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The absorption of inorganic nutrients by plants treated with sulphur dioxide Jack, Wilfred Robert 1937-12-31

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THE ABSORPTION OF IHORGAMIC MDTRIEMIS BY PLANTS TREATED WITH SULPHDR DIOXIDE  W i l f r e d Robert Jack  A T h e s i s submitted f o r the Degree o f MASTER OF ARTS i n the Department o f BOTANY  The U n i v e r s i t y o f B r i t i s h April,  1937  Columbia  THE ABSORPTION OF INORGANIC NUTRIENTS BY PLANTS TREATED WITH SULPHUR DIOXIDE.  I  The Problem Defined.  II  The Environmental Factors and t h e Plant Stock.  III  The Experimental  IV  The Results of the Experiments.  V  Discussion.  VI  Acknowledgements.  VII  Bibliography.  Conditions.  The effect upon vegetation of the gaseous wastes from i n d u s t r i a l enterprises, p a r t i c u l a r l y smelters, has received considerable a t t e n t i o n i n t h i s century both from the l e g a l and from the s c i e n t i f i c points of view. I t was soon agreed that t h i s effect was  caused by  the presence of gaseous sulphur dioxide i n the atmosphere. For a summary of investigations upon t h i s subject p r i o r to 1915 the reader i s referred to the 528 pages published by the Selby Smelter Commission ( l ) . Such s c i e n t i f i c investigations have shown t h a t the conditions under which i n j u r y to vegetation from sulphur dioxide takes place are determined p r i m a r i l y by the concent r a t i o n of the gas, the duration of the exposure, and the humidity of the a i r . Secondary factors are the c h a r a c t e r i s t i c s u s c e p t i b i l i t y of the species, the l i g h t conditions, and other environmental conditions of atmosphere and s o i l .  On  these conditions a large amount of data has been compiled. It i s not the purpose of t h i s paper to disprove any of these findings.  However, i t has been the purpose of these experiments to determine the absorption of inorganic nutrients by treated plants, and. to see what conclusions might be made regarding the metabolism of the plants from t h i s point of view.  The plant environment i s so complex, with so many variable factors entering that i t would be impossible to get data on a l l combinations i n a problem of t h i s kind. The method followed has been the usual r e l a t i v e study employing controls.  In a l l factors excepting sulphur dioxide i n the  atmosphere,' the control plants-were developing i n an environment i d e n t i c a l with that of the experimental plants . Buring treatments with the gas, a i r was supplied i n continuous flow to the plants i n s p e c i a l l y constructed a i r - t i g h t cabinets by blowers ("Sirocco" blowers, driven by General E l e c t r i c motors, 1/20 H.P., 110 V o l t s , 1.3 Amps., 1140\R'.P.M.). The a i r flow delivered by such blowers may be conveniently regulated by adjusting a s l i d i n g panel over the aperature i n a galvanized i r o n box connected t o t h e blower intake.  To determine the volume of the a i r flow and to main-  t a i n equality between control and fumigated cabinets a c a l i b rated anemometer (Short & Mason, Ltd., London,  "Biram's !b.  3132") was introduced into the a i r supply pipe through an a i r - t i g h t s l i d i n g panel.  The l i n e a r v e l o c i t y was then deter-  mined.for an i n t e r v a l of time measured by a stopwatch.  This  l i n e a r T e l o c i t y m u l t i p l i e d by the c r o s s - s e c t i o n of the supply pipe gave the volume of a i r s u p p l i e d . The a i r f u r n i s h e d t o the experimental  cabinet was  passed through a "mixing chamber", a l a r g e g a l v a n i z e d box w i t h b a f f l e p l a t e s i n i t .  iron  A tube l e a d i n g to the a i r e n t r y  end of t h i s chamber c a r r i e d the s u l p h u r d i o x i d e gas f o r i n t i m a t e mixture  i n the c o n c e n t r a t i o n s d e s i r e d .  The sulphur d i o x i d e f o r the treatments was i n s m a l l c y l i n d e r s from the manufacturers,  obtained  A n s u l Chemical  Company, M a r i n e t t e , W i s c o n s i n , through the C e n t r a l S c i e n t i f i c Company.  These 6 l b . c y l i n d e r s c o n t a i n l i q u i d sulphur d i o x i d e  of extreme p u r i t y , no t r a c e of sulphur t r i o x i d e heing f o u n d . They are equipped w i t h a needle v a l v e f o r r e g u l a t i n g t h e f l o w of gas from the c y l i n d e r s . To measure the f l o w of gas through a needle v a l v e , a meter of t h e type d e s c r i b e d by Benton (2) may be used. This a r t i c l e d i s c u s s e s the t h e o r e t i c a l 'and p r a c t i c a l .consid- • e r a t i o n s which l e a d t o the a p p l i c a t i o n of P o i s e u i l l e ' s (for  formula  the f l o w o f l i q u i d s ) to t h e f l o w of gases through cap-  i l l a r y t u b i n g , and c a l c u l a t i o n of volume from the pressure difference.  