 .1 9095 12 . 0 3 6 0 12 .0181 1 .18 5 6 3 1 .1 5565 1 .1 1292 10 . 5 8 9 2 9 .9547 9 . 2460 8 .4848 7 .6929 6 .8913 6 .0993 5 .3336 4 .6080 3 .9335 3 .3174 2 .7643 2 . 2758 1 .8512 1 ,4877 1 .1813 0 .9267 # 9  UPTK  15 . 5 9 6 3 13 . 5 2 8 3 1 .1 7932 10. 3321 9. 0974 8. 0 5 0 2 7 . 1593 6 . 3988 5. 7477 5 . 1887 4 . 7075 4 . 2922 3. 9332 3. 6 2 2 3 3 . 3526 3. 1 186 2 . 9154 2 . 7390 2 . 5862 2 . 4542 2 . 3405 2 . 2433 2. 1609 2. 0 9 1 9 #12  UPTK  1 1 483 .5 12. 0628 1 2 . 5509 12. 9346 1 3 . 2035 1 3 . 3499  O.1229 12 . 2598 0 . 1694 8 .8972 0 . 1 127 13 . 3697 O.1313 10 . 7338 0 . 1693 8 . 3243 0 . 1063 13 .2623 0.1387 9 . 4533 0 .17 11 7 .6631 0 . 1006 13 .0308 . 3750 0.1449 8 0 . 1748 6 .9410 0 .0957 12 .6817 7 . 4636 0.1495 0 . 1803 6 . 1859 0 .0913 12 .2247 0.1521 6 . 6908 0 . 1876 5 . 4243 0 .0872 1 1.6722 4 .6800 0 . 1965 0.1524 6 .0336 0 .0833 1 1.0388 5 .4731 0.1501 0 .2068 3 .9729 0 .0795 10 . 3406 4 . 9942 0 .2181 3 .3184 0 . 1449 0 .0754 9 .5945 4 . 5842 0.1365 2 .7272 0 . 2295 0 .0710 8 .8 177 4 . 2327 . 2394 0.1247 0 2 . 2053 0 .0658 8 .0268 3 . 93 14 0.1094 0 . 245 1 1 . 7546 0 .0594 7 . 2374 3 .6731 1 .3735 0.0904 0 . 24 18 0 .0514 6 .4636 3 .4522 0.0679 0 .2214 1 .0580 0 .0410 5 .7178 3 . 2638 0.0418 0 . 1700 0 .8018 0 .0272 5 .0099 3 . 1039 0 .0643 0.0124 0 .5979 O .0088 4 . 3480 2 .9694 -0.0200 -0 . 1357 0 .4387 -0 .0159 3 . 7376 2 .8575 -0.0551 -0 . 497 1 0 .3167 -0 .0495 3 . 1825 SEEDLING ,S<13 SEEDLING #14 SEEDLING #15 RES UPTK RES UPTK RES UPTK O.0680 30 .8227 0 . 1518 13 . 7956 0 . 1036 20 .2146 25 . 472 1 0.0784 0 . 1648 12 . 1 163 0.. 1 156 17 .2715 21 . 1916 0.0896 0 . 1776 10 .6901 0. 1276 14 .8758 0.1015 17 . 7488 0. 1901 9 .4747 0.. 1394 12..9156 O.1138 14 . .9651 0. 2019 8 .4359 1507 0. 1 1 3039 . 12 .7026 0.1264 0. 2127 7 .5453 0. 1609 9. 9731 0.1389 10. 8546 0. 2223 6 .7795 0. 1699 8. 8697 9 .3377 0.1509 0. 2303 6. 1 192 0. 1772 7. 9520 0.1622 8 . 0868 0. 2363 5 .5484 7 .1866 0. 1825 7 .0504 0.172 1 0. 2401 5 .0539 0. 1853 6 .5472 0. 2412 4 .6244 0.1803 6 . 188 1 0. 1855 6.0127 O.1861 5 .4677 4 .2507 0. 2394 0. 1823 5. 5662 4 .8636 0.1891 0. 2343 3 .9251 0. 1770 5 .1945 4 .3553 0.1887 0. 2257 3 .6410 0. 1682 4. 8866 3 .9263 0.1844 0. 2133 3 .3928 1562 4 .6339 0. 3 .5633 0.1757 0. 1971 3 .1760 4 .4297 0. 14 13 3 .2556 0.1622 0. 1768 2 .9866 4 .2686 0. 1237 2 .9944 0.1436 0. 1524 2 .8214 4 .1465 .0. 1037 7727 • 2. 0.1198 0. 1241 2 .6774 0. 0818 4 .0603 2 .5846 0.0906 0. 0918 2 .5524 0. 0584 4 .0079 0.0562 2. 4254 0. 0558 2 .4443 0. 0342 3. 9881 2 .29 13 0.0168 0. 0163 2 .3515 0096 4. 0. 0003 -0.0273 2. 1792 -0. 0262 2 .2725 4 .0448 -o. 0147 2 .0865 -0.0754 -0. 0714 2 .2062 4 .1228 -o. 0382 CORE USAGE DIAGNOSTICS COMPILE TIME= Execution  $SIGN0FF  OBJECT CODE=  2096 BYTES,ARRAY AREA=  NUMBER OF ERRORS'  0, NUMBER OF WARNINGS'  0.071 SEC,EXECUTION TIME'  /STOP terminated  19:08:08  T=0.197  3348 BYTES,TOTAL AREA AVAILABLE'  RC=1  0.113 SEC, $0.18  102400  0, NUMBER OF EXTENSIONS'  WATFIV - JUL 1973 V1L4  19:08:08  BYTES 2  THURSDAY  1 2 3 4 5 6 7 8 9 10  1 1 12  13 14 15 16 17  18 19 20 21 22 23  24  25 26 27 28 29 30 31 32 33 34  /COMPILE FOR S E E D L I N G S 1 T O 1 5 : #15 R E F E R S 10 AVERAGE C O E F F I C I E N T S C PROGRAM P R I N T S C A L C U L A T E D UPTAKE AND R E S I S T A N C E FOR E A C H S E E D L I N G C C U P T A K E P R E D I C T E D FROM A M O D I F I E D WEIBULL P R O B A B I L I T Y F U N C T I O N C C C REMEMBER TO D E C L A R E I N T E G E R S M S N FOR FORMATTED OUTPUT c INTEGER I,J.K,L.M.N REAL S P S I ( 2 4 ) , N P S I ( 2 4 ) , D P S I ( 2 4 ) , U P T K ( 15 . 