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Some effects of micronutrient elements upon certain enzymes, vitamin C content, and general metabolism… Magel, Harold Alexander 1955

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SOME EFFECTS OF MICRONUTRIENT ELEMENTS UPON CERTAIN ENZYMES, VITAMIN C CONTENT, AND GENERAL METABOLISM OF THE TOMATO (LYCOPERSICUM ESCULENTUM) by HAROLD ALEXANDER MAGEL A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN AGRICULTURE in the Department of Horticulture We accept this thesis as conforming to the standard required from candidates for the degree of MASTER OF SCIENCE IN AGRICULTURE Members of the Department of THE UNIVERSITY OF BRITISH COLUMBIA Apri l , 1?55 i ABSTRACT Tomatoes of the Vetomold 121 variety were grown i n sand cultures i n the University of B r i t i s h Columbia green-house. Interiors of cl a y pots, i n which the plants were grown, were painted with asphalt several times to prevent the plants from absorbing micronutrient elements from the materials used to manufacture the pots* Ripe tomatoes were analyzed f o r vitamin C, ash, and t o t a l soluble sugar; green f r u i t was analyzed f o r catalase; and plant leaves were analyzed f o r catalase, oxidase and peroxidase a c t i v i t y . The f r u i t y i e l d and plant weight were also recorded. Plants receiving boron i n t h e i r nutrient solutions were observed to be l e s s susceptible to tomato leafmold than plants receiving no boron. Boron was also observed to stimu-l a t e fibrous root growth. Boron s i g n i f i c a n t l y decreased oxidase but increased catalase and peroxidase i n the leaves, and s i g n i f i c a n t l y i n -creased plant weight. Trends indicate that boron also i n -creased y i e l d and depressed the sugar content of the f r u i t . Copper tended to increase plant weight, and decrease ash i n the ripe tomatoes and catalase i n the leaves. This micronutrient also s i g n i f i c a n t l y decreased peroxidase. EesXilifesL:^ indicated a tendency f o r manganese to increase vitamin C i n ripe tomatoes and catalase i n green f r u i t , but i i to depress sugar, ash, y i e l d , and plant weight. Manganese s i g n i f i c a n t l y reduced catalase and peroxidase a c t i v i t y i n the leaves. A s i g n i f i c a n t increase i n peroxidase and a s i g n i -f i c a n t decrease In catalase of tomato plant leaves was effected by molybdenum. F r u i t y i e l d was depressed somewhat by the. addition of molybdenum to a complete nutrient sol u t i o n . Zinc appeared to increase catalase i n green f r u i t and oxidase In tomato leaves; however, i t did s i g n i f i c a n t l y increase plant weight and decrease l e a f catalase. ACKNOWLEDGEMENTS The author wishes t o express h i s indebtedness t o Dr. G.H. H a r r i s , P r o f e s s o r of H o r t i c u l t u r e , f o r h i s w i l l i n g a s s i s t a n c e g i ven throughout the working of the experiment. Thanks are a l s o due t o Dr. A.P. Bar ss and Dr. C A . Hornby of the Department of H o r t i c u l t u r e , P r o f e s s o r A. Renney, Department of Agronomy, and Dr. J.J.R. Campbell, Department of D a i r y i n g , of the F a c u l t y o f A g r i c u l t u r e f o r t h e i r i n t e r e s t i n t h i s problem. The h e l p f u l advice of Dr. C.G. Woodbridge, Dominion E x p e r i -mental S t a t i o n , Summerland, B.C., i n the second h a l f of the experiment i s g r a t e f u l l y acknowledged. TABLE OP CONTENTS PAGE I n t r o d u c t i o n " 1 Review of L i t e r a t u r e 3 M a t e r i a l s and Methods 9 Greenhouse Procedure •• 9 Experimental Procedure 11 Determination of Catalase 12 Determination of Oxidase II4. Determination of Peroxidase l£ Determination of V i t a m i n C 17 Determination o f Sugars 18 Determination of Ash 18 R e s u l t s 19 Discussion. 31+ 1 Conclusions 1+2 Summary I4I4. L i t e r a t u r e C i t e d i+7 Appendix 51 SOME EFFECTS OF MICRONUTRIENT ELEMENTS UPON tEETAlNYENZYMES VITAMIN C CONTENT, AND GENERAL METABOLISM OF THE TOMATO (LYCOPERSICUM ESCULENTUM) INTRODUCTION Wallace (ip.) r e c e n t l y s t a t e d , " D i f f i c u l t problems i n biochemistry a l s o await s o l u t i o n and, i n p a r t i c u l a r , i t may be expected t h a t the i n t e n s i f i c a t i o n o f research on the r o l e o f t r a c e elements i n enzyme systems w i l l y i e l d a r i c h harvest of knowledge;" Much i n t e r e s t i s being aroused today regarding the r o l e of m i c r o n u t r i e n t elements i n p l a n t n u t r i t i o n . T h i s study was conducted to a s c e r t a i n some e f f e c t s of the f i v e m i c r o n u t r i e n t elements (boron, copper, manganese, molybdenum, and z i n c ) upon the c a t a l a s e , oxidase and peroxidase enzymes, and the v i t a m i n C, ash, and t o t a l s o l u b l e sugar content o f the tomato. The i n v e s t i g a t i o n undertaken was i n c o n j u n c t i o n w i t h a general program being conducted by the P l a n t N u t r i t i o n s e c t i o n of the Department of H o r t i c u l t u r e at the U n i v e r s i t y of B r i t i s h Columbia. The purpose of the program I s t o d e t e r -mine the e f f e c t s of m i c r o n u t r i e n t elements upon p l a n t growth. Some m i c r o n u t r i e n t elements h a v i been found t o be a p a r t of the molecule of c e r t a i n p l a n t enzymes. Copper i s the p r o s t h e t i c group i n oxidase, w h i l e i r o n i s the p r o s t h e t i c group i n c a t a l a s e and peroxidase. Enzymes are c o l l o i d a l proteinaceous c a t a l y s t s p r o -duced by the l i v i n g c e l l . R e l a t i v e l y l i t t l e work has been done to determine t h e i r a c t i v i t y i n p l a n t s , but i t i s known that c a t a l a s e i s present i n most p l a n t s and animals. This enzyme a c c e l e r a t e s the decomposition of hydrogen peroxide i n p l a n t s i n t o water and molecular, non-activated oxygen. The hydrogen peroxide i s produced by metabolic processes In the p l a n t . Catalase i s very s e n s i t i v e to heat (destroyed above 5>0 degrees Centigrade) and a c i d s , and p r e f e r s a n e u t r a l t o s l i g h t l y a l k a l i n e medium. Peroxidase i s an enzyme having the same f u n c t i o n as c a t a l a s e ; however, the hydrogen peroxide Is broken down i n t o water and atomic oxygen. Oxygen i n t h i s form w i l l cause ox-i d a t i o n of r e a d i l y o x i d i z a b l e substances. Peroxidase ex-h i b i t s i t s g r eatest a c t i v i t y at a pH of approximately Oxidases are enzymes which w i l l reduce molecular oxygen to atomic oxygen. The atomic oxygen I s then d i r e c t e d to any s l o w l y s e l f - o x i d i z i n g compounds. The a c t i o n of o x i d -ase i s independent of hydrogen peroxide ( t h i s c h a r a c t e r i s t i c d i s t i n g u i s h e s oxidases from p e r o x i d a s e s ) . Mono- and p o l y -phenol oxidases c a t a l y z e the o x i d a t i o n of mono- and p o l y -phenols to quinones. Oxidase i s the most a c t i v e when the f>H of the medium i s s l i g h t l y a l k a l i n e . V itamin C ( a s c o r b i c a c i d ) i s an important v i t a m i n i n human n u t r i t i o n . Along w i t h c i t r u s f r u i t s and s t r a w b e r r i e s , tomatoes are recognized as a v a l u a b l e source of v i t a m i n C (approximately 15 to 25> m i l l i g r a m s per 100 grams of f r e s h 3 weight of f r u i t ) . A s c o r b i c a c i d i s capable of undergoing a l -t e r n a t e o x i d a t i o n and r e d u c t i o n , and hence has the q u a l i t i e s r e q u i r e d by an Intermediate i n b i o l o g i c a l o x i d a t i o n s . Because m i c r o n u t r i e n t elements can i n f l u e n c e the ab-s o r p t i o n r a t e of other minerals by p l a n t r o o t s (12, 13» 32, 36, 39), i t was a l s o decided to determine the t o t a l ash eon-t e n t of the f r u i t . Furthermore, i f a p l a n t ' s a b s o r p t i o n r a t e i s i n f l u e n c e d , then i t s v i g o u r and photosynthetic a c t i v i t y should be s i m i l a r l y a f f e c t e d . Photosynthesis i s necessary f o r sugar formation i n p l a n t s ; hence the t o t a l s o l u b l e sugar content of the tomato f r u i t s was determined. REVIEW OF LITERATURE E z e l l and C r i s t (11+) s t a t e t h a t disease and n u t r i e n t element a v a i l a b i l i t y could a l t e r the enzymatic c o n d i t i o n of a p l a n t , i n order to have t h e i r f u l l e f f e c t upon the p l a n t ' s growth. I n other words, these f a c t o r s could be considered as growth r e g u l a t o r s . The same workers found a d i s t i n c t c o r r -e l a t i o n e x i s t i n g between p l a n t growth and c a t a l a s e a c t i v i t y of l e t t u c e , r a d i s h and.spinach p l a n t s , as a f f e c t e d by c a l -cium n i t r a t e and calcium a c i d phosphate f e r t i l i z e r t r e a t -ments. A s i m i l a r r e s u l t occurred w i t h l l j . to 17 year o l d Mcintosh apples t r e e s i n an experiment conducted by H e i n i c k e ( 2 1 ) . He found t h a t the c a t a l a s e a c t i v i t y of the leaves v a r i e d I n v e r s e l y w i t h the crop y i e l d ; however, t r e e s which y i e l d e d crops r e g u l a r l y from year!.to year showed no v a r i a t i o n i n t h e i r c a t a l a s e content. I n