I n t h i s way a U-ma,nometer of the i n c l i n e d gauge  type w i t h a s e a l e d - i n c a p i l l a r y tube between the arms has been used. for  The manometer was f i l l e d w i t h l i q u i d  petrolatum.,  low c o n c e n t r a t i o n s i t has heen found more convenient  to  use c a p i l l a r y t u b i n g l e a d i n g from the c y l i n d e r o u t l e t i n t o , a r e s e r v o i r of p e t r o l a t u m w i t h t h e the l i q u i d .  o u t l e t above the s u r f a c e of  Counts of gas-bubble r a t e s of f l o w lip through  6.  the petrolatum correlated with concentration determinations gave a useful empirical method of adjusting the concentration to the d e s i r e d amount. A constant pressure from the cylinder was maintained by temperature control.. A l i n e d gas-metering box was constructed.  I t contained the c y l i n d e r almost Immersed i n a  water-bath; the c a p i l l a r y U-manometer; and a thermo-regulator (Geneo De Khotinsky) governing a 100  Watt  lamp connected to  supply heat when needed. To ensure p u r i t y of ;air supply and to guard against any possible accident, the gas-meter box was kept i n an adj o i n i n g room and the outlet passed through the w a l l t o the mixing box. The frequent and accurate determination of the sulphur dioxide content i n the atmosphere during any treatment is.essential.  The most r e l i a b l e method yet devised i s the  sampling system, of Thomas and Abersold, (3).." In t h i s method the gas i s oxidized i n absorbers by d i l u t e hydrogen peroxide and s u l p h u r i c a c i d s o l u t i o n , and the changing conductivity i s measured upon a Leeds and Hbrthrup recording Wheat stone bridge.  The s p e c i a l apparatus f o r t h i s method was not a v a i l -  able f o r these experiments.  But determinations of low con-  centrations of sulphur dioxide- using the methiod  described by  G r i f f i n and Skinner ( 4 ) are accurate * and the necessary apparatus i s e a s i l y assembled.  This l a t t e r method was used  throughout. The apparatus -consisted of two absorbers, ah a i r  flowmeter of the U-manoineter type, a suction pump, a.nd small pressure-equalizing tank.  The a i r flowmeter was calibrated  by means of a wet meter which i n turn had been checked against a standard "prover". The manometer readings were determined for a wide range on either side of the optimum flow, which i s about 1 gram-molecular volume of gas i n 3 minutes.  The  arrangement f o r c a l i b r a t i n g the flow rate i s i l l u s t r a t e d i n the accompanying photograph.  The absorbers contain 100 c c . of an iodinepotassium iodide-starch solution about 0.00003 N i n iodine. Through one absorber containing t h i s solution a metered volume of a i r i s drawn as a "blank" using a soda-lime tower.  Through  the other absorber the same volume of a i r i s drawn from the  8. experimental flasks  and t i t r a t e d to  a standard The  cabinet.  The  s o l u t i o n s are  the  same l i g h t - b l u e  sodium t h i o s u l p h a t e  titration  i s made  w h i c h may b e r e a d t o  by t h e 0.02  two t i t r a t i o n s r e p r e s e n t s dioxide be  absorbed.  sulphur  solution dioxide  arrangement  is  in  of  a small precise  equivalent  of  1 c . c . of to  1 gram-molecular  photograph.  for  volume  0.0015 l i . buret  between  the  sulphur  sodium  per m i l l i o n of  the  p e r m i l l i o n may  0 . 0 0 2 l\  1 part  into with  0.001C H to  The d i f f e r e n c e  equivalent  of t h e a p p a r a t u s  accompanying  use  end-point  concentration in parts  calculated directly, since  sulphate  the  The  solution,  c.c. the  then drained  air.  a determination  is  thioof  The shown  in  '-. As has been mentioned, the other atmospheric condition of primary importance in;gas treatments i s the humidity. This i s a very d i f f i c u l t f a c t o r to c o n t r o l .  In a l l treatment  a record of the r e l a t i v e humidity has been kept. Temperature of the atmosphere has a l s o been recorded. The l i g h t f a c t o r was v a r i a b l e during two experiments The plants i n t hese experiments were grown i n tthe greenhouse, under the s o l a r i l l u m i n a t i o n received by t h i s part of the P a c i f i c coast i n the late f a l l . Tight from day to day:  There i s a wide v a r i a t i o n of  c l e a r bright sunlight, d i f f u s e l i g h t  on foggy days, or very l i t t l e l i g h t i n cloudy weather. other experiments were c a r r i e d out under " a r t i f i c i a l " i n a t i o n i n s p e c i a l l y constructed cabinets.  The illum-  Excepting f o r  the gradual decrease i n i n t e n s i t y of the r a d i a t i o n from the bulbs, the l i g h t .factor was s i m i l a r f o r each 24-hour period. The construction of the cabinets i n which the l i g h t factor was c o n t r o l l e d followed the general plan of Davis and Hoagland ( 5 ) .  