2 4 ) , R E S ( 1 5 , 2 4 ) , A ( 1 5 ) . B ( 1 5 ) X,C(15),ABAR,BBAR,CBAR.SUMA,SUMB.SUMC REAL D ABAR=0 BBAR=0 CBAR=0 SUMA'O SUM8=0 SUMC=0 D=-2.2  c c c c c c  c c  c c c c c c c c c  A B A R , B B A R , AND CBAR ARE AVERAGE C O E F F I C I E N T S SUMA, SUMB, SUMC A C C U M U L A T E C O E F F I C I E N T S FOR C A L C U L A T I N G READ(5,1) (A(K),B(K).C(K),K-=1.14) 1 F0RMAT(3F1O.5) " CALCULATING  AVERAGES  OF  COEFFICIENTS  DO 3 0 0 L= 1 . 14 SUMA=SUMA+A(L) SUMB=SUMB+B(L) SUMC=SUMC+C(L) 3 0 0 CONTINUE ABAR=SUMA/14 BBAR=SUMB/14 CBAR=SUMC/14 A(15)=ABAR B(15)=BBAR C(15)=CBAR P R I N T I N G AVERAGE C O E F F I C I E N T S AVERAGE C O E F F I C I E N T S A . B , S C PRINT, A ( 1 5 ) , B ( 1 5 ) , C ( 1 5 ) AVERAGE C O E F F E C I E N T S A . B . & C  A . B . SC  LOOP 2 0 0 I N D E X E S S E E D L I N G S 1 TO 14 LOOP 1 0 0 G E N E R A T E S S O I L WATER P O T E N T I A L V A L U E S DO 2 0 0 1 = 1 . 1 5 . 1 DO 10O J = 1 . 2 2 , 1 SPSK J)=J*(-0. 1 ) NPSI ( J ) = - 2 . 1 9 2 6 1 + ( 0 . 0 2 0 8 2 0 1 * S P S I ( J ) ) DPSI( J ) = S P S I ( J ) - N P S I ( J ) UPTK(I,J)=A(I)»<1-(EXP(-B(I)*(<SPSI(J)D)»* C(I))))) IF ( U P T K ( I , J ) . G T . 0 . 0 0 0 1 ) UPTK(I,J)=0 RES(I.J)=0 GO TO 3 3  GO TO 22  AVERAGES  5. . 0 0 0 6 000 7 000 8 OOO 9 OOO 10 . 0 0 0 1 .1 0 0.0 12.. 0 0 0 13. OOO 14 . 0 0 0 1 5 .. 0 0 0 16.. 0 0 0 17 . 0 0 0 18 . 0 0 0 19 ooo 20 .000 21 ooo 22 ooo 23 ooo 2 4 . ooo 2 5 .. 0 0 0 26 . 0 0 0 27 . 0 0 0 28 0 0 0 2 9 .. 0 0 0 30 .000 31 ooo 3 2 .. 0 0 0 3 3 . ooo 34 0 0 0 35. 000 36 . 0 0 0 37 . 0 0 0 38 ooo 39 . 0 0 0 40 .000 4 1 ooo 42 . 0 0 0 43 . 0 0 0 44 .ooo 45 .000 46 .000 47 . 0 0 0 48 . 0 0 0 4 9 .ooo 5 0 ooo 51 . 0 0 0 52 ooo 53 . 0 0 0 54 . 0 0 0 55 .000 56 ooo 57 . 0 0 0 58 . 0 0 0 59 . 0 0 0 60 .000 61 . 0 0 0 62 .ooo 6 3 .ooo  35 36 37 38 39  22 CONTINUE RES(I,d)=DPSI(J)/UPTK(I, d) 33 CONTINUE ' 100 CONTINUE 200 CONTINUE  c c c c PROGRAM PRINTS CALCULATED UPTAKE AND RESISTANCE FOR EACH SEEDLING c FOR 14 SEEDLINGS AND FOR AVERAGE: AVERAGE IS # 15 c OUTPUT UPTAKE AND RESISTANCE c FOR SEEDLINGS 1 TO 15: #15 REFERS TO AVERAGE COEFFICIENTS c c c c c OUTPUT: WRITING RES AND UPTK FOR 14 SEEDLINGS AND FOR AVERAGE AVERAGE IS # 15 ^ c WRITES RES AND UPTK FOR 3 SEEDLINGS (OR AVERAGE FOR #15) PER PAGE c c  DO 400 M=1,15,3 40 WRITE(6,405)M,M+1. M+2 41 EXTENSION'' OTHER COMPILERS MAY NOT ALLOW EXPRESSIONS IN OUTPUT LISTS • OTHER COMPILERS MAY NOT ALLOW EXPRESSIONS IN OUTPUT LISTS EXTENSION' 405 F0RMAT(2X, 'SEEDLING #' ,I 2.8X, 'SEEDLING #' ,I 2.8X, 'SEEDLING #'.I2) 42 ON FORMAT WRITE STATEMENT #405 C THANK YOU FOR THE EXTENSIONS C C  43 44  WRITE(6,415) 4 15 FORMAT(3X,'RES',7X,'UPTK',6X,'RES',7X,'UPTK',6X,'RES',7X.  „, . UPTK )  DO 500 N=1,22,1 WRITE(6,505)RES(M,N),UPTK(M,N),RES(M+1,N).uPTK(M+1,N),RES(M+2.N),U XPTK(M+2,N) 505 FORMAT(3(F10.4.F10.4)) 500 CONTINUE 49 50 51  400 CONTINUE STOP END  /EXECUTE 39E 01 O.1647898E 01 O. 0.8520613E 01 SEEDLING # 3 SEEDLING # 2 SEEDLING # 1 UPTK ES UPTK RES UPTK RES 8 4958 0. 2466 10 6302 0. 197 1 5 7983 0. 3613 8 3870 2381 0. 10 5584 0. 1891 5 7591 0. 3467 8 2654 2297 0. 4733 1813 10 7127 0. 5 0. 3324 8 1295 0. 22 15 10 3722 0. 1736 5 6576 0. 3183 7 9777 2135 0. 10 2525 0. 1661 5 5923 0. 3045 7 8080 2056 0. 10 1 109 1587' 0. 5 5151 0. 2910 7 6184 0. 1978 9 9437 0. 1516 5 4238 0 2779 7 4065 0. 1903 9 7464 0. 1446 5 3162 0. 2651 7 1698 1829 0 5140 9 0. 1378 5 1895 0 2527 6 9055 0. 1757 9 2409 0. 1313 5 0405 O 2407 6 6102 0 1688 8 9206 0. 