d i r e c t c o n t r a s t t o these f i n d i n g s , Appleman (Ij.) showed c a t a l a s e to be higher I n e t -i o l a t e d than green b a r l e y s e e d l i n g s . B a i l e y and McHargue ( 6 ) showed t h a t tomato f r u i t s have l e s s c a t a l a s e than the f o l i a g e . E a r l i e r , these same workers, (7) found the c a t a l a s e a c t i v i t y of tomato f r u i t s t o d e c l i n e as the f r u i t s r ipened; but, i n the case of the p l a n t ' s l e a v e s , the c a t a l a s e content increased u n t i l the time of senescence. I n v e r s e l y , Gustafson and others (17) found a p o s i t i v e c o r r e l a t i o n between the c a t a l a s e a c t i v i t y and the r e s p i r a t i o n r a t e of r a p i d l y growing tomato f r u i t s . E a r l i e r work by He i n i c k e (19) l e a d him t o suggest th a t the determination of c a t a l a s e may be an i n d i c a t i o n of the n u t r i t i v e c o n d i t i o n of f r u i t t r e e s . Appleman ( 3 ) , studying the catalase a c t i o n of potatoes, found more oxygen evolved from l a r g e mature potatoes than from those which were small and immature. When Reed (35) worked w i t h pineapples he found c a t a l a s e to be l a c k i n g i n green f r u i t s , but t h a t r i p e pineapples were r i c h i n c a t a l a s e ; thus he concluded t h a t c a t a l a s e must develop i n pineapples as the f r u i t s r i p e n . The p h y s i o l o g i c a l and n u t r i t i o n a l c o n d i t i o n s of apple t r e e s g r e a t l y i n f l u e n c e the presence of the enzyme c a t -a l a s e , according to Heinicke ( 2 1 ) . He found t h a t the c a t -alase a c t i v i t y of apple leaves exposed to s u n l i g h t was greater than that of leaves removed from shaded p o r t i o n s of 5 the tree. The catalase a c t i v i t y of apple leaves from trees growing i n a sod cover crop was less than i n the leaves from trees growing under clean c u l t i v a t i o n ; leaves of trees grow-ing i n clay loam s o i l s were higher i n catalase than those i n sandy s o i l s . Further experiments by the same worker showed that the catalase a c t i v i t y i n leaves from pruned apple trees growing i n a sod cover crop was greater than i f they had not been pruned. Bailey and McHargue (6) found the catalase a c t i v i t y of tomato f r u i t s to decline s t e a d i l y with an increase i n the copper content of the nutrient solution; however, the leaves of a l f a l f a plants showed maximum catalase action when a concentration of 0.01 ppm. of boron, copper, or zinc were used, but that one ppm. of manganese depressed i t . Alexander (1) observed catalase to be greater i n boron de-f i c i e n t squash plants than the plants receiving boron. Loustalot and others (28) found that the photosyn-t h e t i c rate of tung trees was greater i n the forenoon than i n the afternoon, and that copper and zinc d e f i c i e n c i e s d e f i n i t e l y reduced the photosynthetic rate (a copper de-f i c i e n c y more so than a zinc d e f i c i e n c y ) , and that carbon dioxide assimilation was greatly depressed, even though the leaves showed no v i s u a l deficiency symptoms. In tomato leaves, Bailey and McHargue (6) showed that polyphenol oxidase increased with increasing concent-rations of copper (up to a maximum concentration of 0.10 ppm.). They found that with a l f a l f a plants, one ppm. of 6 boron or z i n c produced the gr e a t e s t oxidase a c t i v i t y , but one ppm. of manganese depressed i t . E a r l i e r , the same auth-ors (7) presented evidence t h a t oxidase c o n c e n t r a t i o n i s highest i n the youngest tomato l e a v e s : as the leaves in«-creased i n m a t u r i t y , the oxidase content s t e a d i l y d e c l i n e d . Nason (33) found t h a t polyphenol oxidase of tomato leaves decreased when the p l a n t became d e f i c i e n t i n copper. K l e i n (26) was unable t o e s t a b l i s h a d i r e c t e f f e c t of boron upon the t y r o s i n a s e (polyphenol oxidase) a c t i v i t y of tomato p l a n t s . Arnon (5) e s t a b l i s h e d polyphenol oxidase t o be l o c a l i z e d i n the l e a f c h l o r o p l a s t s of beet l e a v e s . MacVicar and B u r r i s (29) found polyphenol oxidase t o be g r e a t e r i n boron d e f i c i e n t tomato, tobacco and soybean p l a n t s than i n p l a n t s which r e c e i v e d adequate boron. Peroxidase was found by B a i l e y and McHargue (6) t o decrease w i t h an increase of copper i n tomato l e a v e s . I n a l f a l f a p l a n t s , they were able to produce maximum peroxidase a c t i o n w i t h separate, one ppm. a p p l i c a t i o n s of z i n c , mang-anese, and boron. I n e a r l i e r work, these men (7) concluded t h a t peroxidase decreased c o n s i s t e n t l y as the tomato f r u i t s approached m a t u r i t y . Leaf a n a l y s i s showed peroxidase t o i n -crease p r o g r e s s i v e l y w i t h i n c r e a s i n g m a t u r i t y , and then t o decrease g r a d u a l l y as the leaves approached senescence. Hivon and h i s co-workers ( 23 ) were unable t o f i n d a c o n s i s t e n t r e l a t i o n s h i p between the manganese n u t r i t i o n of fie l d - g r o w n soybean 1plants and the v i t a m i n C content of the 7 le a v e s . S i m i l a r r e s u l t s were obtained by Gum and others (16) when they analyzed the leaves and f r u i t s of manganese d e f i c i e n t tomato p l a n t s . S c o t t and Walls (38) analyzed 11 v a r i e t i e s of tomatoes and found the as c o r b i c a c i d content to vary from 17.1 to 25.1 m i l l i g r a m s per 100 grams of f r e s h f r u i t . I n the U.S.S.R., Dmi t r i e v (11) observed that boron and manganese lowered the ash content of red c l o v e r stems and l e a v e s . Anderssen (2) of South A f r i c a discovered a c h l o r o t i c c o n d i t i o n e x i s t i n g i n f r u i t t r e e s which was due to a copper d e f i c i e n c y . The t r e e s were growing i n a w e l l -drained a c i d s o i l . I t was the c h l o r o t i c t i s s u e which always showed a higher ash content when compared w i t h the ash i n normal l e a f t i s s u e . Muhr ( 3 D showed th a t a l a c k of boron In sugar beets s i g n i f i c a n t l y increased the i r o n , magnesium, and c a l c i u m content of the r o o t s compared w i t h those which r e c e i v e d adequate boron. Gum and other (16) grew beet and tomato p l a n t s i n sand c u l t u r e s . They found that a boron de^ f i c i e n c y decreased the dry matter content of both vegetables, th a t a manganese d e f i c i e n c y decreased the dry matter content of beet r o o t s , t h a t the boron and manganese content of the f o l i a g e v a r i e d d i r e c t l y w i t h the amount of these elements a v a i l a b l e t o the plants., t h a t manganese has i t s g r e a t e s t c o n c e n t r a t i o n i n the f o l i a g e of young p l a n t s , and t h a t the boron and manganese content of beet and tomato leaves must be extremely low before d e f i c i e n c y symptoms w i l l appear. 8 The t o t a l sugar content of leaves and stems of boron d e f i c i e n t tomato p l a n t s was shown by Johnston and Dore (2I4.) t o be higher than i n p l a n t s which r e c e i v e d adequate boron. They concluded t h a t the increase was due t o the breaking down of the conducting system caused by the boron d e f i c i e n c y . The reducing and t o t a l sugar content of tomato f r u i t s were found to be lower I n boron and manganese d e f i c -i e n t p l a n t s than i n the c o n t r o l p l a n t s ( 1 6 ) . Cook and M i l l a r (9) and Keese (25) found t h a t boron increased the (37) sugar content of sugar beets; however, Saruy^calculated a s l i g h t decrease i n the sugar content of the r o o t s when boron was a p p l i e d . H a r r i s ( 1 8 ) showed the percentage of s o l u b l e carbohydrates of red r a s p b e r r i e s t o inc r e a s e from s o i l app-l i c a t i o n s of manganese or z i n c . G i l y a r o v s k i i and Chernov (lf>) o f the U.S.S.R. c l a i m t h a t boron a p p l i c a t i o n s to tomato and cucumber p l a n t i n g s caused an inc r e a s e i n y i e l d s , a sh o r t e r p e r i o d of r i p e n i n g , and improved f r u i t q u a l i t y . Manganese, according t o Vlasyuk ( l j . 0 ) , when a p p l i e d t o s o l u t i o n s c o n t a i n i n g n i t r a t e n i t r o g e n , increased the sugar content of sugar beets by 1 . 