One of the modifications adopted by Swain and  Johnson (6) was introduced, however: theolights.were placed d i r e c t l y over the plants. Each cabinet followed the same pattern:  a counterpoised l i f t piece (6 f t . long x 2 f t . wide  x 5 f t . high), rested upon a f i x e d base ( 6 f t . long x 2 f t . widejx 2 f t . h i g h ) .  The e n t i r e construction was  air-tight.  The top, sides, and ends of each l i f t piece were of f a i r l y heavy high-grade glass.  Lights were seated i n 52 cm. r e f l e c -  tors and f i x e d inppo-sitio.n. 2 feet above the cabinet top.  10,  The l i g h t s were controlled by a time switch.  Air circulation  was provided f o r through an intake i n the base and outlets at the cabinet top.  Some of the d e t a i l s are i l l u s t r a t e d  i n the accompanying photographs.  11.  12.  The plants w.ere grown i n water culture s o l u t i o n . The nutrient solution used was the "Rubideux" solution described by Eaton (7).  The constituents  of t h i s solution and  t h e i r concentration are as follows: Millimoles per l i t e r .  Constituent  Grams per 10G l i t e r s .  Calcium n i t r a t e , Ca(¥0 ) .4H 0  4  94  Potassium n i t r a t e ,  3  30  2  27  Magnesium sulphate, MgS0^.7H 0  2  49  Potassium acid phosphate,  0.2  - 3  3  2  2  KNO3  Ammonium sulphate, (IH^gSG^ 2  Boric a c i d ,  H3BO3  KH2PO4  _  0.6  Manganese chloride, MnC^^H^Q;; -  0.1  Zinc sulphate,  0.04  ZnSO4.7H.2O  '  Iron was made a v a i l a b l e to the plants "by addition of the t a r t rate (freshly prepared 0.5 percent solution) i n small amounts at frequent i n t e r v a l s as required.  The concentration was  maintained by replacement of nutrients as.these were absorbed, and by changing the whole s o l u t i o n .  The hydrogen ion concen-  t r a t i o n varied from pH 6.0 to 6.5. This s o l u t i o n has been used i n preference t o the older widely-used Hoagland's solution because i t seems to possess the advantages described by the author.  The supply  of some of the nitrogen as ammonium tends to maintain a better ion balance, the 'low-level of phosphate makes i t easier to maintain available i r o n . The solution containers were wide-mquth glass jars of  13. 1.75 l i t e r volume (2-quart Mason) . The tops were closed "by corks with 5 holes f o r plants and an opening f o r additions of water.  Five cereal seedlings were grown i n each j a r , and  enough jars were used to make a t o t a l of 110 plants f o r both control and treated i n each experiment. The plant stock used i n these experiments was barley. Barley has been widely used i n experimental work of this kind because i t grows with vigor i n balanced nutrient solutions of pH 5.5 - 7.0. The v a r i e t y grown f o r experiment was Duckb i l l , the purest l i n e .seed obtainable from the Department of Agronomy, U n i v e r s i t y of B r i t i s h Columbia.  The importance of  homogeneous stock i n experimental work has been r i g h t l y stressed by many i n v e s t i g a t o r s . , The i-seed was soaked f o r 24 hours i n culture s o l u t i o n at 20.5°C.  I t was then placed on mesh screen over culture  solution to germinate i n a medium l i g h t i n t e n s i t y . .When the seedlings were 9 cm. i n length, a uniform s e l e c t i o n was transplanted t o the culture j a r s . The importance of guarding against plant diseases such as rusts and smuts has been recognized, and the plants were free from i n f e c t i o n . ... Measures were taken to prevent i n f e s t a t i o n by plant parasites such as aphids.  14.  Ill The conditions under which treatments took place, and the environmental factors are recorded i n the following pages, .Experiment 1. Sulphur Dioxide Records:F i r s t Treatment: 27 days a f t e r dry seed was soaked. Healthy plants with 3-4 leaves. Duration;  6 hours gas h i s t o r y . Treatments on 2 successive days. Each treatment of 3 hours duration. Time of day: morning.  Concentration:  Average - 0.40 p.p.m. Maximum - 0.50 p.p.m. Minimum - 0.30 p.p.m.  Humidity:  Very high, 90-100% r e l a t i v e humidity,  Temperature:  65-75° F.  V i s i b l e Injury  oQ% of the total,, leaf area.  Env i r onment Re e o r d s : Light:..  Sunlight i n l a t e f a l l .  15. Light:  P l a n t s were grown i n a greenhouse  Temperature:  Average Maximum - 1 0 5 ° Pv Minimum -  Humidity:  55° P.  Average  (high)  Maximum - 100^ r e l a t i v e h u m i d i t y . Minimum The dry  10% r e l a t i v e h u m i d i t y .  experiment was d i s c o n t i n u e d  56 days a f t e r t h e  seed was soaiked.. . E x p e r i m e n t 2.  Sulphur Dioxide First  Records T r e a t m e n t : 27 days a f t e r  d r y s e e d was soaked.  H e a l t h y p l a n t s w i t h 3-4 l e a v e s . Duration:  12 h o u r s gas h i s t o r y . 3 t r e a t m e n t s a t 10-day  intervals.  Time o f d a y : m o r n i n g . Concentration:  Average  - 0.30 p.p.m.  