1250 4 8658 O 2293  64 .000 65.000 66 .000 67 .000 68.000 69 . OOO 70.000 71.000 72.000 73.000 74 .000 75.000 76.000 77.000 78 .000 79.000 80.000 81.000 82 .000 83.000 84 . OOO 85.000  86.000 87.000 88.000 89.000 90.000 91.000 92.000 93.000 I 94.000 95.000 CO 96.000 ~-> 97.000 I 98 .000 99.000 100.000 '101 .000 102.000 103.000 104 .000 105.000  6 .2805 4 .6612 0 1620 0 1191 8 . 5456 0 2 183 0 2080 4 .4224 0 1 1 34 8 . 1077 0 1555 5 .9125 4 1443 7 . 5980 0 1 194 5 .5017 0 1082 0 1983 3 8 2 17 0 1033 7 .0064 0 1.135 5 0434 o 1894 3 4488 6 . 3227 0 138 1 4 .5321 0 0990 0 1815 3 0198 5 . 5363 0 1333 3 9620 o 1748 0 0954 4 .6374 0 1293 3 3268 2 5295 0 0927 0 1700 1 6 8 3 1 9 7 3 9 0 9 1 8 3 . 6 1 8 8 0 I2S8 2 6 194 0 0 1 3524 2 .4793 0 1278 1 8326 0 1732 0 0945 0 6739 0 1 103 1. 2356 0 142 1 0 9595 0 2023 O OOOO 0 OOOO 0 OOOO 0 OOOO 0 OOOO o OOOO SEEDLING # 4 SEEDLING # 5 SEEDLING # 6 RES UPTK RES UPTK RES UPTK 3 8655 0 2992 7 OOOO 0 1790 1 1 7047 0 54 19 3 8394 7 OOOO 0 17 11 1 1 6707 0 2853 0 5201 3 8085 0 2713 7 OOOO 0 1634 1 1 6220 0 4986 3 7717 7 OOOO 1 15 5 3 3 0 4775 0 2573 0 1559 O 4568 3 7282 0 2433 7 OOOO 0 I486 1 14 5 7 8 3 6767 0 2293 7 OOOO 0 14 17 1 13 2 7 0 0 4366 3 6 159 0 2 153 7 OOOO 0 1352 1 11509 O 4168 3 544 1 0 2013 6 9999 0 129 1 10 9 1 7 8 0 3976 3 4597 0 1874 6 9988 0 1235 10 6 1 4 5 0 3790 3 3603 0 1737 6 987 1 0 1 186 10 2 2 7 2 0 36 1 1 6 9179 0 1 145 9 7421 O 3439 3 2438 0 1612 3 1075 0 1527 6 6640 0 1113 9 1469 0 3275 2 9483 6 0454 0 1091 8 4325 0 1521 0 31 19 2 7629 4 9821 0 1082 7 5955 0 2974 0 1649 2 8 4 1 5 4 7 8 1998 3 6 2 3 6 0 1090 6 6405 2 0 0 2 2992 2 2774 0 112 1 5 5831 0 2722 0 2748 2 0132 0 4363 1 2101 0 1 186 4 4522 O 2623 1 6863 0 5236 0 1306 3 2931 0 2550 0 8215 1 3159 1 9530 0 1701 0 1532 2 1687 0 2524 o 2598 0 9016 6 9042 0 0339 0 2016 1 1619 64 1445 0 002 1 0 3577 0 381 1 o 3034 0 4493 0 OOOO 0 OOOO 0 OOOO 0 OOOO 0 OOOO 0 OOOO SEEDLING # 8 SEEDLING # 9 SEEDLING # 7 UPTK UPTK RES UPTK RES RES 9 7218 0 54 19 3 8655 , 0 2457 8 5266 0. 2155 9 667 1 0 5201 3 8394 b 2346 8 5096 O. 2066 9 599 1 0 4986 3 8085 0 2238 8 4833 0 . 1978 9 5 149 3 77 17 0 2 133 8 4436 O . 1893 0 4775 9 4 1 10 3 7282 0 203 1 8 3847 O. 1810 0 4568 3 6767 8 2995 9 2835 0 4366 0 1934 0 . 1729 9 1278 0 4 168 3 6159 0 1843 8 1790 0 . 1651 8 9384 3 544 1 0 1 759 8 0124 0 3976 0 . 1577 8 7094 0 3790 3 4597 0 1684 7 7880 O . 1506 0 16 19 7 4930 8 4340 0 361 1 3 3603 0 . 1439 8 1050 3 2438 0 . 1568 7 1 153 0 3439 0 . 1376 7 7 145 0 3275 3 1075 0 . 153 1 6 6451 0 . 1319 7 2546 2 9483 0 . 15 14 6 0765 O . 1268 0 3119 6 7 177 2 7629 0 15 19 5 4 101 0 2974 0 . 1223 6 0968 0 284 1 2 5478 0 . 1555 4 6553 0 . 1 187 5 3870 0 2722 2 2992 0 . 1633 3 8323 0 . 1 162 4. 5869 0 2623 2 0132 0 . 1776 2 9728 o . 115 1 3. 7005 1 6863 0 . 2028 2 12 10 o . 1 162 0 2550 2 . 7414 0 . 2524 1 3159 0 . 2495 1 3313 0 . 1212 0 6665 1 7. 3 9 9 0 . 2598 0 9016 0 . 3515 0 . 1346 0 . 698 1 0 1953 0 . 7597 0 . 3034 0 4493 0 . 1795 0 . OOOO 0 . OOOO 0 OOOO 0 . OOOO 0 OOOO 0 . OOOO S E E D L I N G #10 S E E D L I N G #11 S E E D L I N G #12  UPTK RES 0 . 2 178 9 .6165 0 . 2076 9 . 6 165 9 .6165 0 . 197S 0 . 1873 9 .6165 9 .6164 0 .1771 9 .6164 O . 1669 9 .6160 0 . 1S67 9 .6114 0 . 1466 0 . 1369 9 .5814 9 .4533 0 . 1284 0 . 1229 9 .0758 8 . 2700 0 . 1230 O . 1320 6 .9690 5 .3182 0 . 1545 3 .5184 0 . 2000 2 . 1630 0 . 2894 0 . 4749 1 .1119 0 . 4720 0..9112 2 . 1864 0 . 1519 7. . 7 5 6 0 0 .0302 72 . 0 9 7 3 0 .0019 0 . OOOO 0 .0000 S E E D L I N G #13 UPTK RES 9 .0000 0. 2327 0. 2219 9 .0000 9 .0000 0 . 21 10 0 . 2001 9.. 0 0 0 0 8 .9996 0 . 1892 8 . 9974 0 . 1784 8 . 