1 percent. Boron, copper, manganese, and z i n c , as shown" by H a r r i s ( 1 8 ) , decreased the sugar content of c a r r o t s grown on a peat s o i l . But, both boron and z i n c Increased the sugar i n the r o o t s when the c a r r o t s were grown on a c l a y s o i l . When t h i s same crop was grown I n a sandy loam s o i l and f e r t i l i z e d w i t h the f o u r 9 mentioned m i c r o n u t r i e n t elements, no s i g n i f i c a n t e f f e c t upon t h e i r sugar content r e s u l t e d . MATERIALS AND METHODS GREENHOUSE PROCEDURE I n October, 1952, and September, 1953* at the Univ-e r s i t y of B r i t i s h Columbia, tomato seed of the Vetomold 121 v a r i e t y was sown i n greenhouse f l a t s c o n t a i n i n g c a r e f u l l y washed fresh-water sand. The r e s u l t i n g p l a n t s were thinned and t r a n s p l a n t e d i n t o s i m i l a r f l a t s c o n t a i n i n g sand. When the p l a n t s were approximately 5 inches i n h e i g h t , they were i n t r a n s p l a n t e d i n t o 1 0-inch p o t s . The pots used^the e x p e r i -ment were o r d i n a r y c l a y pots f i l l e d w i t h f r e s h water-washed sandj the i n t e r i o r of each pot was given s e v e r a l coatings of an a s p h a l t - v a r n i s h p a i n t manufactured by the B r i t i s h American P a i n t Company. The purpose of the c o a t i n g was t o prevent the piLant r o o t s from o b t a i n i n g any m i c r o n u t r i e n t element, ' i m p u r i t i e s which may be present i n the m a t e r i a l s used t o manufacture the p o t s . The sand was pl a c e d i n the pots and leached w i t h d i s t i l l e d water f o r a p e r i o d of one week p r i o r to the t r a n s p l a n t i n g date. The source of d i s t i l l e d water came from a t i n - l i n e d s t i l l . The pots were arranged i n f o u r randomized blocks of 12 p l a n t s each and were p l a c e d i n the same room i n the Univ-10 ersity greenhouse. Each block consisted of 12 treatments with one plant per pot. The treatments were as l i s t e d below: i treatment 1. complete (major nutrients and iron only — Hoagland's solution) (C) 2. complete plus boron only (C+B) 3. complete plus copper only (C+Cu) I4.. complete plus manganese only (C+Mn) 5. complete plus molybdenum only (C+Mo) plus 6. complete^zinc only (C+Zn) 7. complete plus boron, copper, manganese, molybdenum, and zinc (C+M) 8. complete plus copper, manganese, molyb-enum and zinc O(M-B) 9« complete' plus boron, manganese, molyb-denum, and zinc C+(M-Cu) 10. complete plus boron, copper, molybdenum, and zinc C+(M-Mn) 11. complete plus boron, copper, manganese, and zinc C+(M-Mo) 12. complete plus boron, copper, manganese, and molybdenum C+(M-Zn) Chemically pure reagents were used to prepare the stock solutions* The plants were fed the six macronutrients twice each week, the micronutrient elements once every tenu 11 days, i r o n once per week, and d i s t i l l e d water whenever i t was r e q u i r e d by the p l a n t s . D i s t i l l e d water was used as the source of water throughout the experiment. For the complete s o l u t i o n , 500 ml. per p l a n t of Hoagland's n u t r i e n t s o l u t i o n were used, and f o r the micro-n u t r i e n t a p p l i c a t i o n s , 500 ml. were a p p l i e d per p l a n t of a s o l u t i o n c o n t a i n i n g the f o l l o w i n g c o n c e n t r a t i o n s : 1.0 ppm. boron as b o r i c a c i d 0.1 ppm. copper as c u p r i c s u l f a t e 1.0 ppm. manganese as manganous c h l o r i d e 2.0 ppm. molybdenum as molybdic a c i d 1.0 ppm. z i n c as z i n c s u l f a t e f e r r i c I r o n was a p p l i e d at a c o n c e n t r a t i o n of 1.0 ppm. as^ c i t r a t e * The 1952 p a r t of the experiment terminated May 20, 1953, and the 1953 p a r t ended A p r i l 7, 195k' EXPERIMENTAL PROCEDURE In determining the c a t a l a s e a c t i v i t y , the gasov. metric water displacement method d e s c r i b e d by Landon (27) was used.. The height of water i n the l e v e l l i n g f l a s k was moved downward and kept at the same height as the water i n the graduated b u r e t t e tube, thus m a i n t a i n i n g a constant pressure of one atmosphere f o r the e v o l v i n g oxygen gas t o act a g a i n s t . The oxygen entered the b u r e t t e tube at the 12 top and f o r c e d the water downward. In 1952, c a t a l a s e determinations were made on green f r u i t s and the j u i c e of r i p e f r u i t s , but no analyses were c a r r i e d out f o r oxidase of peroxidase.. I n the 1953 p a r t of the experiment, c a t a l a s e , oxidase and peroxidase were d e t e r -mined on the p l a n t l e a v e s . The o l d e r leaves of new shoots were gathered each sampling date at 11:00 a.m. The s e l e c t e d leaves were approximately 5 centimeters i n l e n g t h and 2 centimeters I n width, and were l o c a t e d near the top o f each p l a n t . A number of t r i a l s were run f o r each b l o c k t r e a t -ment . The j u i c e of the r i p e f r u i t s was analyzed f o r c a t a -l a s e , v i t a m i n C, ash ( c o n d u c t i v i t y ) , and t o t a l s o l u b l e sugar:; during the 1952 phase of the experiment, and f o r v i t a m i n C, ash, and t o t a l s o l u b l e sugar ; I n the 1953 p a r t of the experiment. As soon as the f r u i t s r ipened on the v i n e s , they were harvested, weighed, and analyzed the same day. To e x t r a c t the j u i c e , the f r u i t s were ground t o a f i n e mass i n an o r d i n a r y meat g r i n d e r and s t r a i n e d through c l o t h . The r e s u l t i n g j u i c e was used f o r the d e s i r e d d e t e r -minations . DETERMINATION OP CATALASE In order to c a l c u l a t e the volume of oxygen gas evolved as a r e s u l t of c a t a l a s e a c t i o n , 5 m i l l i l i t e r s of 13 f r u i t j u i c e were n e u t r a l i z e d w i t h one gram of powdered, ch e m i c a l l y pure c a l c i u m carbonate. The j u i c e and chalk mixture were then p i p e t t e d i n t o one arm of the c a t a l a s e tube. I n the other arm was p l a c e d 5 m i l l i l i t e r s of 10 v o l -ume (3 percent) hydrogen peroxide. The rubber tubing from the top of the b u r e t t e tube was connected t o the neck of the c a t a l a s e tube, and the l a t t e r was then p l a c e d i n t o a water bath (20 degrees Centigrade) f o r s e v e r a l minutes u n t i l the contents of the c a t a l a s e tube reached a constant temperature. The contents of the tube were then a g i t a t e d back and f o r t h at a r a t e of 20 times per minute f o r a t o t a l o f 5 minutes f o r each sample. The volume of oxygen e v o l v i n g was recorded at one-minute i n t e r v a l s . As the oxygen evolved, the water l e v e l i n the l e v e l l i n g bulb was kept at the same height as the water i n the b u r e t t e . When determining the c a t a l a s e a c t i o n i n tomato l e a v e s , a half-gram sample of l e a f t i s s u e was ground t o a f i n e suspension w i t h a half-gram of powdered calcium c a r -bonate i n a mortar. Seven m i l l i l i t e r s of d i s t i l l e d water were added t o the mortar before the l e a f t i s s u e was ground. A f t e r a smooth suspension was formed, a 5 m i l l i l i t e r a l i -quot was withdrawn w i t h a p i p e t t e and placed i n t o one arm of the c a t a l a s e tube. I n t o the other.airm was p l a c e d 5 m i l l i -l i t e r s of 3 percent hydrogen peroxide, and the procedure was then continued i n the same manner as :for the determination of t h i s enzyme i n the f r u i t j u i c e . While e v a l u a t i n g f o r c a t a l a s e i n apple l e a v e s , Heinicke (19) found t h a t the a g i t a t i o n of the supernatant l i q u i d of the l e a f p r e p a r a t i o n , Immediately b.efore the l e a f sample i s withdrawn, i s not harmful to the enzyme. However, the contents of the c a t a l a s e tube must be a g i t a t e d back and f o r t h during the r e a c t i o n , or e l s e the e v o l u t i o n of molec-u l a r oxygen w i l l be q u i t e slow. DETERMINATION OP OXIDASE The oxidase a c t i v i t y o f tomato leaves was analyzed c o l o r i m e t r i c a l l y by Using the KLett-Summerson p h o t o e l e c t r i c c o l o r i m e t e r w i t h a number 5k f i l t e r . B a i l e y and McHargue (7) and Arnon (5) found t h i s method s a t i s f a c t o r y f o r d e t e r -mining the oxidase a c t i o n i n tomato and beet l e a v e s , r e -s p e c t i v e l y . The procedure used i n t h i s experiment was the same as the one used by these workers. A l l t e s t s were r e -peated a number of times. A half-gram sample of l e a f was p l a c e d i n a mortar w i t h 10 m i l l i l i t e r s of potassium phosphate b u f f e r (pH 7.3). A f t e r the t i s s u e was f i n e l y ground, i t was poured i n t o a 100 m i l l i l i t e r graduate. Then one m i l l i l i t e r of a one p e r -cent p y r o g a l l o l s o l u t i o n was added. Ten minutes l a t e r the graduate's contents were made up t o a t o t a l volume of 100 m i l l i l i t e r s w i t h d i s t i l l e d water, then set aside f o r 5 15 minutes. F i n a l l y , a sample of the solution was placed into the colorimeter tube and the reading was recorded. A high reading indicates low oxidase, since the solution remains greenish i n color; i n contrast, high oxidase a c t i v i t y i s Indicated by an amber-brown solution which forms; as a r e -su l t of the p y r o g a l l b l being oxidized to purpurogalline. A very high oxidase content produces a c l e a r amber s o l -u tion. DETERMINATION OF PEROXIDASE The method used f o r determining peroxidase a c t i v i t y In t h i s experiment was that of Davis'CLO). A buffered sub-strate consisted of the following reagents: 20 m i l l i l i t e r s of 2 percent soluble starch 10 m i l l i l i t e r s of 0.1 Normal sodium t h i o s u l f a t e ij..5 grams of potassium iodide s u f f i c i e n t 0.2 Normal sodium acetate buffer (pH i^.7) to make one l i t e r of buffered substrate. This substrate i s good f o r 2\\. hours only, and must be pre-pared f r e s h l y each day. The reaction time f o r a blank test i s determined f o r each f r e s h l y prepared substrate