Maximum - 0.33 p.p.m. Minimum - 0.27 p.p.m. Humidity:  V e r y h i g h , 90-100% r e l a t i v e h u m i d i t y ,  Temperature:  65-80° P.  Visible  5% o f t h e t o t a l l e a f  Environment  Injury:  area.  Records:-  Light:  Sunlight  i n late  fall.  P l a n t s were grown i n a greenhouse Temperature:  Average  ------  Maximum - 1 0 5 ° P.  16. Temperature:  Minimum - 55° F.  Humidity:  Average Maximum - 100% r e l a t i v e humidity. Minimum -  70% r e l a t i v e humidity.  The experiment was discontinued 56 days a f t e r the dry seed was soaked.  Ebrperiment 3. Sulphur Dioxide Records:F i r s t Treatments 15 days a f t e r dry seed was soaked. Healthy plants with 3-4 leaves. Durat ion:  480 hours gas h i s t o r y . Continuous treatment. 24 hours each day.  Concentration  Average - 0.27 p.p.m. Maximum - 0.41 p.p.m. Minimum - 0.18 p.p.m.  Humidity:  F a i r l y low.  Usually a maximum of  60-62% r e l a t i v e humidity "before the i l l u m i n a t i o n , then a decrease to a minimum of about 40% at the end of the i l l u m i n a t i o n period. Temperature:  F a i r l y high.  Usually a minimum of  about 67-68° F before i l l u m i n a t i o n , then an increase to a maximum of about 90-94° F at the end of the i l l u m i n a t i o n period.  17. V i s i b l e Injury:  None. Leaf colour of the experimental plants was a l i g h t e r green than that of the control plants.  Environment Records:Light:  Plants were grown i n cabinets. Illumination from four 1000-Watt gas f i l l e d bulbs seated i n 52 cm. r e f l e c tors at a distance of 7 feet. 16 hours i l l u m i n a t i o n each day.  Temperature:  Average -:84° P. Maximum  98° P.  Minimum - 66° P. Humidity:  Average - A9% r e l a t i v e humidity. •Maximum - 66% r e l a t i v e humidity. Minimum - 37% r e l a t i v e humidity.  The experiment was discontinued 35 days a f t e r the dry.lseed was soaked.  Experiment 4. Sulphur Dioxide Records:P i r s t Treatment: 15 days a f t e r dry seed was soaked. Healthy plants with 3-4 leaves. Duration:  51 hours gas h i s t o r y . Treatments on 17 consecutive days. Each treatment of 3 hours duration. Treatment began 1 hour a f t e r the i l l u m i n a t i o n had commenced.  18.  Concentration;  Average - 0.36 p.p.m. Maximum -0.58 p.p.m. Minimum - 0.25 p.p.m.  Humidity:  Medium - 55-70% r e l a t i v e humidity.  Temperature:  75-85° F.  V i s i b l e Injury:  None.  Environment Records:•Light:  Plants were grown i n cabinets. Illumination  from three 1000-Watt  gas f i l l e d bulbs seated i n 52 cm. r e f l e c t o r s at a distance of 7 feet 16 hours i l l u m i n a t i o n each day. Temperature;  Average - 78° F-. Maximum - 90 F. Minimum - 70° F.  Humidity:  Average - 59% r e l a t i v e humidity?. Maximum - 77% r e l a t i v e humidity. Minimum - 4 7% r e l a t i v e humidity.  The experiment was discontinued 32 days a f t e r the dry seed was soaked.  19.  IV  The results of the experiments have been recorded as y i e l d data and as analyses.  These records are grouped i n  tables i n the following pages. The plants i n experiment 4 had the least f i n a l green and dry weights. these plants.  I t seemed of interest to show the size of  The accompanying picture was taken 12 days a f t e r  fumigation commenced, or 5 days before the experiment was d i s continued .  20. Fa Me go..- 1. Y i e l d Data. Tillers per Plant.  Green Weights (Grams per 100 Plants)  •loisture (as %•)  Dry Weights (Grams per 100 Plants)  230 119 71 420  91.1 92.0 93.4 91.8  20.5 9.5 4.7 34.7  168 94 65 327  89.2 93.0 93.9  91.2  18.1 6.6 4.7 28.7  205 100 66  90.8 93.0 93.4  91.9  18 .8 7.0 4.4 30.2  Sxperiment 1. Control  1.61  Leaves •Stems Roots Total  Treated  1.51  Leaves Stems Roots Total  Experiment 2. Control  1.26  Leaves Stems  Roots Total Treated, Leaves Stems Roots Total  371  1.16 177 94 63  334  90.0 93.1 93.4 91.5  17.7 6.5 4.1  28.3  21. Table Ho. 1 (continued). Y i e l d Data. Tillers per Plant.  Green Weights (Grams per 100 Plants)  Moisture (as %)  Dry Weights (Grams per 100 Plants)  109 102 119 330  81.9 89.3' 93.5 88.4.  19.7 11.0 7.7  102 99 109 '310  82.3 89.0 93.7 88.5  18.1 10.9 6.8 35.8  77.3 59.7 65.4  202.4  84.7 90.6 92.1 88.8  11.87 5.63 5.16  77.0 61.1 65 .6 203.7  84*8. 90.7 92.1 88.9  11.73 5.77 5.23 22.63  Experiment 5.Control  3.02  Leaves Stems Roots. Total  Treated  38.4  2.93  Leaves Stems Roots Total  Experiment 4. Control  2.09  Leaves Stems Roots . Total Treated Leaves Stems Roots Total  22.66  2.14  22 «  The  t r a n s p i r a t i o n by  to keep the c o n t a i n e r s has been determined. "has been d e r i v e d .  the p l a n t s has been measured as water added  at l e v e l . I n t h i s way  The  evaporation from blank  a net t r a n s p i r a t i o n v a l u e f o r the  and Shantz (8)  j  Volume o f water t r a n s p i r e d . freight o f dry m a t e r i e l formed.  Transpiration Coefficient:  r e s u l t s of t h i s c a l c u l a t i o n are i n the next t a b l e  T a b l e Mo.  