9859 0 . 1677 8 . 9416 0 . 1576 8 . 8 100 0 . 1488 8 . 5008 0 . 1427 7 .9133 0 . 14 10 6. 9918 0 . 1455 5 . 7755 0 . 1592 4 . 4024 0 . 1867 3 . 0588 0 . 2366 0 . 3279 1 9091 . 1 0.4 6 9 0. 5043 8 8 7 1 0 . 4 848 0. 1 .8 9 6 2 0 . 1752 5. 7141 0. 0410 4 0 . 2503 0 . 0034 0 . OOOO 0 . OOOO CORE  USAGE  OBJECT  DIAGNOSTICS COMPILE  UPTK RES UPTK RES 7 .9985 12 -OOOO 0 . 2619 0 . 1 74S 0 . 2497 7 .9962 12 .OOOO 0 . 1664 7 .9909 0 . . 1582 12 .OOOO 0 . 2376 0 . 2257 7 .9796 0 . 1501 12 .OOOO 0 .2140 7 .9569 0 . 14 19 1 .19 9 9 9 7 .9137 0 . 1338 1 .1 9991 0 . 2028 7 .8367 0 . 1257 1 .1 9942 0 . 1923 7 .7075 0 . 1 177 1 .19 7 2 0 0 . 1828 0 . 1748 7 .5035 0 . 1 103 1 .18 9 4 2 7 .2002 0 . 1039 1 .16 7 8 7 0 . 1685 0 . 1646 6 .7753 0. 0996 1 .1 1 9 8 8 0 . 1637 6 .2150 0. 0986 10 . 3 2 3 0 0 . 1666 5 .5192 0 . 1023 a .9918 4 .7071 0 . 1 129 7 .2803 0 . 1746 0 . 1896 3 .8173 0 . 1342 5 .3952 2 .9058 0 . 1739 3 .5995 0..2154 0 . .2591 2 .0378 0 . 2497 2 . 1 145 0. 3367 1 . 2775 0 . 408 1 1 .0538 0 .6773 0. 8019 0 .4142 0. 4904 0. 8734 0 .2682 2 . 1658 0 . 1082 2. 5 5 8 3 0 .0533 12 . 7444 0 .0107 0 . OOOO 0 OOOO 0 . OOOO 0 .0000 S E E D L I N G #14 S E E D L I N G #15 RES UPTK RES UPTK 8 .9001 0. 2460 8 .5165 0. 2354 0 . 2253 8 .8633 0. 2346 8 .5121 8 .8143 0 . 2233 8 . 5035 0 . 2154 0 . 2058 8 . 7497 0 . 2122 8 .4876 8. . 6 6 5 2 0 . 20 13 8. . 4 5 9 3 0 . 1965 8 . 5559 0 . 1908 8. .4 109 0 . 1876 8 . 4157 0 . 1809 8. . 3 3 1 8 0 . 1791 8. 2 3 7 9 0 . 17 17 8. 2 0 7 3 0 . 17 11 8 .0150 0 . 1635 8. . 0 1 9 8 0 . 1636 7. 7 3 8 6 7 7492 0 . 1568 0 . 1566 7 . 4003 0 . 15 13 7. 3 7 5 1 0 . 1507 6. 9918 6. 8 8 0 8 0 . 1479 0 . 1455 6 . 5059 0. M 7 0 6. 2568 0 . 1414 5. 937 1 5. 5 0 6 2 0 . 1492 0 . 1384 5 . 2835 0 . I557 4 .6478 0 . 1370 0 . 1376 4 . 5482 0 . 1683 3. 7 1 8 3 3. 7413 2 . 77 1 1 0 . 14 1 1 0 . 1905 0 . 1492 2. 8828 0 . 2298 1 8. 7 1 8 2. 0059 10.9 0 0 0 . 1656 0 . 3047 1 1626 . 0. 4782 0. 4899 0 . 2015 0 . 4351 1. 1400 • 0 . 1196 0 . 3134 0 . OOOO 0 . OOOO 0 . OOOO 0 . OOOO  TIME'  CODE=  NUMBER 0.068  OF  2288  BYTES.ARRAY  ERRORS'  SEC.EXECUTION  O. TIME'  AREA'  NUMBER OF 0.106  /STOP Execution  SSIGNOFF  terminated  15:37:37  T=0.19  RC'1  $0.19  SEC.  3348  BYTES,TOTAL  WARNINGS' WATFIV  O. -  JUL  AREA  AVAILABLE'  NUMBER OF  1973 V 1 L 4  102400  EXTENSIONS' 15:37:36  BYTES 2  THURSDAY  - 100 -  APPENDIX IV Observed Uptake Rates i n Ponderosa Pine Compared to Average Uptake Curves  - 101 -  1  -3.0  =  z  1  1  -2.0  1  1  -1.0  Soil Water Potential (MPa)  Figure IV A OBSERVED UPTAKE IN PONDEROSA PINE PER UNIT ROOT AREA vs SOIL WATER POTENTIAL (Uptake Curve for Regression Model I is Shown)  r  0  - 102 -  Mac-  Figure IV B OBSERVED UPTAKE IN PONDEROSA PINE vs SOIL WATER POTENTIAL . (Uptake Curve for Regression Model II is Shown)  - 103 -  APPENDIX V Uptake and Needle Water P o t e n t i a l Data f o r Ponderosa P i n e  - 104 -  UPTAKE DATA  Seedling Number  fs(MPa)  U (mg s"1 m-2)  Seedling Number  ¥ s(MPa)  U (mg s 1 rn" ) -  1  -0.38 -0.43 -0.48 -0.49 -0.52 -0.68 -0.96 -2.06  3.933 3.865 2.906 5.782 1.653 1.482 2.471 1.455  6  -0.06 -0.30 -0.43 -0.67 -0.81 -1.57 -1.93 -2.10  2  -0.80 -0.90 -0.91 -0.96 -2.81 -3.23  6.254 3.809 7.659 9.676 2.526 2.355  7  3  -0.40 -0.43 -0.48 -0.50 -1.94  8.623 8.210 7.591 7.522 2.026  -0.64 -0.71 -0.76 -0.81 -0.94 -1.12 -1.85 -2.04 -3.06  9.557 8.836 9.137 8.949 9.095 7.392 2.493 2.371 0.07567  8  -0.82 -0.94 -0.99 -1.05 -1.38 -1.91 -2.08 -2.21 -3.09  10.43 10.13 9.513 9.732 4.849 1.178 0.8754 1.037 0.5946  -0.46 -0.52 -0.67 -0.88 -1.12 -1.84 -2.26 -1.89 -2.39  4.460 4.013 1.726 1.364 1.508 1.455 1.248 1.143 0.5685  9  -0.31 -0.34 -0.42 -0.41 -0.44 -0.57 -1.67 -2.11 -3.13  4.015 4.172 3.992 4.009 3.818 1.716 1.685 1.584 0.1641  -0.40 -0.51 -0.61 -0.66 -0.79 -1.70  9.393 9.940 8.437 7.889 5.694 2.701  10  -0.77 -1.02 -1.04 -1.03 -1.46 -2.13 -2.99  8.838 7.462 9.820 11.78 3.936 1.274 1.175  4  5  11.96 11.13 12.03 11.78 10.52 4.766 2.244 2.167  2  - 105 UPTAKE DATA  Seedling Number  ^s(MPa)  U (mg s 1 m~2) _  11  -0.01 -0.94 -1.16 -1.04 -1.28 -1.28 -1.53 -1.69 -2.00  9.008 5.968 7.860 9.211 3.333 1.847 3.067 3.536 1.514  12  -0.01 -0.26 -0.38 -0.72 -2.39  11.78 10.22 13.98 13.58 3.137  13  -0.75 -0.98 -1.14 -1.08 -1.35 -1.73 -2.37 -2.45 -3.04  8.739 7.626 7.950 7.745 3.838 1.483 1.328 0.1826 0.09314  14  -0.38 -0.46 -0.49 -0.86 -1.24 -1.87 -2.98  9.638 8.799 8.053 7.491 3.143 2.375 1.083  106 -  NEEDLE WATER POTENTIAL DATA  *S(MPa)  YN(MPa)  -0.54  *S(MPa)  l'N(MPa)  *S(MPa)  ^(MPa)  -2.57  -0.08  -1.34  -2.32  -.1.70  -0.19  -2.26  -1.91  -2.17  -2.08  -2.13  -0.75  -2.59  -0.77  -2.36  -2.78  -2.21  -1.16  -2.26  -0.01  -2.12  -1.69  -2.64  -1.12  -2.50  -0.61  -2.06  -2.43  -2.21  -0.40  -2.23  -1.36  -2.16  -1.86  -2.55  -0.46  -2.32  -1.24  -2.09  -1.71  -2.00  -0.47  -2.40  -0.57  -2.68  -1.96  -2.14  -0.01  -2.14  -0.61  -2.50  -1.41  -1.97  -0.26  -2.45  -1.14  -2.41  -2.25  -2.01  -0.50  -2.65  -0.95  -2.26  -1.67  -2.31  -0.63  -2.89  -1.03  -1.85  -2.42  -2.00  -0.01  -2.02  -4.22  -1.92  -3.24  -2.51  -0.21  -1.99  -4.67  -2.78  -2.91  -1.94  -0.04  -1.63  -2.09  -2.21  -3.71  -2.22  -0.15  -1.73  -3.47  -2.35  - 107 -  APPENDIX VI Data f o r Douglas-Fir Experiment  - 108 -  CONTROL TREATMENT  Resistance Seedlino Number  ¥ s (MPa)  ¥ N (MPa)  Seedling (TPa s Kg-1)  Root Area Needle Area (TPa s m Kg" ) (TPa s m Kg" ) 2  1  2  1  -0.24 -0.48 -0.74 -1.14 -2.06 -2.55  -2.29 -2.82 -2.10 -2.14 -2.98 -3.12  10.4699 12.6350 11.0749 16.1996 15.0548 16.8739  0.1669 0.2014 0.1766 0.2583 0.2400 0.2690  0.09104 0.02291 0.09630 0.1409 0.1309 0.1467  2  -0.78 -0.49 -0.58 -0.57 -0.70 -1.01  -2.95 -2.86 -3.20 -2.61 -2.70 -3.14  13.2317 14.9338 18.4507 14.3662 28.6164 26.5553  0.3178 0.3587 0.4431 0.3450 0.6873 0.6378  0.1331 0.1503 0.1856 0.1446 0.2879 0.2672  3  -0.09 -0.80 -0.91 -1.37 -2.07 -2.35  -2.24 -2.86 -2.96 -3.11 -3.12 -3.17  9.0256 11.1231 11.6876 25.6259 31.4937 29.7857  0.2305 0.2840 0.2985 0.6544 0.8042 0.7606  0.1184 0.1459 0.1533 0.3362 0.4131 0.3907  4  -0.09 -0.29 -0.88 -0.93 -2.43 -2.26  -2.68 -2.33 -2.94 -3.11 -3.10 -3.28  11.6382 12.0496 14.0903 16.0530 19.5237 25.8364  0.3347 0.3465 0.4502 0.4615 0.5614 0.7430  0.1505 0.1558 0.1822 0.2075 0.2524 0.3340  5  -0.07 -0.30 -0.43 -0.83 -2.62 -2.78  -2.81 -3.04 -2.87 -2.69 -3.11 -3.35  14.3832 19.9128 16.6553 17.1113 17.6386 17.4365  0.4153 0.5750 0.4809 0.4941 0.5093 0.5035  0.1552 0.2149 0.1797 0.1847 0.1904 0.1882  1  - 109 PLANTED TREATMENT  Seedling Number  Resistance s (MPa) V  * N (MPa)  Seedling (TPa s Kg-1)  Root Area Needle Area (TPa s m2 Kg-1) (TPa s m2 K g " ) 1  1  -0.186 -0.48 -0.48 -0.48 -0.51 -0.82  -2.89 -2.76 -3.37 -3.25 -2.52 -2.22  102.2113 98.4592 104.0403 98.7232 103.3425 131.7647  2.832 2.728 2.883 2.735 2.863 3.651  0.8966 0.8637 0.9126 0.8265 0.8652 1.103  2  -0.079 -0.54 -0.51 -0.38 -0.47 -0.58  -2.42 -2.60 -2.68 -2.94 -2.68 -2.34  110.6123 83.4302 39.0600 67.8978 118.7463 117.5355  2.343 1.762 0.8250 1.434 2.508 2.482  1.867 1.408 0.6593 1.146 2.004 1.984  3  -0.30 -0.72 -0.95 -0.91 -1.01 -0.99  -2.37 -2.68 -2.34 -2.53 -2.06 -2.22  77.9804 67.8905 57.9167 56.9658 62.3697 56.0722  1.963 1.709 1.458 1.434 1.570 1.411  0.9326 0.8120 0.6927 0.6813 0.7459 0.6706  4  -0.094 -0.49 -0.48 -0.59 -0.66 -0.66  -2.67 -2.61 -2.22 -2.46 -2.59 -3.00  54.0960 49.0629 47.4807 43.9750 46.9984 51.2270  1.911 1.734 1.678 1.554 1.661 1.810  0.5232 0.4745 0.4592 0.4253 0.4546 0.4955  5  -0.58 -0.23 -0.25 -0.30 -0.37 -0.37  -2.27 -2.23 -2.59 -2.14 -2.27 -2.22  41.0296 48.4919 41.9358 37.2600 40.3943 56.0076  1.476 1.745 1.509 1.341 1.453 2.015  0.5608 0.6628 0.5732 0.5093 0.5521 0.7656  - 110 PLANTED AND VIBRATED TREATMENT  Resistance Sf*f*H 1 i nn  V> t» V_y l _ i J  L 1 |U  Number  (MPa)  ¥N (MPa)  Seedling (TPa s Kg" ) 1  Needle Area Root Area (TPa s m Kg" ) (TPa s m Kg" ) 2  1  2  1  -0.185 -0.43 -1.36 -2.00 -2.85 -2.93  -2.66 -2.17 -2.72 -2.86 -3.11 -3.22  4.9239 14.0940 12.2400 12.2857 14.9614 17.0779  0.1400 0.4007 0.3480 0.3493 0.4254 0.4856  0.06730 0.1926 0.1673 0.1679 0.2045 0.2334  2  -0.30 -0.25 -0.31 -0.39 -0.48 -0.99  -3.12 -2.86 -2.66 -2.70 -2.93 -2.72  7.1064 11.6547 14.1000 11.6962 13.2935 14.8286  0.1769 0.2934 0.3554 0.2948 0.3351 0.3738  0.08505 0.1413 0.1709 0.1418 0.1611 0.1797  3  -0.079 -0.41 -0.51 -0.64 -1.56 -2.36  -2.95 -2.97 -3.20 -2.60 -2.55 -2.47  6.9542 15.3600 16.4136 10.9490 14.8500 15.9141  0.1516 0.3349 0.3579 0.2387 0.3238 0.3470  0.07294 0.1611 0.1722 0.1148 0.1558 0.1669  4  -0.20 -0.62 -0.67 -0.75 -1.36 -1.84  -2.37 -2.59 -2.44 -2.60 -2.45 -2.70  6.6150 10.6380 09.9562 13.6227 12.7985 12.7323  0.2076 0.3339 0.3125 0.4276 0.4018 0.3997  0.06229 0.1002 0.09375 0.1283 0.1205 0.1199  5  -0.11 -0.55 -0.49 -0.58 -0.51 -2.35  -2.76 -3.13 -2.93 -3.06 -2.12 -2.14  5.5038 10.7169 12.5486 12.5550 18.1125 19.8427  0.2396 0.4665 0.5462 0.5465 0.7884 0.8637  0.1069 0.2083 0.2438 0.2439 0.3519 0.3855  1  - 111 -  MEANS AND STANDARD DEVIATIONS (S.D.)  Control  Planted  Planted and Vibrated  T s (MPa)  Mean  -1.095  -0.516  -0.936  S.D.  0.859  0.249  0.837  17.318  70.433  12.478  R(TPa s Kg" ) (per seedling) 1  Mean  6.1290  S.D.  28.652  3.6360  R(TPa s m Kg" ) (per root area) 2  1  Mean  0.43365  1.9493  0.37554  S.D.  0.18210  0.62642  0.15889  Mean  0.19175  0.85104  0.16406  S.D.  0.088282  0.43669  0.074710  R(TPa s m Kg" ) (per needle area) 2  1  -  112  -  APPENDIX V I I S t a t i s t i c a l Summaries f o r R e s i s t a n c e in Douglas-Fir Seedlings  - 113 -  COVARIANCE ANALYSIS PER SEEDLING  ANOVA D.F.  Source  Treatment  Within  Treatment  Error  Total  SS  MS  F  Probability  73.7506  0.0000  4.9085  0.0000  2  9.554574  4.777287  12  7.77315 x 10-1  6.47762 x IO"  74  89  9.765572 x 10-1  2  1.319672 x IO" 2  11.371552  Covariate  Regression Coefficient  PSI  -0.10577  Standard Error  F Value  Probability  0.01887  31.4127  0.0000  - 114 -  COVARIANCE ANALYSIS PER UNIT ROOT AREA  ANOVA D.F.  Source  Treatment  Within  Treatment  Error  Total  SS  MS  F  Probability  2  9.265220  4.632609  28.6792  0.0000  12  1.938382  1.615318 x 10-1  12.2485  0.0000  74  9.759058 x 10-1  1.318792 x IO"  89  2  12.225852  Covariate  Regression Coefficient  Standard Error  F Value  Probability  PSI  -0.10581  0.01886  31.4835  0.0000  - 115 -  COVARIANCE ANALYSIS PER NEEDLE AREA  ANOVA  Treatment  Within  SS  D.F.  