mixture. Each set of analyses f o r a block treatment Is accompanied by a blank t e s t . The reaction time f o r each i n i t i a l blank i s compared with the reaction time of one of the blanks a r b i -t r a r i l y selected. The r e s u l t of the comparison gives a 16 correction f a c t o r f o r the i n i t i a l blank readings of each newly prepared substrate. A l l the blank reaction times f o r one substrate are then m u l t i p l i e d by t h e i r corresponding correction factor In order to standardize the r e s u l t s . To prepare the l e a f tissue f o r analysis, one gram of l e a f was ground i n a mortar with 10 m i l l i l i t e r s of the buffered solution. Then t h i s was made up to 30 m i l l i l i t e r s with the buffered substrate and poured into a 125 m i l l i r l i t e r f l a s k . F i n a l l y , at zero time, one m i l l i l i t e r of one percent hydrogen peroxide was added to the f l a s k , and the flas k ' s contents were agitated immediately. The zero time and the reaction time ( i n seconds) f o r a dark blue color to appear i n the center of the f l a s k were recorded. Each de-termination was repeated several times. The reading of each block treatment was subtracted from Its respective corrected blank reading. Consequently, high peroxidase a c t i v i t y corresponded with a high peroxidase reading (iafter the above calc u l a t i o n s were completed). Several reactions occur between the beginning and the end of each reaction. In the case of the blank, the hydrogen peroxide slowly oxidizes the sodium t h i o s u l f a t e . As soon as the t h i o s u l f a t e i s completely oxidized, the hy-drogen peroxide releases free iodine from the potassium iodide; the iodine then reacts with starch to produce a deep blue color. The reactions are the same when the l e a f 17 t i s s u e i s added to the substrate and the hydrogen peroxide, except t h a t the peroxidase present i n the t i s s u e speeds up the r e a c t i o n . DETERMINATION OP VITAMIN C The indophenol method (8) was used to determine' the v i t a m i n C content of the f r e s h l y e x t r a c t e d tomato j u i c e . A 10 m i l l i l i t e r a l i q u o t o f j u i c e was p i p e t t e d i n t o I4.O m i l l i l i t e r s of 0.I4. percent o x a l i c a c i d . This mixture was a g i t a t e d and then c e n t r i f u g e d at 2f>00 r e v o l u t i o n s per minute f o r 10 minutes. Ten m i l l i l i t e r s of the c l e a r supernatant l i q u i d from the c e n t r i f u g e d j u i c e were then plac e d i n a p o r c e l a i n evaporating d i s h ; a 0.2 percent s o l u t i o n of sodium - 2 , 6-dichlorobenzenone-indophenol was added dropwise to the evaporating d i s h u n t i l a pink c o l o r appeared and p e r s i s t e d f o r a p e r i o d of 30 seconds. During the dropwise a d d i t i o n of the i n d i c a t o r dye, the j u i c e was c o n t i n u a l l y a g i t a t e d w i t h a g l a s s s t i r r i n g r o d . The num-ber of drops of dye r e q u i r e d to reach the endpoint was r e -corded. The analyses of the j u i c e samples and the standard were made i n d u p l i c a t e , and the standard was prepared d a i l y . While the tomato j u i c e was being c e n t r i f u g e d , 10 m i l l i l i t e r s of an as c o r b i c a c i d s o l u t i o n , c o n t a i n i n g 0.25 m i l l i g r a m s of v i t a m i n C per 10 m i l l i l i t e r s , were used to standardize the dye before each t i t r a t i o n of j u i c e . 18 The indophenol method i n v o l v e s the t i t r a t i o n of the dye (standardized against a standard s o l u t i o n of as-c o r b i c a c i d ) i n t o the unknown s o l u t i o n of v i t a m i n C. The ascorbic a c i d becomes o x i d i z e d and reduces the dye t o g i v e a c h a r a c t e r i s t i c p ink c o l o r a t i o n . DETERMINATION OF SUGARS The t o t a l s o l u b l e sugar, content of the j u i c e was measured w i t h a refrac t o m e t e r . DETERMINATION OF ASH The ash (mineral) content of the j u i c e was mea-sured by determining i t s e l e c t r i c a l c o n d u c t i v i t y ~by means of a s o l u - b r i d g e ; the r e s u l t s were recorded as percent KC1. RESULTS A l l data i n t h i s experiment were analyzed s t a t i s t i c -a l l y by a n a l y s i s of v a r i a n c e . A t y p i c a l i l l u s t r a t i o n of the method of c a l c u l a t i o n i s shown i n the Appendix. A l l tomato p l a n t s grown during the 1952 - 1953 ex-periment were vigorous and h e a l t h y , whereas the p l a n t s i n the 1953-1951+ experiment contacted tomato leafmold l a t e r i n the experiment on March 2 , 195U-- The o l d or senescent leaves were the f i r s t to show signs of the di s e a s e . The leaves were c a r e f u l l y removed from the p l a n t s at the f i r s t 19 sign of infection. However, the uncontrollably high green-house temperatures (up to and above 80 degrees Fahrenheit) TABLE 1. The Effect of Treatment Upon the Catalase Activity of Green Fruits (each weighing approximately 25 grams) From Four Blocks of Tomatoes (1952-1953) treatment total oxygen volume (c.c.) of evolved i n 5 minutes treatment mean A B C D 1. C 1.8 1.6 3.0 0.6 1.8 2. C+B 2.0 1.8 2.0 0.8 . 1.7 3. C+Cu 1.6 1.8 2.1 1.0 1.6 4. C+Mn 2.6 1.9 2.3 1.7 2.1 5. C+Mo 2.8 1.8 2.1 2.0 6, C+Zn 1.6 3 . 4 * 2.0 1.6 2.2 7. C+M 1.6 2.7 2.0 2.0 2.1 8 . C+(M-B) 2.2 2.3 2.7 1.4 2.2 9. C+(M-Cu) 2.0 1.6 2.0 . 1.5 1.8 10. C+(M-Mn) 1.9 2.5 . 1-4 1.1 1.7 11. C+(M-Mo) 2.1+ 2.3 t 1.7 2.2 2.2 12. C+(M-Zn) 1.7 2.1 2.6 2.0 2.1 block total 22.7 26.866 25.6AA 18.0 S.D. between block totals at .05 level = S.D. between block totals at .01 level = 4*86 c.c. 6.64 c.c. 20 were i d e a l f o r the spread of the disease. No evidence of micronutrient d e f i c i e n c i e s were v i s i b l e on the plants. TABLE 2. The E f f e c t of Treatment Upon the Catalase A c t i v i t y of Tomato Leaves (1953-195k). t treatment t o t a l volume of oxygen treatment evolved i n 5 minutes (c,c.) mean A B C D 1. C 30.2 29-5 31.0 . 30.1 30.2 2. C+B 3k.O 30.0 33.-2 33.3 32.6$ 3. C+Cu 28.8 28 .k 30.2 27-6 28.8 k- C+Mn 28.k 26.3 27.2 28.7 27.7AA 5. C+Mo 26.k 29.6 30.0 26.5 28.1 A 6. C+Zn 28.6 26.2 28.7 27.1 27-74A 7. C+M 32.6 30.0 31.2 33.0 31.7 8. G+(M-B) 27-3 27.9 29.0 26.6 27.7AA ,9. C+(M-Cu) 32.7 33.6 31.k 33.2 32.7AA 10. G+(M^Mn) 29.0 31.5 31.0 31.7 30.8 11. C+(M-M6.) 33.0 28.8 32.0 31.0 31.2 12. C+(M-Zn) 32.k 30.6 32.7 3k.2 32.5* block t o t a l 363. k 352.k 367.6". 363.0 S.D . between treatment means at .05 l e v e l = 1.9 c c . S.D. between treatment means at .01 l e v e l = 2.5 "c .c. 21 When the t o t a l p l a n t weights were being recorded, i t was observed t h a t p l a n t s which r e c e i v e d boron i n t h e i r n u t r i e n t s o l u t i o n s had many more f i b r o u s r o o t s than p l a n t s not r e c e i v i n g boron. CATALASE Table 1 shows th a t there were no s i g n i f i c a n t d i f f -erences i n the c a t a l a s e a c t i v i t y of the green f r u i t s as a r e s u l t of the va r i o u s m i c r o n u t r i e n t element treatments. I t does show th a t there was a s i g n i f i c a n t d i f f e r e n c e due to the p o s i t i o n of the b l o c k s i n the greenhouse. Table 2 shows th a t the c a t a l a s e a c t i v i t y of the leaves was very much gr e a t e r than t h a t i n the green f r u i t s , and t h a t there were s i g n i f i c a n t d i f f e r e n c e s due t o t r e a t -ments. A comparison of the complete n u t r i e n t s o l u t i o n w i t h s o l u t i o n the complete n u t r i e n t ^ p l u s the v a r i o u s s i n g l e microelements r e v e a l s t h a t boron s i g n i f i c a n t l y i n c r e a s e d c a t a l a s e a c t -i v i t y , whereas the other m i c r o n u t r i e n t s depressed i t . A comparison of the complete p l u s a l l m i c r o n u t r i e n t s added (C+M) w i t h those where one of the r e s p e c t i v e f i v e micro-n u t r i e n t s had been omitted, shows th a t where boron was omitted, the c a t a l a s e decreased. Prom Table 3 i t was noted t h a t where boron only was added t o the complete s o l u t i o n , there was a tr e n d to de-crease v i t a m i n Cj where boron was omitted from the complete p l u s other m i c r o n u t r i e n t s (C+(M-B)), v i t a m i n C tended t o 22 i n c r e a s e . This i n d i c a t e s a tendency f o r boron to depress v i t a m i n C, although the r e s u l t s are not s i g n i f i c a n t . Where manganese only was added to the complete n u t r i e n t s o l u t i o n v i t a m i n C tended to inc r e a s e more than from any other t r e a t -ment, but when i t was omitted from the complete p l u s micro-TABLE 3. The E f f e c t of Treatment Upon the Vitamin C Content of Tomato F r u i t s (1952-1953) treatment b l o c k averages of v i t a m i n treatment C (mg. per 100 g. f r e s h wt.) mean A B C D 1. C 15.2 15-6 13.4 16.8 15.3 2. C+B III-. 7 14-3 15.1 14.2 14.6 3. C+Cu 14-3 18.8 16.9 15.1 16.3 4- C+Mn 18.8 20 .5 18.5 16.1 18.5 5. C+Mo 13 . 4 18.0 17.9 18.4 I 6 . 9 6. C+Zn 16.1 16.7 16.6 16.2 16.4 7. C+M 17 .0 . 