Experiment 1  Plants  2  Transpiration"Coefficient  Control Treated  255 152  Control Treated  194 180  Control Treated  692 548  Control Treated  653 618  To determine the absorption  o f n u t r i e n t s from the  s o l u t i o n , analyses were made o f the dry p l a n t m a t e r i a l . have been done according by  plants  Using these values a t r a n s p i r a t i o n c o e f f i c i e n t has  been c a l c u l a t e d as f o l l o w s , a f t e r t h a t o f B r i g g s  The  containers  to the recognized  culture  These analyses  -quantitative methods advocated  the A s s o c i a t i o n o f O f f i c i a l A g r i c u l t u r a l Chemists i n the p u b l i c a t i o n  (9) " O f f i c i a l and T e n t a t i v e  Methods.....", 4th e d i t i o n , 1936.  r e s u l t s o f these analyses are recorded i n the f o l l o w i n g t a b l e .  The  25.  Table Ho. 3 Plants Analysed.  As percent of dry plant material.  Ash  Ca  MgO  K  Control-  18.14  1.19  0.79  7.78  1.98  0.85  6.IS  Treated  17.56  1.17  Q.71  7.54  1.71  0.76  5.72  Control  17.66  1.46  0.82  7.46  1.92  0.75  6.48  Treated  17.47  1.46  0.85  7.59  1.84  0.74  6.51  Control  18.11  1.42  0.64  7.38  2.06  1.02  6.61  Treated  18.16  1.27  0.53  6.73  2.84  0.95  5.87  Control  18.56  1.24  0.60  7.61  1.94  1.31  6.34  Treated  17.73  1.18  0.58  7.23  1.98  1.27  6.30  I (total)  Experiment 1  Experiment 2.  Experiment '3.  Experiment 4.  24.  The r e l a t i o n of the extent of r i s i b l e i n j u r y to the factors involved has been the subject of exhaustive  research  by many i n v e s t i g a t o r s . This work has been done c h i e f l y under f i e l d conditions.  The r e s u l t s are obtained by observing the  extent and character of the i n j u r y , by measurements of y i e l d , and by c o r r e l a t i n g these with the experimental  conditions.  The r e s u l t s are compared with controls as standards.  Inform-  a t i o n of t h i s nature i s extensive, and has an all-important bearing upon the subject of sulphur dioxide e f f e c t s . Growth of plants i n culture solutions i n a cont r o l l e d environment and treated with low concentrations of sulphur dioxide has been studied by Swain and Johnson ( 6 ) . This research shows c a r e f u l study throughout and the authors reached the following conclusion: "The r e s u l t s of t h i s study, indicate that wheat plants grown i n nutrient solutions under optimum conditions of a r t i f i c i a l l i g h t and humidity which were favorable to rapid and uniform growth, and which at the same time could be accurately controlled and recorded, w i l l t o l e r a t e an exposure to sulphur dioxide of several, hours d a i l y i n concentrations  25  below those at which t y p i c a l f o l i a r markings are produced without showing any signs of injurious actions i n t h e i r general appearance, i n t h e i r rate of growth, or i n the dry weight of tissue which they develop." Studies have been made of the effects of sulphur dioxide exposure upon stomatal movement i n plants to determine possible c o r r e l a t i o n s .  These morphological investigations  are based upon the reasonable assumption that any i n j u r y to the plant organism takes place through the leaf stomata. The effect of exposure to sulphur dioxide upon the chemical composition of the growing plant has been studied ever since f o l i a r accumulation of sulphur compounds was f i r s t observed'.  Correlations of sulphur content with the exposure  have "been made. Data r e l a t i n g to the effect upon' carbohydrate metabolism and protein eontent has been compiled. The w r i t e r has been p r i v i l e g e d i n observing an extremely i n t e r e s t i n g study of the effect of sulphur dioxide treatments on the carbon dioxide-oxygen metabolism of plants, carried out "by Dr. M . Katz and associates. ( 1 0 ) In t h i s research the problem has been approached  from  the point of view of inorganic nutrient absorption through the roots.  The assumption has been made that the chemical con-  s t i t u t i o n , as shown by a n a l y s i s , i s a measure of the nutrient absorption.  Prom the r e s u l t s of experiments with wheat plants  ( 1 1 ) , the w r i t e r reached the conclusion that the problem could only be considered when plants werejaot severely injured.  The  reasons f o r t h i s are - that the chemical composition of plants may vary widely according to the stage of growth and, since the  26. study i s a r e l a t i v e one, i t would be unwise t o i n c r e a s e the p r o b a b i l i t y or error. The experimental c o n d i t i o n s and r e s u l t s have been recorded i n s e v e r a l t a b l e s above.  The y i e l d d a t a , t r a n s p i r a t i o n c o e f f i c i e n t s and  analyses have been grouped i n the f o l l o w i n g t a b l e .  