Source  Treatment  MS  F  Probability  2  11.731708  5.865853  50.7649  0.0000  12  1.386592  1.155493 x 10"  8.7554  0.0000  1  Error  74  9.76618 x 10" 1  Total  89  1.31975 x IO" 2  14.176537  Covariate  Regression Coefficient  Standard Error  F  PSI  -0.10589  0.01887  31.4793  Value  Probability  0.0000  - 116 -  APPENDIX V I I I Stomata! R e s i s t a n c e Measurements and E s t i m a t e d Maximum T r a n s p i r a t i o n Rates  - 117 -  STOMATAL RESISTANCE  Stomatal R e s i s t a n c e ( s / c m )  Species  d  Ponderosa P i n e  2.91  Douglas-Fir  2.28  Douglas-Fir  2.78  Douglas-Fir  2.64  Douglas-Fir  2.87  d  Measured i n the growth chamber with a v e n t i l a t e d d i f f u s i o n porometer; s o i l water content was s l i g h t l y below f i e l d c a p a c i t y f o r a l l s e e d l i n g s .  An e s t i m a t e of the maximum t r a n s p i r a t i o n r a t e f o r s e e d l i n g s i n t h e growth chamber can be made u s i n g t h e r e l a t i o n s h i p : l T = —  " a  l  + a  p  r  where T i s the t r a n s p i r a t i o n d e n s i t y i n the l e a f  p  r  r a t e i n mg  (mg c m ) , P - 3  a  c m , Pj_ i s the water vapour -2  i s the water vapour d e n s i t y i n the a i r  (mg c m ) , r ^ i s l e a f r e s i s t a n c e (s c m ) , and r - 3  resistance (at  - 1  (s cm""1).  a  i s boundary l a y e r  Assuming t h a t l e a f and a i r temperatures a r e equal  20°C) and boundary l a y e r r e s i s t a n c e i s low ( a p p r o x i m a t e l y 0.5  s cm 1) i n the w e l l v e n t i l a t e d  growth chamber,  _  6.058 x I O " mg 3  cm  then f o r ponderosa p i n e :  - 3  T =  = 1.7 x 1 0 " mg s ~ 3  3.5 s c m  1  cm"  2  -1  and f o r D o u g l a s - f i r ( u s i n g the average of the four measurements o f stomatal r e s i s t a n c e ) : 6.058 x 1 0 ~ mg cm" 3  T =  3  = 1.9 x 1 0 3.14 s c m  -1  - 3  mg s ~  1  cm  - 2  - 118 -  The average maximum observed t r a n s p i r a t i o n about 1.6 x 1 0 3 g - 1 _  m  s  c m  -2.  -  - 2  f o r ponderosa pine was  The maximum p r e d i c t e d  f o r uptake Models I and I I a r e 5 x 1 0 mg s 1 c m  rate  - 3  transpiration  mg s"1 c m  - 2  and 2 x 1 0  - 3  respectively.  Maximum observed r a t e s  for Douglas-fir  seedlings  (at s o i l  potentials  g r e a t e r than -0.1 MPa) a r e about 1.8 x 1 0  1.8 x 1 0  mg  - 3  rates  respectively.  cm  - 2  f o r control  - 3  treatment s e e d l i n g s  water  , 1.7 x 1 0 3, 4 and 5  - 3  and  APPENDIX IX S e e d l i n g and Water Pathway Dimensions  I  - 120 -  PONDEROSA PINE INDIVIDUAL SEEDLING DIMENSIONS  0D Root: Shoot R a t i o  Root Area (cm2)  Needle Area (cm2)  Needle Area: Root Area R a t i o  1  146  36  0.25  2.35  2  164  53  0.33  1.95  3  145  61  0.42  1.60  4  84  40  0.48  2.11  5  125  35  0.28  2.31  6  97  48  0.49  1.14  7  121  42  0.35  2.01  8  132  36  0.27  2.37  9  73  36  0.50  1.17  10  102  50  0.50  2.21  11  89  31  0.35  1.79  12  128  65  0.51  1.16  13  94  46  0.49  2.13  14  120  48  0.40  1.57  Seedling Number  - 121 -  DOUGLAS-FIR INDIVIDUAL SEEDLING DIMENSIONS  Treatment Control  Planted  P l a n t e d and Vibrated  S e e d l i n g Root Area Needle Area Needle Area: 0D Root: Number (cm ) (cm ) Root Area R a t i o Shoot R a t i o 2  2  1  159  87  0.54  0.40  2  240  101  0.42  0.71  3  255  131  0.51  --  4  288  129  0.45  --  5  289  108  0.37  0.69  1  277  84  0.30  0.81  2  211  107  0.51  0.39  3  252  120  0.48  --  4  353  97  0.27  --  5  360  98  0.27  0.66  1  284  137  0.48  2  252  121  0.48  0.28  3  218  105  0.48  --  314  94  0.30  1.02  435  194  0.45  0.40  5  - 122 -  AVERAGE  7b  S o i l Volume (cm3)  T o t a l Root Length (cm)  (cm)  (cm)  360  1210  0.19  0.0132  Douglas-Fir Control  370  2198  0.14  0.0141  Douglas-Fir Planted  370  2608  0.12  0.0144  Douglas-Fir P l a n t