14.3 15.2 18.5 16.3 8. C+(M-B) 17.2 18.1 14.8 18.8 17.2 9. C+(M-Cu) 17.7 20 .5 16.1 16.4 17.7 10. C+(M-Mn) 18.8 17.6 17.0 17.1 17 .6 11. C+(M-Mo) 15.2 17 .9 17.5 17.0 16.9 12. C+(M-Zn) 16.1 I6.7 15.1 18.6 16.6 block t o t a l 194.5 209.0 194.1 203.2 N.S.D. 23 n u t r i e n t s treatment i t s t i l l i n c r e a s e d . I t i s noted t h a t TABLE /+. The E f f e c t of Treatment Upon the Vi t a m i n C Content of Tomato F r u i t s (1953-1954) treatment mg. of v i t a m i n C per treatment 100 g. f r e s h weight mean A B C D 1. C 22.7 21.3 22.8 23.8 22.7 2. C+B 25.3 21.9 21.9 25.1 23.6 3. C+Cu 0 0 0 0 0 4- C+Mn 0 0 0 0 0 5. C+Mo 18.7 23.2 20.9 23.8 21.7 6. C+Zn 0 0 0 0 0 7. C+M 25.3 21.9 21.9 25.1 23.6 8. C+(M-B) 26.2 14.5 18.5 20.1 19..3 9. C+(M-Cu) 28.4 26.5 28.2 28.0 27.84* 10. C+(M-Mn) 17.5 23.9 21.7 21.2 21.1 11. C+(M-Mo) 24.1 22.0 30.0 23.9 25.0 12. C+(M-Zn) 22.3 26.4 27.0 28.1 26.04 block t o t a l 211.9 198.2 215.1 220.4 S.D. between treatment means at .05 l e v e l = 3«3 mg. S.D. between treatment means at .01 l e v e l = 4*5 mg. 2k there was a tendency f o r a l l micronutrients other than boron to increase vitamin C. Unfortunately, the complete plus the single micro-TABLE 5. The E f f e c t of Treatment Upon the Percentage of Toital Soluble Sugar, i n Tomato F r u i t s (1952-1953) treatment t o t a l soluble sugar . (%) treatment mean A . B C D 1. C k.5o 3.50 k.75 k.5o k-3i 2. C+B 3.50 3.75 k.OO • 3.75 3.75 3. e+cu k.75 k.OO 3.75 k.5o k.25 k- C+Mn 3.85 k.50 k.25 3.50 k.03 5. C+Mo 6.50 k»5o 3.50 k.75 k.8l 6. C+Zn 5.00 k.OO 3.5o k.50 k.25 7. C+M 5.00 k.OO 3.50 3.75 k.06 8. C+(M-B) 5.00 k-25 k.50 3.50 k.31 9. C+(M-Cu) k.oo 3.75 k.OO k.5o k.06 10. C+(M-Mn) k.OO k.OO k.50 k.75 k.31 11. C+(M-Mo) 6.00 k.25 k.OO k.OO k-56 12. C+(M-Zn) 5.50 3.25 3.75 k.5o k.25 block t o t a l 57.60 k7.75 k8.00 50.50 S.D. between block t o t a l s at .05 l e v e l = 6.0$ S.D. between block t o t a l s at .05 l e v e l =8.0^ 2 5 elements of copper, manganese, and zinc did not produce f r u i t , as shown i n Table 4» In 1 9 5 3 - 1 9 5 4 * boron had the opposite e f f e c t upon vitamin G than i t had i n the 1 9 5 2 -1 9 5 3 season (Table 3 ) ; however, the resu l t s i n both Tables indicate a trend only and are not s i g n i f i c a n t . Omitting zinc or copper from'the complete plus micronutrient TABLE 6 . The E f f e c t of Treatment Upon the Ash (Conductivity) Content of Tomato F r u i t s ( 1 9 5 2 - 1 9 5 3 ) treatment block averages read as treatment % potassium chloride mean A B C D 1. C 0.54 0.57 0.53 0.53 0.54 2. C+B 0.58 0 .62 0.52 0.54 o.57 3. C+Cu o.5o 0.51 ^0.59 0.66 o.57 4- C+Mn o.5i 0.59 o.5i 0.69 0.58 5. C+Mo o.74 0.53 0.61 0 .60 0.62 6. C+Zn o.57 0.61 o.5i 0.53 0.56 7. C+M 0 .62 0.75 0.55 0.49 0 .60 8. C+(M-B) 0 .60 0.65 0.55 0.53 • 0.58 9. C+(M^Cu) 0.59 0.61 0.65 0.55 0.60 10.- C+(M-Mn) 0.54 0.69 o.5o o.55 0.57 11. C+(M-Mo) 0.68 0.55 0.59 0.60 0.61 1 2 . C+(M-Zn) 0.57 0.52 0.59 0.54 0.56 . block t o t a l 7.04 7.20 6.70 6.81 N.S.D. 26 solution s i g n i f i c a n t l y increased vitamin C i n compari-son with the complete (C) treatment (basis f o r a l l com-parisons). Where copper was omitted from the complete plus microelements solution, the vitamin C i n the f r u i t was s i g n i f i c a n t l y increased at the .01 l e v e l . The ommission of manganese from the complete plus micronutrients solution tended to depress the vitamin C content of the f r u i t * TOTAL SOLUBLE SUGAR.' The t o t a l soluble sugar*, content of the tomato f r u i t s grown i n the 1953-1951+ season was very si m i l a r to that of the 1952-1953 crop; hence the re s u l t s of the f i r s t season only are presented (Table 5)« Prom Table 5» i t can be seen that the addition of molybdenum to the com-plete solution gave tomatoes with the highest t o t a l s o l -uble sugari content; where molybdenum was omitted from the complete plus micronutrients solution, the. sugar content of the tomatoes was reduced, but i t was s t i l l greater.than i n those f r u i t s of the complete solu t i o n . The addition of boron or manganese to a complete solution resulted in. a tendency to depress the sugar content of the f r u i t . - the complete plus micronutrient solution, and the omission of copper from th i s same solution, also depressed the sugar content of the f r u i t . The presence of zinc i n a complete solution, or i t s absence from a complete plus microelements solution did not af f e c t the t o t a l soluble .sugar; ..content. 27 ASH Ash r e s u l t s f o r the 1953-1954 season were not r e -corded. The 1952-1953 r e s u l t s (Table 6) showed th a t no TABLE 7. The E f f e c t of Treatment Upon the Y i e l d (grams) of Tomato F r u i t s (1952-1953) treatment t o t a l f r u i t y i e l d (g.) treatment 1. C 1079 1662 783 2236 1440 2. C+B 1733 1719 2044 1464 1740 3. C+Cu 1732 1443 1210 1102 1372 4 . C+Mn 645 1533 1568 598 1086 5. C+Mo 539 1201 1674 1009 1106 6. C+Bn 701 1345 2030 1268 1336 7. C+M 1163 1371 1331 1766 11+08 8. C+(M-B) 917 815 1457 877 1017 9. C+(M-Cu) 1201 1398 1167 1169 1234 10. C+(M-Mn) 1079 1434 1298 1619 1358 11. C+(M-Mo) 959 1100 1675 I5ii4 1320 12. C+(M-Zn) 798 1670 1247 1625 1335 b l o c k t o t a l 12,546 16,691* 17,4844- 16,2774 S.D. between block t o t a l s at .05 l e v e l = 2377 g. 28 s i g n i f i c a n t d i f f e r e n c e s occurred between treatments or TABLE 8 . The E f f e c t of Treatment Upon the Oxidase A c t i v i t y ^ of Tomato Leaves ( 1 9 5 3 - 1 9 5 k ) treatment A co l o r i m e t e r reading treatment mean B C D 1 . C 162 1 9 2 1 8 0 2 1 8 1 8 8 . 0 2 . C+B 2 3 5 2 5 2 2 k k 2 6 9 250.044 3 . C+Cu 1 8 6 1 8 k 2 0 7 2 0 8 1 9 6 . 3 1+. C+Mn 1 8 1 2 0 6 1 9 6 2 1 0 1 9 8 . 3 5 . C+Mo - 232 196 203 1 9 5 2 0 6 . 5 6 . C+Zn 1 5 1 1 7 6 1 8 3 1 5 9 1 6 7 . 3 7 . C+M 2 6 3 278 2 5 3 2 8 9 2 7 0 . 8 4 4 8 . C+(M~B) 190 1 7 9 1 8 6 1 6 0 1 7 8 . 8 9. CH-(M-Cu) 261+ 261 278 266 2 6 c 7 . 3 * 4 1 0 . C+(M-Mn) 2 7 1 297 3 0 5 2 7 2 2 8 6 . 3 4 * U . C+(M-Mo) 2 3 9 2 7 3 2 6 9 2 5 0 2 5 7 . 8 4 4 1 2 . G+(M-Zn) 2 7 1 3 0 8 321 3 0 8 302.044 block t o t a l 2 6 k 5 * 2 8 0 2 2 8 2 5 2 8 0 k # A h i g h reading i n d i c a t e s low oxidase a c t i v i t y . S.D. between treatment means at .05 l e v e l = 21;8 u n i t s S.D. between treatment means at .01 l e v e l = 29.k u n i t s S.D. between block t o t a l s at .05 l e v e l = l 5 l u n i t s 29 blocks; however, trends did exist to show that molybdenum added to a complete solution produced the lowest ash content i n the f r u i t . When copper or molybdenum were omitted from the complete plus micr©nutrients solution, the ash (conductivity) decreased. The presence of the f i v e microelements i n a complete nutrient solution also decreased the ash content. Table 6 shows that the highest amount of ash was found i n f r u i t from plants receiving the complete nutrient s o l u t i o n . YIELD Block A i n Table 7 produced s i g n i f i c a n t l y l e s s f r u i t than the other blocks. No r e a l differences appeared between treatment means, although boron produced the greatest y i e l d when i t was added to the complete s o l u t i o n . Omitting boron from the complete plus micronutrients s o l u -t i o n gave the smallest f r u i t y i e l d . The addition or omis-sion of the other microelements from the complete solution . resulted i n the y i e l d s being le s s than the y i e l d s f o r eith e r the complete or the complete plus a l l micronutrients solu-t i o n * , OXIDASE Oxidase was markedly affected by zinc. Table 8 shows that the addition of zinc to a complete solu t i o n caused oxidase to be increased (the lowest reading In the Table); but, when zinc was omitted from a complete plus 30 micronutrient sol u t i o n (C+(M-Zn)), the smallest amount of oxidase was produced. Adding boron to a complete solution caused t h i s enzyme to decrease s i g n i f i c a n t l y belowVthe complete treatment. Omitting boron from a complete plus TABLE 9. The E f f e c t of Treatment Upon the Peroxidase A c t i v i t y of Tomato Leaves (1953-1951+) treatment reaction time i n seconds treatment A B G p mean 1. C 31+3 313 320 253 307 2. C+B 585 553 663 1+85 57244 :-3. C+Cu 195 21+0 275 230 235* i+. C+Mn 138 170 138 125 11+344 5. C+Mo 380 1+15 1+33 kOO 1+0744 6. C+Zn 21+8 200 333 315 27k 7. C+M 560 1+1+5 1+78 k88 1+93*4 8. C+(M-B) 335 285 1+33 290 336 9. C+(M-Cu) 525 603 508 k75 52844 10. C+(M-Mn) i+60 1+98 1+85 1+60 1+7644 11. C+(M-Mo) 508 1+80 1+98 1+15 1+7544 12. C+(M-Zn) 605 565 1+15 1+28 50344 block t o t a l 1+882 1+767 1+979 k36k S.D. between treatment means at .05 l e v e l = 72 seconds means S.D. between treatment >at .01 l e v e l = 97 seconds 31 TABLE 10. The E f f e c t of Treatment Upon the Total Plant Weight (Root Plus Top) i n 1952-1953 treatment t o t a l plant weight (root plus top) i n grams A B C D 1. C 225 264 269 244 251 2. C+B 253 273 274 220 255 3. C+Cu 247 286 261 231 256 4- C+Mn 193 248 222 240 226 5. C+Mo 215 251 262 222 238 6. C+Zn 258 318 329 295 300AA 7- C+M 214 268 241 258 245 8. C+(M-B) 234 242 268 206 238 9. C+(M-Cu) 209 249 274 233 241 10. C+(M-Mn) 238 287 274 321 2804 11. C+(M-Mo) 195 259 242 219 229 12. C+(M-Zn) 224 238 251 207 230 block t o t a l 2705 3183 3167 2896 S.D. between treatment means at .05 l e v e l = 26 grams S.D. between treatment means at .01 l e v e l = 35 grams S.D. between block t o t a l s at .05 l e v e l =178 grams S.D. between block t o t a l s at .01 l e v e l = 240 grams 32 a l l the micronutrients solution gave the same result, as a complete solution. Omitting copper, manganese, molyb-denum or zinc from a complete plus microelements solution TABLE 11. The E f f e