An accompanying  t a b l e o f r e l a t i v e v a l u e s with the c o n t r o l as 100 has been made. The w r i t e r i s unprepared.to defend t h i s l a t t e r on e i t h e r l o g i c a l o r mathematical grounds.  These t a b l e s may be conveniently r e f e r r e d to i n  this discussion. V i s i b l e i n j u r y i s not dependent  upon the c o n c e n t r a t i o n o f the  gas alone, as has been shown by the c l a s s i c a l experiments i n t h i s T h i s f i n d i n g i s r e a f f i r m e d by these experiments. o f primary importance.  field.  Humidity i s a f a c t o r  A t 90 - 100$ humidity, v i s i b l e i n j u r y was  produced by sulphur d i o x i d e a t c o n c e n t r a t i o n s o f 0.40 p.p.m. and 0.50 p.p.m.j while a t 37 - 66$ r e l a t i v e humidity, no i n j u r y was produced by 0.27 p.p.m.j and a t 55 - 70$ humidity, no i n j u r y was produced by 0.56 p.p.m. The d u r a t i o n f a c t o r i s a l s o shown; 6 hours on 2 successive days a t a c o n c e n t r a t i o n o f 0.40 p.p.m. produced v i s i b l e i n j u r y on 30$ o f the t o t a l l e a f area o f p l a n t s i n t h e 3-4 l e a f stage; b u t 12 hours on 3 days a t 10-day i n t e r v a l s with a c o n c e n t r a t i o n o f 0.30 p.p.m. produced v i s i b l e i n j u r y on only 5$ o f t h e t o t a l l e a f area.  Where no v i s i b l e i n j u r y was  produced, t h e e f f e c t s o f d u r a t i o n a r e seen i n comparing the dry weight yields.  27.  Table Hp. 4 Exper. 1  Exper. 2  Exper. 5  Exper. 4  C. Absolute Values.  Tillers Green weight % Moisture Dry weight Transpiration % Ash % Calcium % Magnesium % Potassium % Sulphur % Phosphorus % Mitrogen  1.61 420 91.8 34.7 235 18.14  1.51 327 91.2 28.7 152 17.56  1.19  1.17  0.79 7.78  0.71 7.54 1.71 0.76 5.72  1.98 0.85 6.13  1.26 371  91.7 30.2 194 17.66 1.46 0.82 7.46 1.92 0.75 6.48  1.16 334 91.5 28.3 180 17.47 1.46 0.83 7.59 1.84 0.74 6.51  5.02 350 88.4 38.4 692 18.11 1.42 0.64 7.38 2.06 1.02 6.61  2.93 2.09 310 202.4 88.5 88.8 55.8 22.7 548 653 18.16 , 18.56 1.27 1.24 0.53 0.60 6.73 7.61 2c 84 1.94 0.95 1.31 5.87 6.54  2.14 205.7 88.9 22.6 618 17.73 1.18 0,58 7.23 1.98 1.27 6,30  R e l a t i v e Values. ( C o n t r o l as 100) Tillers Green weight % Moisture Dry weight Transpiration % Ash % Calcium $ Magnesium % Potassium % Sulphur % Phosphorus /S H i t r o g e n  100 100 100 100 103 100 100 100 100 100 100 100  93,7 77.9 99.3 82.7 65.0 95.7 98.3 89.9 96,8 86.4 89.4 93.3  G§ - the c o n t r o l p l a n t s . T# - the t r e a t e d  plants.  100 100 100 100 100 100 100 100 100 100 100 100  92,1 90.0 99.8 93.7 95.0 98.9 100.0 101.2 97.8 95.8 98.7 100.5  100 100 100 100 100 100 100 100 100 100100 100  97.0  95.9 100.1 93.2 80.0 100.3 89.5 82e 8 91.2 157.4 95.1 .88.8  100 100 100 100 100 100 100 100 100 100 100 100  102.4 100.6 100.1 99.9 95.0 95.5 95.1 96.6 95.0 102.1 96.9 99.4  28.  T i l l e r i n g by t r e a t e d p l a n t s d i d not d i f f e r g r e a t l y from the t i l l e r i n g by the c o n t r o l s .  The r a t i o o f t r e a t e d to c o n t r o l v a r i e d from  92,1 s 100 t o 102.4 : 100. Dry weights o f the t r e a t e d p l a n t s were decreased i n rough proportion  t o t h e degree o f treatment w i t h i n t h e experimental c l a s s e s (a)  producing v i s i b l e i n j u r y , and (b) producing no v i s i b l e i n j u r y .  Plants  with v i s i b l e i n j u r y t o 30$ o f the l e a f area produced a dry weight 82.7$ as heavy as t h a t o f t h e c o n t r o l s ; while p l a n t s with i n j u r y t o 5$ o f the l e a f area produced dry weight 93.7$ as heavy as the c o n t r o l s .  P l a n t s with no  v i s i b l e i n j u r y but exposed t o 480 hours csf treatment with sulphur at a c o n c e n t r a t i o n  dioxide  o f 0.27 p.p.m. produced a d r y weight 93,2$ as heavy as  t h a t o f the c o n t r o l s ; while p l a n t s exposed t o 51 hours o f gas a t a conc e n t r a t i o n o f 0.36 p.p.m. produced 99.9$ as much as t h e c o n t r o l dry weight. T r a n s p i r a t i o n d i f f e r e n c e s were very n o t i c e a b l e between the c o n t r o l and t r e a t e d p l a n t s i n each experiment.  I n order o f experiment  number, the r a t i o s o f c o n t r o l t r a n s p i r a t i o n c o e f f i c i e n t to those o f t h e t r e a t e d ares  100:65,  100:95,  100:80,  100:95.  The d i f f e r e n c e i n  t r a n s p i r a t i o n c o e f f i c i e n t s o f i n j u r e d p l a n t s i s greater than the t r u e t r a n s p i r a t i o n d i f f e r e n c e , due to t h e r e d u c t i o n o f t r a n s p i r a t i o n surface through i n j u r y ,  nevertheless,  t r a n s p i r a t i o n d i d take p l a c e .  a diminishment o f the per u n i t  surface  T h i s might be explained p a r t l y upon the  b a s i s o f p a r t i a l c l o s u r e o f t h e stomata through some e f f e c t o f the sulphur dioxide. Scarth  Such an e f f e c t might be due to a change i n a c i d i t y , h e l d by  (12) to be o f great importance i n stomatal movement.  