e d and Vibrated  370  2411  0.14  0.0158  Treatment  Ponderosa P i n e  a  WATERFLOW PATHWAY DIMENSIONS  Z i s t h e c a l c u l a t e d average d i s t a n c e t h a t water must move through t h e s o i l to a root surface  &r i s the c a l c u l a t e d weighted average r o o t  radius  -  123  -  APPENDIX X Soil Physical  Properties  - 124 -  SOIL PHYSICAL  %  Treatment  Sand*  %  SiLt  a  %  PROPERTIES  Clay  d  Average Bulk D e n s i t y (Kg m ) K b(cm -3  s  Control  79  17  4  995  0.96  Planted  79  17  4  1020  1.00  P l a n t e d and Vibrated  79  17  4  984  0.95  a  B a s e d on 2 mm  D  Saturated  and s m a l l e r f r a c t i o n ,  hydrometer method  h y d r a u l i c c o n d u c t i v i t y , c o n s t a n t head permeameter  method  d" ) 1  — I  1  -1.0 -0.1 Soil Water Potential (MPa)  Figure  -0.01  X A  SOIL WATER RETENTION CURVE FOR DOUGLAS FIR (ALL TREATMENTS) (Determined by Pressure Plate Extraction Method)  - 126 -  APPENDIX XI E s t i m a t e s of S o i l  Resistance  - 127  -  DOUGLAS-FIR CONTROL SEEDLINGS  MMPa)  a  6(m3  m  Kb (Kg m-1  -3)a  R c Pa"  s" )  1  1  (TPa  s Kg" )  Rs/RT  2.9  x 10-*  --  1  -0.53  .06  6.25  x  10"  -0.81  .03  4.25  x  10"  1 2  4.3  x 10-3  --  -1.5  .015  3.27  x  10" 3  5.5  x 10-2  .006  V o i u m e t r i c water content e s t i m a t e d  11  1  from water r e t e n t i o n  curve  bUnsaturated h y d r a u l i c c o n d u c t i v i t y e s t i m a t e d by the method shown by dackson (1972); c o n v e r t e d from cm d a y t o p r e s s u r e u n i t s by: - 1  1 cm Calculated  soil  day"  1  = 1.18  x 10~  8  r e s i s t a n c e a f t e r Gardner  Kg n r  1  Pa"  1  s  _ 1  (1960)  d R a t i o of c a l c u l a t e d s o i l r e s i s t a n c e to minimum observed r e s i s t a n c e (shown a t = -1.5 MPa o n l y )  total  d  - 128 -  DOUGLAS-FIR PLANTED SEEDLINGS  Ys(MPa)  a  e(m  3  nr ) 3  Kb (Kg n r  d  1  Pa"  1  s" ) 1  R c (TPa s Kg-1)  3.5 x  -4  -0.53  .06  6.73 x 10-1  -0.81  .03  4.48 x 1 0 "  1 2  5.2 x 10"  3  -1.5  .015  3.47 x 1 0 "  1 3  6.8 x I O "  2  1  1 0  Rs/RT  V o l u m e t r i c water content e s t i m a t e d from water r e t e n t i o n  -.002  curve  bUnsaturated h y d r a u l i c c o n d u c t i v i t y e s t i m a t e d by the method shown by Jackson (1972); c o n v e r t e d from cm day"1 t o p r e s s u r e u n i t s by: 1 cm d a y " Calculated  soil  1  = 1.18 x 1 0 "  8  r e s i s t a n c e a f t e r Gardner  Kg n r  1  Pa" s " 1  1  (1960)  ^ R a t i o o f c a l c u l a t e d s o i l r e s i s t a n c e to minimum observed r e s i s t a n c e (shown a t ^ s = -1.5 MPa o n l y )  total  d  - 129 -  DOUGLAS-FIR PLANTED AND VIBRATED SEEDLINGS  Vs(MPa)  a  6 (m  3  nT ) 3  Kb (Kg n r  a  1  Pa"  s" )  1  1  R C (TPa s Kg" ") -  -4  -0.53  .06  5.00 x 1 0 "  1 1  3.5 x  -0.81  .03  3.36 x 1 0 "  1 2  5.3 x 1 0 "  3  -1.5  .015  2.55 x 1 0 "  1 3  6.9 x 1 0 -  2  10  V o l u m e t r i c water c o n t e n t e s t i m a t e d from water r e t e n t i o n  Rs/RT  —  —  .014  curve  bUnsaturated h y d r a u l i c c o n d u c t i v i t y e s t i m a t e d by t h e method shown by Oackson (1972); c o n v e r t e d from cm d a y t o p r e s s u r e u n i t s by: 1  1 cm d a y  1  = 1.18 x 1 0 " Kg n r 8  1  Pa" s" 1  1  C a l c u l a t e d s o i l r e s i s t a n c e a f t e r Gardner (1960) d R a t i o o f c a l c u l a t e d s o i l r e s i s t a n c e t o minimum observed r e s i s t a n c e (shown a t ^ s = -1.5 MPa o n l y )  total  d  

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