c t of Treatment Upon the Total Plant Weight (Root Plus Top) i n 1953-1954 t o t a l plant weight (root treatment TREATMENT plus top) i n grams mean A B C D 1. C 185 191 234 224 209 2. C+B 765 638 724 663 69844 3. C+Cu 368 321 283 34+ 32944 4- C+Mn 191 224 171 162 187 5. C+Mo 299 317 236 278 283S 6. C+Zn 202 181 252 245 220 7. C+M 809 685 744 728 74244 8. C+(M-B) 261 196 212 217 222 9. C+(M-Cu) 797 648 780 669 72444 10. C+(M-Mn) 655 709 632 757 68844 11. C+(M-Mo) 665 738 788 701 72344 12. C+(M-Zn) 669 780 746 796 74844 block t o t a l 5866 5628 5802 5784 S.D. between treatment means at .05 l e v e l = 69 grams S.D. between treatment means at .01 l e v e l = 93 grams 33 resulted i n s i g n i f i c a n t l y l e s s oxidase a c t i v i t y . The omission of zinc from the complete plus micronutrients solution produced s i g n i f i c a n t l y l e s s oxidase than the com-plet e solution when the f i v e micronutrients were added. PEROXIDASE When copper or manganese were added to the com-plet e solution the peroxidase decreased s i g n i f i c a n t l y ; when boron or molybdenum were added to the complete s o l -a c t i v i t y s olution the peroxidase/\increased s i g n i f i c a n t l y . Omitting boron from the complete plus micronutrients solution gave the same eff e c t as a complete solution would give. When-ever boron was added to a solution the peroxidase a c t i v i t y increased. PLANT WEIGHT In 1952-1953" the t o t a l plant weight was increased s i g n i f i c a n t l y by the addition of zinc to a complete s o l -ution and by the omission of manganese from a complete plus micronutrients solution (Table 10) . A l l other treatments did not a f f e c t plant weight s i g n i f i c a n t l y , although there was a tendency f o r i t to be reduced i n comparison with the complete treatment plant weight. Blocks A and D produced s i g n i f i c a n t l y l e s s plant weight than the other two blocks. The t o t a l plant weight i n 1953-1951+ (Table 11) was s i g n i f i c a n t l y Increased,in comparison with the complete solutions t o t a l plant weight, by the following treatments: 34 the addition of boron, copper or molybdenum to a complete solution; the addition of a l l f i v e micronutrients to a complete solution; the omission from a complete plus micro-nutrients solution of e i t h e r of four nutrients - copper, manganese, molybdenum or zinc. In comparison with the com-plete treatment, t o t a l plant weight tended to decrease when manganese only was added to a complete s o l u t i o n . DISCUSSION BORON It was observed that plants being fed nutrient s o l -utions containing boron were quite r e s i s t a n t to leafmold (Cladasporium spp.) i n f e c t i o n , whereas plants receiving no boron were le s s r e s i s t a n t to the disease. In Table 1 a trend shows that when boron was omitted from the complete plus micronutrients solution, the catalase of immature tomato f r u i t s Increased. However, i n tomato leaves the opposite i s true «r adding boron to a complete solu t i o n In-creased the catalase s i g n i f i c a n t l y . The l a t t e r r e s u l t d i s -agrees with Alexander (1). Comparing Tables 1 and 2 con-c l u s i v e l y indicates that tomato plant leaves are much more abundant i n catalase than immature tomato f r u i t s . Experi-mental t r i a l s showed that r i p e tomato f r u i t s lacked cata-l a s e . This r e s u l t i s i n agreement with Bailey and 3 5 McHargue (6) who concluded that tomato fruits have less catalase than the foliage, and that catalase declines as tomatoes ripen. Boron did not have any significant effect upon vitamin C i n ripe tomato f ru i t s . In Table 3 omitting boron from a complete plus micronutrients solution caused an upward trend, but i n Table k the same treatment resulted i n a downward trend for vitamin C. Adding boron to a complete solution produced a decrease i n the total soluble sugar content of ripe tomatoes, but when this micronutrient was omitted from a complete solution containing the other four micronutrients, the sugar content increased to the same level as the complete (control) treatment. Consequently, It appears that boron depresses sugar formation i n ripe tomatoes. Saru ( 3 7 ) obtained similar results , although two other.workers ( 9 * 2 5 ) found that boron increased sugar formation. Prom Table 6 i t can be seen that boron had no definite effect upon the amount of ash i n ripe tomato f ru i t s . Dmitriev (11) found that boron depressed the ash content of red clover stems and leaves. The y ie ld of ripe tomatoes i n this experiment showed a downward trend when boron was deleted from a complete plus micronutrients solution, but by adding boron only to a complete solution the f ru i t y ie ld was 36 i n c r e a s e d . As shown i n Table 7, boron i s necessary i n tomato f r u i t p r o d u c t i o n ; G i l y a r o v s k i i and Chernov (15) obtained s i m i l a r r e s u l t s w i t h t h i s crop. The oxidase a c t i v i t y of tomato leaves was decreased by the a d d i t i o n of boron (Table 8). Any treatment which contained boron r e s u l t e d i n a s i g n i f i c a n t l y h i g h i n c r e a s e of the enzyme i n t h i s experiment. On the other hand, K l e i n (26) was unable to e s t a b l i s h a d i r e c t e f f e c t of t h i s element upon oxidase. Tables 9 and 11, r e s p e c t i v e l y , showed th a t peroxidase i n tomato leaves and the t o t a l p l a n t weight of tomatoes do e x h i b i t h i g h l y s i g n i f i c a n t i n c r e a s e s as a r e -s u l t of boron a p p l i c a t i o n s . COPPER Although copper had no e f f e c t (Table 1) on the c a t a -r e s u l t s l a s e a c t i v i t y of green tomato f r u i t s , Table 2/jshowed a s i g n i -f i c a n t l y h i g h increase i n the c a t a l a s e content of tomato leaves when copper was omitted from a complete n u t r i e n t s o l u t i o n c o n t a i n i n g the remaining f o u r m i c r o n u t r i e n t elements. B a i l e y and McHargue ( 6 ) , however, d i d f i n d t h a t c a t a l a s e de-creased w i t h an i n c r e a s e i n copper. I n s p i t e of the f a c t t h a t copper showed a t r e n d to increase v i t a m i n C i n r i p e tomatoes., when compared w i t h the complete treatment; o m i t t i n g t h i s microelement from the complete s o l u t i o n c o n t a i n i n g boron, manganese, molybdenum, 37 and z i n c , tended to give a g r e a t e r increase i n v i t a m i n C. During the second season (1953 - 195k) the l a t t e r treatment produced a s i g n i f i c a n t i n c r e a s e . The t r e n d i n Table 6 shows th a t the ash content of r i p e tomatoes decreased when copper was omitted from the complete p l u s m i c r o n u t r i e n t s s o l u t i o n . This t r e n d does not correspond w i t h Anderssen's r e s u l t s (2)« But when copper was omitted from treatment 9 i n Table 7* the tomato f r u i t y i e l d d e c l i n e d . By not i n c l u d i n g copper i n the complete p l u s m i c r o n u t r i e n t s s o l u t i o n , the oxidase a c t i v i t y decreased (Table 8) and t o t a l p l a n t weight (Table 11) of tomatoes i n -creased very s i g n i f i c a n t l y ; when copper alone was added t o the complete s o l u t i o n , p l a n t weight increased (Table 11). Copper caused a s i g n i f i c a n t decrease i n peroxidase (Table 9) when added to the complete s o l u t i o n ; by o m i t t i n g t h i s micro-element from the complete p l u s m i c r o n u t r i e n t s s o l u t i o n the peroxidase content i n c r e a s e d very s i g n i f i c a n t l y . However, the omission of copper from treatment 9 i n Tables 8, 9» and i l i s ndt what was thought to have caused the s i g n i f i c a n c e -a c t u a l l y , the presence of boron i n t h i s treatment s o l u t i o n was the i n d i r e c t cause* MANGANESE The c a t a l a s e a c t i v i t y o f green tomato f r u i t s showed an upward tr e n d when manganese was i n c l u d e d i n any n u t r i e n t 38 solution. However, i n Table 2 t h i s treatment reduced the catalase In the plant leaves to a s i g n i f i c a n t l y low l e v e l . Analysis of Table 2 shows that the exclusion of boron i n -d i r e c t l y caused the decline. Vitamin C i n tomatoes appeared to be affected by treatment 1+ (G+Mh). Leaving out the manganese from a complete plus micronutrients solution resulted i n a tendency f o r vitamin C to increase (Table 3)« The f a i l u r e of the complete plus manganese treatment to produce f r u i t (Table was unfortunate, hence no comparison of t h i s treatment between the two seasons can be made. Two other workers (16, 23) were unable to e s t a b l i s h any e f f e c t of manganese upon vitamin C i n tomatoes and soybean leaves, respectively. When manganese was not added to a complete plus micronutrients treatment, the t o t a l soluble sugar content was greater than when manganese was added to a complete solution. This trend indicates that manganese i s d e t r i -mental to sugar formation i n ripe tomato f r u i t s . The ef f e c t of t h i s element upon the ash of ripe tomatoes shows no d e f i n i t e trend, consequently I t i s concluded that man-ganese has no influence upon the ash. Manganese added to a complete treatment sol u t i o n tended to reduce the y i e l d of f r u i t . The omission of manganese from