diminishment o f t r a n s p i r a t i o n might be explained respiration rate.  Or t h e  by p o s t u l a t i n g a  diminished  According t o evidence advanced by workers i n t h i s f i e l d ,  t r a n s p i r a t i o n i s a process r e q u i r i n g a l a r g e expenditure o f energy by the  29»  p l a n t , and consequently factor.  the r a t e o f r e s p i r a t i o n may become a l i m i t i n g  These explanations would f u l l y account f o r the measured decreases  i n t r a n s p i r a t i o n recorded. Calcium, magnesium, and potassium content per u n i t dry weight were diminished i n a l l the t r e a t e d p l a n t s excepting those i n experiment 2, where the treatment was o f 12 hours d u r a t i o n o n l y . i n d i c a t i n g a diminished  T h i s i s regarded  a b s o r p t i o n r a t e o f these elements.  as  The magnesium  content i n the p l a n t s grown i n the greenhouse was about 0.8$, while i n the p l a n t s grown i n cabinets i t was about 0.6$.  T h i s d i f f e r e n c e might  be c o r r e l a t e d with the l i g h t -conditions and the l e a f f r a c t i o n s o f the t o t a l green weights. Nitrogen  ( t o t a l ) content was diminished  appreciably i n the  treatment where v i s i b l e i n j u r y t o 30$ o f the l e a f area was produced (H r a t i o was 100s95.3) and i n the treatment o f 480 hours d u r a t i o n i n which no v i s i b l e i n j u r y was produced (N r a t i o ,  was 100:88.8).  In the other two  experiments there were no a p p r e c i a b l e d i f f e r e n c e s between c o n t r o l and treated plants.  The n i t r o g e n content was q u i t e high i n a l l p l a n t s .  Phosphorus was a l s o f a i r l y h i g h . 1 and 3 are d i f f e r e n t .  The r a t i o s o f P:N i n experiments number  T h i s might be regarded  as an e f f e c t upon the  phosphoproteins i n the growing p a r t s o f the p l a n t ; that i s , the amount o f phosphorus was l e s s i n the p l a n t s which showed a diminished the l a t e r  growth d u r i n g  stages. Sulphur content showed great v a r i a t i o n i n the t r e a t e d p l a n t s .  Arranged i n order o f experiment numbers, the r a t i o s o f c o n t r o l to t r e a t e d ares  100:'86.4,  100:95.8,  100:137.4,  100:102.1.  C o n s i d e r a t i o n o f the  treatments and the r e s u l t s o f analyses suggests t h a t the sulphate absorption by t r e a t e d p l a n t s from the n u t r i e n t s o l u t i o n i s lowered, but  30.  t h a t treatment o f s u f f i c i e n t d u r a t i o n w i l l i n c r e a s e the t o t a l I n experiment 3, the p l a n t s were t r e a t e d f o r 480 was  sulphur.  hours; the sulphur  content  i n c r e a s e d 37.4$ f o r the whole p l a n t . Ash  analyses  i n c l u d e s i l i c o n t r a c e s which cannot be  i n c u l t u r e work o f t h i s k i n d . . number 3, was  The  ash f o r a l l experiments,  excepting  lower i n the treated, p l a n t s than i n the c o n t r o l p l a n t s .  R a t i o s o f c o n t r o l to t r e a t e d p l a n t s are 100:95.7, 100:95.5.  excluded  100:98.9,  100:100.3,  T h i s agrees, i n general, with the r e s u l t s found i n separate  a n a l y t i c a l procedures.  The  s l i g h t l y higher ash found i n the t r e a t e d  p l a n t s o f experiment number 3 i s e v i d e n t l y the r e s u l t o f the h i g h  sulfur  content o f the t r e a t e d p l a n t s . Absorption of transpiration.  i s regarded as a p h y s i o l o g i c a l f u n c t i o n independent  At the present  time both f u n c t i o n s can only be  ex-  p l a i n e d by p o s t u l a t i n g expenditure o f energy by the p l a n t t o maintain them. Other researches seem to e s t a b l i s h t h i s f a c t . a lowered t r a n s p i r a t i o n was  recorded  n u t r i e n t s per gram dry weight was the absorption)  I n these experiments  and a lowered absorption o f i n o r g a n i c  measured (assuming analyses  to i n d i c a t e  i n t r e a t e d p l a n t s as compared to c o n t r o l p l a n t s .  This  can be explained on the hypothesis t h a t sulphur d i o x i d e lowers the r e s p i r a t i o n r a t e o f the p l a n t without a f f e c t i n g the r a t e o f i n the same degree. substantiate t h i s  No d i r e c t texperimental  photosynthesis  evidence i s advanced to  explanation.  SUMMARY: Four experiments upon b a r l e y p l a n t s have been conducted under d e s c r i b e d environmental c o n d i t i o n s .  These experiments c o n s i s t e d o f  treatments with sulphur d i o x i d e i n a range o f low concentrations  with  31. averages from 0.27  p.p.m. to 0.40  p.p.m.  The d u r a t i o n o f exposure i n  the treatments v a r i e d from 6 t o 480 hours; and the range o f between experiments was  from 40$ r e l a t i v e humidity  V i s i b l e i n j u r y was was  to  100$.  