a complete plus micronutrients solution resulted i n a tendency to decrease 39 the f r u i t y i e l d very s l i g h t l y . Hence i t appears that manganese i s of no benefit to f r u i t production. Manganese added to a complete solution showed no eff e c t on the oxidase a c t i v i t y of tomato leaves (Table 8); however, i t did appear to decrease t o t a l plant weight (Tables 10 and 11). Peroxidase s i g n i f i c a n t l y declined as a resu l t of manganese addition to the nutrient media (Table 9)» but when manganese was omitted from the com-plete micronutrient media (C+M-Mn), oxidase decreased and peroxidase a c t i v i t y increased i n tomato leaves. When manganese was omitted from a complete plus micronutrients solution, the t o t a l plant weight increased s i g n i f i c a n t l y . MOLYBDENUM Adding molybdenum alone to a complete solu t i o n caused an upward trend i n the catalase of immature f r u i t s (Table 1), while omitting i t from the complete plus micronutrients treatment also s l i g h t l y increased the catalase a c t i v i t y . However, these r e s u l t s are non-signi-f i c a n t . The catalase a c t i v i t y of tomato leaves (Table 2) s i g n i f i c a n t l y decreased when molybdenum was included i n the complete nutrient solution (treatment 5 ) . but by omitting i t (treatment 11), catalase . v increased s l i g h t l y above the control treatment. Molybdenum did not appear to have any e f f e c t on vitamin C formation i n ripe tomatoes. When molybdenum ko was added to the complete solution there was a tendency f o r the sugar content of ripe f r u i t to increase; but when i t was omitted from the complete plus micronutrients solution, the sugar content was s t i l l greater i n compari-son with the complete treatment. The complete plus molybdenum solution resulted i n the lowest ash content of tomatoes, but when i t was omitted from the complete plus micronutrients solution, the ash con-tent s t i l l remained low. Apparently- then, the additive e f f e c t of other micronutrients i n treatment 11 must be as e f f e c t i v e as molybdenum only i n increasing the ash content of tomato f r u i t s . The tomato y i e l d s (Table 7) were decreased by the addition of molybdenum to a complete solution, but when molybdenum was not included i n the complete plus micro-nutrients solution, the y i e l d increased, although the complete treatment y i e l d was greater. Hence, t h i s trend seems to indicate that molybdenum depresses y i e l d . The t o t a l plant weight and the oxidase a c t i v i t y of the leaves d i d not appear to benefit from the addition of molybdenum. However, Table 9 showed that molybdenum plus a complete solution did increase peroxidase a c t i v i t y i n tomato leaves. On the other hand, omitting molybdenum (treatment 11) resulted i n highly s i g n i f i c a n t increases 6f peroxidase and plant weight, and a s i g n i f i c a n t decrease i n oxidase over the complete treatment plants* ZINC Adding zinc without the addition of other micro-nutrients, or omitting i t from the presence of other microelements, treatments 6 and 12 (plus and minus zinc) i n Table 1 produced s i m i l a r amounts of catalase i n green tomatoes. Since manganese, when added to a complete solution, produced an upward trend i n the catalase a c t i v i t y , i t i s very probable that its presence i n treatment 12 caused catalase to increase despite the omission of z i n c . However, Table 2 r e s u l t s s i g n i f i c a n t l y show that the complete plus zinc treatment decreased catalase, while the complete plus micronutrients minus zinc solution increased catalase a c t i v i t y i n tomato leaves. Although no results were obtainable i n 1953-1951+ on the e f f e c t of a complete plus zinc solution upon vitamin C i n ripe tomatoes, the 1952-1953 r e s u l t s indicated that there were no s i g n i f i c a n t e f f e c t s of zinc on vitamin C production. But, i n 1953-1951+, omitting zinc from the complete plus" micronutrients treatment resulted i n a s i g n i f i c a n t increase' of vitamin C. The t o t a l soluble sugar and ash contents of ripe tomatoes were not affected by z i n c . Zinc tended to increase oxidase In tomato leaves. When zinc was omitted from the complete plus micronutrients solution, oxidase a c t i v i t y decreased s i g n i f i c a n t l y . Peroxidase a c t i v i t y and t o t a l plant weight tended to 42 be reduced and s i g n i f i c a n t l y increased, respectively, by the addition of zinc to a complete s o l u t i o n . The ommission of zinc from the complete plus micronutrients treatment produced very s i g n i f i c a n t increases i n the peroxidase a c t i v i t y andvplant weight. I t i s thought that the increases are due to the i n c l u s i o n of boron i n the nutrient s o l u t i o n . CONCLUSIONS The e f f e c t of boron was very noticeable throughout the experimental r e s u l t s . Plants receiving boron were l e s s susceptible to tomato leafmold Infection than plants which received no boron. No s i g n i f i c a n t e f f e c t of boron upon the vitamin C or ash content of ri p e tomatoes occurred. How-ever, the addition of boron to a complete nutrient solution tended to reduce the t o t a l soluble sugar content of ripe tomatoes. Ripe tomatoes exhibited no catalase a c t i v i t y , while immature tomatoes were found to possess a very small amount. On the other hand, tomato leaves proved to be abundant i n catalase. I t was evident that catalase In tomatoes de-creased as the f r u i t approached maturity. Omitting boron showed a tendency to increase catalase i n immature f r u i t s , while adding boron to a nutrient solution s i g n i f i c a n t l y increased catalase i n tomato leaves. The 43 oxidase and peroxidase content of tomato leaves was very s i g n i f i c a n t l y reduced and increased, respectively, when boron was included i n the various nutrient solutions. The tomato y i e l d tended to increase when boron was included in a nutrient solution; however, i n certain instances the y i e l d decreased because other micronutrients, present in the solutions, counteracted the effect of boron and consequently depressed y i e l d . In 1953 - 1954, whenever boron was added to a nutrient solution the plant weight i n -creased very s i g n i f i c a n t l y ; from the addition of boron these same plants produced an abundance of healthy fibrous roots. Copper appeared to decrease the t o t a l soluble sugar and ash in ripe tomatoes, and to increase plant weight and decrease oxidase a c t i v i t y in the leaves. Catalase decreased i n the leaves and green f r u i t of tomatoes when copper was added to a complete nutrient solution. Copper s i g n i f i c a n t l y depressed peroxidase, in the plant leaves, while no s i g n i f i -cant effect upon the f r u i t y i e l d could be determined when copper was added or omitted. The omission of copper from the complete plus micronutrients solution tended to increase vitamin C. The vitamin C content of ripe tomatoes appeared to increase when manganese was added to a complete solution; furthermore, a trend suggested that the ash in ripe f r u i t and catalase i n green f r u i t were s l i g h t l y decreased and i n -creased, respectively, by t h i s treatment. The addition of manganese to a complete nutrient so l u t i o n caused a depress-ing trend upon t o t a l soluble sugar , y i e l d , and t o t a l plant weight; catalase and peroxidase a c t i v i t y i n plant leaves was very s i g n i f i c a n t l y reduced by t h i s treatment. Upon oxidase, the l a t t e r treatment appeared to be i n e f f e c t i v e . The plant weight and t o t a l soluble sugar content of ripe tomatoes, and the catalase action of green f r u i t were s l i g h t l y Increased by molybdenum; the peroxidase enzyme i n -creased s i g n i f i c a n t l y at the one percent l e v e l when molybdenum was added to the complete solution. This treatment also indicated that molybdenum depressed f r u i t y i e l d and s i g n i -f i c a n t l y reduced catalase i n the leaves, whereas vitamin C and oxidase d i d not appear to be influenced by molybdenum. Zinc very s i g n i f i c a n t l y reduced the catalase content of tomato leaves, while a trend Indicated that zinc reduced f r u i t y i e l d and l e a f peroxidase, but increased l e a f oxidase. Plant weight, vitamin C i n ripe f r u i t , and catalase i n green f r u i t tended to increase when zinc was added. There was no apparent e f f e c t of zinc upon sugar or ash i n r i p e tomatoes. SUMMARY The e f f e c t of boron, copper, manganese, molybdenum, and zinc upon the catalase. oxidase, peroxidase, vitamin C, ash, and t o t a l soluble sugar content, and the y i e l d and plant weight of the tomato (Vetomold 121 variety) were studied i n the greenhouse at the University of B r i t i s h Columbia* Plants receiving boron were observed to be l e s s susceptible to tomato leafraold than plants which received no boron. Boron was also observed to stimulate fibrous root growth. Manganese, copper, and zinc tended to increase vitamin C i n ripe tomatoes, while boron d i d not appear to have any e f f e c t . A trend indicated that either boron or manganese depressed t o t a l soluble sugar- i n ripe tomatoes, whereas copper and zinc produced no e f f e c t . Molybdenum was the only micronutrient that appeared to increase the sugar content. A l l micronutrient elements tended to decrease the ash content of ripe tomatoes, p a r t i c u l a r l y molybdenum, but the results were non-significant. Ripe tomatoes were found to be lacking i n catalase, while immature tomatoes possessed very l i t t l e of the enzyme. Hence i t was concluded that catalase decreases as the f r u i t matures. In contrast with the f r u i t , tomato leaves proved to be abundant i n catalase. Boron and copper both s l i g h t l y depressed catalase 4 6 i n green tomato f r u i t s , whereas manganese, molybdenum, and zinc a l l produced an increase. Boron, however, s i g n i f i -cantly Increased the catalase a c t i v i t y of the leaves, but manganese, molybdenum, and zinc s i g n i f i c a n t l y decreased i t . Although significance i s lacking, copper d i d reduce catalase below the control treatment. Boron s i g n i f i c a n t l y decreased oxidase but markedly Increased peroxidase i n tomato leaves. Molybdenum tended to decrease oxidase and to s i g n i f i c a n t l y increase peroxidase. Zinc was the only micronutrient that appeared to increase oxidase, while copper and manganese s i g n i f i c a n t l y decreased peroxidase. The f i v e micronutrients d i d not e f f e c t any r e a l differences i n f r u i t y i e l d , although boron gave the greatest y i e l d while manganese and molybdenum tended to decrease i t . Boron s i g n i f i c a n t l y augmented the t o t a l plant weight, while copper, molybdenum, and zinc showed a tendency to increase i t . On the other hand, manganese indicated a trend to de-press plant weight. k i LITERATURE CITED (1) Alexander, T.R. Anatomical and p h y s i o l o g i c a l r e -sponses of squash to v a r i o u s l e v e l s of boron supply. Bot.G-az. l03:tyZ5-k91. 1 9 k l . (2) Anders sen, P.G-. C h l o r o s i s of deciduous f r u i t t r e e s due t o a copper d e f i c i e n c y . Jour. Pomol. Hort. S c i . 10:130-lk6. 1932. (3) Appleman, CO. Some observations on c a t a l a s e . Bot. Gaz. 50:182-192. 1910. (k) Appleman, D. C a t a l a s e - c h l o r o p h y l l r e l a t i o n s h i p i n b a r l e y s e d d l i h g s . P l a n t P h y s i o l . 27:613-621. 1952. (5) Arnon, D.I. Copper enzymes i n I s o l a t e d c h l o r o p l a s t s . PSlyphenoloxidase i n Beta V u l g a r i s . P l a n t P h y s i o l . 2 k : l - l 5 . 19k9. (6) B a i l e y , L.P., and J.S. McHargue. E f f e c t of boron, copper, manganese, and z i n c on the enzyme a c t -i v i t y of tomato and a l f a l f a p l a n t s grown i n the greenhouse. P l a n t P h y s i o l . 19:105-116. 19kk. (7) : Enzyme a c t i v i t y i n tomato f r u i t s and leaves at d i f f e r e n t stages of development. Amer. Jour. Bot. 30:763-766. 19k3< (8) Bessey, O.A., and C.G. King. The d i s t r i b u t i o n of v i t a m i n C i n p l a n t and animal t i s s u e s and i t s determination. Jour. B i o l . Chem. 103:687-698. 1933. (9) Cook, R.L., and C.E. M i l l a r . The e f f e c t of borax on the y i e l d , appearance, and mineral Composition of spinach and sugar beets. S o i l S c i . Soc. Am. Proc. 5:227-23k. 19k0. (10) D a v i s , W.B. Q u a n t i t a t i v e f i e l d t e s t f o r e s t i m a t i o n of peroxidase. Ind. Eng. Chem. A n a l . Ed. I k : 952-953. 19k2. (11) D m i t r i e v , K.A. The e f f e c t of micro-elements i n the limed podzol s o i l s on the development and the crop of r e d c l o v e r . Pedology (U.S.S.R.), 1939, No. k : l l l+ - l 3 3 . (Chem. Abs. 3k, 705k. 19k0). 48 (12) Drosdoff, M. The use of minor elements. U.S.D.A. Yearbook 1943 - 1947, p. 577 - 582. 1947. (13) Erkama, J . On the e f f e c t of copper and manganese on the i r o n s t a t u s of higher p l a n t s . Trace Elements i n P l a n t Physiology, p. 55. Chronica Botanica Co., Waltham, Mass., U.S.A. 1950, (14) E z e l l , B.D., and J.W. C r i s t . E f f e c t of c e r t a i n n u t r i e n t c o n d i t i o n s on a c t i v i t y of oxidase and c a t a l a s e . Mich. Agr. Exp. St a . Tech. B u l l . 78, May, 1927. (15) G i l y a r o v s k i i , I.P., and I.S. Chernov. The e f f e c t o f boron on the increase of the tomato and cucumber crops. Ovoshchevodstvo (U.S.S.R.) 1940, No. 11-12, 28-29. (Chem*, Abs. 35:5934* 194U. (16) Gum, O.B., H.D. Brown, and R.C. B u r r e l l . Some e f f e c t s of boron and manganese on the q u a l i t y o f beets and tomatoes. P l a n t P h y s i o l . 20:267-275. 1945. (17) Gustafson, P.G., et a l . Catalase a c t i v i t y i n tomato f r u i t s at d i f f e r e n t stages of t h e i r development. P l a n t P h y s i o l . 7:155-160. 1932. (18) H a r r i s , G.H. Some e f f e c t s of micro-elements on growth and storage of c a r r o t s and t u r n i p s . Proc. Amer. Soc. Hort. S c i . 43:219-224. 1943. (19) H e i n i c k e , A.J. Catalase a c t i v i t y as an I n d i c a t o r of the n u t r i t i v e c o n d i t i o n of f r u i t t r e e t i s s u e s . Proc. Amer. Soc. Hort". S c i . 22:209-214. 1922. (20) Catalase a c t i v i t y i n dormant apple twigs; i t s r e l a t i o n to the c o n d i t i o n s of the t i s s u e , r e s p i r a t i o n , and other f a c t o r s . C o r n e l l Univ. Agr. Exp. St a . Mem. 74. 1923. (21) Factors i n f l u e n c i n g c a t a l a s e a c t i v i t y i n ap p l e - l e a f t i s s u e . C o r n e l l Univ. Agr. Exp. S t a . Mem. 62. 1923. (22) Seasonal v a r i a t i o n i n the n u t r i e n t c o n d i t i o n of apple t r e e s as i n d i c a t e d by c a t a l a s e a c t i v i t y . Proc. Amer. Soc. Hort. S c i . 25:234-239. 1928. £9 (23) Hivon, K.J., D.M. Doty, and F.W. Quackenbush. Ascorbic acid and ascorbic-acid-oxidizing enzymes of manganese-deficient soybean plants grown i n the f i e l d . S o i l S c i ; 71:353-359. 1951. (2k) Johnston, E.S., and W.H. Dore. The infleunce of boron on the chemical composition and growth of the tomato plant. Plant Physiol. k:31-62. 1929. (25) Keese, H. Action of boron and manganese on plant growth with special consideration of the effect of liming. Bodenkunde u. Pflanzenernahr. 27:116-13k. 19k2. (26) KLein, R.M. The r e l a t i o n of gas exchange and tyrosinase a c t i v i t y of tomato tissues to the l e v e l of boron n u t r i t i o n of plants. Arch. Biochem.30:207-21k. 1951. (27) Landon, R.H. The effect of cer t a i n chemicals on the catalase a c t i v i t y i n plants. Amer. Jour. Bot. 21:583-591. 193k. (28) Loustalot, A.J., P.W. Burrows, S.G. G i l b e r t , and A. Nason. Ef f e c t of copper and zinc d e f i c i e n -cies on the photosynthetic a c t i v i t y of the foliage of young tung trees. Plant P h y s i o l . 20:283-288. 19k5. (29) MacVicar, R., and R.H. B u r r i s . The r e l a t i o n of boron to ce r t a i n plant oxidases. Arch. Biochem. 17:31-39. 19k8. (30) McHargue, J.S. The role of manganese i n plants. Jour. Am. Chem. Soc. kk : l 5 9 2 - l 5 9 8 . 1922. (31) Muhr, G.R. Plant symptoms of boron deficiency and the effects of borax on the y i e l d and chemical composition of several crops. S o i l S i c . 5k:55-65« 19k2. (32) Mulder, E.G. Importance of copper and molybdenum i n the n u t r i t i o n of higher plants and micro-organisms. Trace Elements i n Plant Physiology, p. kk. Chronica Botanica Co., Waltham, Mass., U.S.A. 19£0. (33) Nason, A l v i n . Metabolism of micronutrient elements i n higher plants. 11. Effect of copper deficiency on the i s o c i t r i c enzyme i n tomato leaves. Jour. B i o l . Chem. 198:6k3-£53. 1952. 50 (34) Paterson, D.D. Statistical Technique i n Agricultural Research. McGraw-Hill Book Co., Inc., New York, U.S.A. 1939. (35) Reed, G.B. The relation between oxidase and catalase in plant tissues. Bot. Gaz. 62:409-412. 1916. (36) Rodriquez, A.G. Effect of boron on the ava i l a b i l i t y of iron, abst. Amer. Jour. Bot. 20:679. 1933. (37) Saru, Natalie. The effect of boron on sugar beets. Kuhn-Arch. 48:1-51. 1939. (Chem. Abs. 34,568. 1940). (38) Scott, L.E., and E.P. Walls. Ascorbic acid content and sugar-acid ratios of fresh f r u i t and processed juice of tomato varieties. Proc. Amer. Soc. Hort. Sci. 50:269-272. 1947. (39) Shear, C.B., and H.L. Crane. Nutrient - element balance. U.S.D.A. Yearbook 1943-1947. pp. 592-601. 1947. (40) Vlasyuk, P.A. The importance of manganese in the u t i l i z a t i o n of the ammonia and nitrate forms of nitrogen for the water culture of sugar beets. Compt. rend. acad. s c i . U.S.S.R. 28:184-186. 1940. (Chem. Abs. 35, 2564. 194L). (41) Wallace, T. Trace Elements in Plant Physiology, p . v i i i of preface. Chronica Botanica Co., Waltham, Mass. 1950. 51 APPENDIX ILLUSTRATION OP CALCULATION OP STATISTICAL-..ANALYSIS: v, e.g. LEAP CATALASE ( 1 9 5 3 - 1 9 5 4 ) Total S.S. . . . 3 4 - 2 2 • 4 8 = 252.5 . . . 3 6 3 . 0 2 - 1446.4 2 1 2 4 » = 10.1|. Treatments S.S. = 1 2 0 . 8 2 + 1 3 0 . 5 2 + . . . - 1 4 4 6 . 4 2 4 4 8 = 1 8 6 . 6 ANALYSIS OP VARIANCE FOR LEAP CATALASE factor S.S. degrees variance P value freedom calculated table ( . 0 5 ) ( . 0 1 ) total 2 5 2 . 5 4 7 treatments 1 8 6 . 6 1 1 1 6 . 9 6 10.1044 2 . 1 0 2 . 8 4 blocks 1 0 . 4 3 3 . 4 7 2 . 0 7 2 . 8 9 4 . 4 4 error 5 5 . 5 3 3 1 . 6 8 S.D. between treatment means at . 0 5 level = .•/1.6B x 2 x 2.03 = 111 9 52 S.D. between treatment means at .01 l e v e l = / 1.68 x 2 x 2.73 = 2 ^ 

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