only produced when the r e l a t i v e  extremely high and the extent was  humidity  humidity  i n f l u e n c e d by the d u r a t i o n o f  posure as w e l l as the c o n c e n t r a t i o n o f the gas.  No v i s i b l e i n j u r y  exwas  produced i n medium and f a i r l y low h u m i d i t i e s by lengthy, treatments o f sulphur d i o x i d e at the same c o n c e n t r a t i o n s . T i l l e r i n g by the p l a n t s was not a p p r e c i a b l y a l t e r e d by  treatments.  The y i e l d , as measured by dry weights, was  reduced i n three experiments.  The r e d u c t i o n i n y i e l d could be considered  as roughly p r o p o r t i o n a l to the  s e v e r i t y o f the treatment (a) producing v i s i b l e i n j u r y to the p l a n t s and (b) producing  no v i s i b l e i n j u r y to the p l a n t s .  T r a n s p i r a t i o n by the p l a n t s was measured. t r a n s p i r a t i o n by  the t r e a t e d p l a n t s was  decreased.  In each experiment Transpiration  c o e f f i c i e n t s have been c a l c u l a t e d . The  dry m a t e r i a l was  analyzed by recognized  q u a n t i t a t i v e methods  f o r the f o l l o w i n g elements; calcium, magnesium, potassium ( i n c l u d i n g sodium t r a c e ) , s u l f u r , phosphorus, and n i t r o g e n ( t o t a l ) ; and an ash determination  was made.  The r e s u l t s showed a decrease i n the  o f these elements per u n i t dry weight i n the more severe S u l f u r was higher i n the experiments o f l o n g d u r a t i o n .  concentration  treatments. Consideration of  the r e s u l t s l e a d to the c o n c l u s i o n t h a t absorption o f i n o r g a n i c n u t r i e n t s from s o l u t i o n i s decreased  by treatment with sulphur d i o x i d e .  T r a n s p i r a t i o n and a b s o r p t i o n are now r e q u i r i n g an energy expenditure by the p l a n t . and a b s o r p t i o n o f i n o r g a n i c n u t r i e n t s may  regarded  as p l a n t f u n c t i o n s  The decrease i n t r a n s p i r a t i o n  be explained i f the  sulphur  d i o x i d e i s assumed to decrease the r e s p i r a t i o n r a t e o f the p l a n t  without  32. the same relative decrease i n the rate of photosynthesis.  •55.  VI  For constant d i r e c t i o n and a i d i n t h i s study, and f o r generous provision of laboratory f a c i l i t i e s , s i n c e r e a p p r e c i a t i o n to Dr. A.H.  the w r i t e r wishes to express h i s  Hutchinson o f the U n i v e r s i t y .  To Dr. M o r r i s Katz o f the National Research C o u n c i l thanks are a l s o due  f o r suggesting  t h i s study, and f o r h e l p i n g i n s e v e r a l  problems i n v o l v e d . The  suggestions  made by Dr. F. Dickson and Dr. H. H a r r i s have  been o f great v a l u e i n t r e a t i n g the  data.  T h i s study would not have been p o s s i b l e without the a i d o f grants f o r equipment from the N a t i o n a l Research C o u n c i l , Ottawa, and from the U n i v e r s i t y o f B r i t i s h Columbia.  VII  Bibliography,  Holmes, J.A,, F r a n k l i n , E.C„, and Gould, R.A. Report o f the Selby Smelter Commission, U.S. Bureau o f Mines, B u l l e t i n 98, (528 pp.), Washington, D.C., 1915,  Benton, A.F, Gas f l o w meters f o r s m a l l r a t e s o f flow, Ind. & Eng. Chem., 2, pp. 62S-629, 1919,  Thomas, M,D»,  and Abersold, J.N,  Automatic apparatus  f o r the determination o f  small concentrations o f s u l f u r d i o x i d e i n the a i r . Ind, & Eng. Chem,, A n a l . Ed. 1, 14, 1929.  G r i f f i n , S.W.,  and Skinner,  W'.W.  Small amounts o f s u l f u r d i o x i d e i n the atmosphere. I.  Improved method f o r determination o f SOg when  p r e s e n t i n low c o n c e n t r a t i o n i n a i r . Ind, & Eng. Chem., 24, 1932.  Davis, A.R.,  and Hoagland,  D.R.  An apparatus f o r the growth o f p l a n t s i n a c o n t r o l l e d environment. 3, pp.  Swain, R.E.,  277-292,  Plant  Physiology,  1928.  and Johnson,  A.B.  E f f e c t of sulphur d i o x i d e on wheat development. A c t i o n at low c o n c e n t r a t i o n s . Chem., 28, pp.  42-47,  Ind. &  Eng.  1936.  Eaton, E.M.  -  Automatically  operated sand-culture  equipment.  J o u r . Agr. Research, 53, pp. 433-444,  B r i g g s , L . J . and Shantz,  1936.  H.L.  R e l a t i v e Water Requirements of P l a n t s . J o u r . Agr. Research 3,, And  pp. 1-64,  1914,  subsequent p u b l i c a t i o n s .  The A s s o c i a t i o n o f O f f i c i a l A g r i c u l t u r a l Chemists. O f f i c i a l and T e n t a t i v e Methods o f A n a l y s i s , 4 t h e d i t i o n , Washington, D.C.,  Ketz,  Dr. M.,  and  1936.  associates.  Unpublished d a t a . N a t i o n a l Research C o u n c i l , Canada.  •  56.  11.  Jack,  W.R. Unpublished d a t a .  12.  S c a r t h , G.W.  _  Report to I t h a c a Congress o f P l a n t Science, 1926.  Bot. Gaz., 82, p. 454, And subsequent  1926.  publications.  

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