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Some effects of urea and nitrate nitrogen on the growth and composition of cranberry. Leschyson, Margaret Ann 1969

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SOME EFFECTS OF UREA AND NITRATE NITROGEN ON THE GROWTH AND COMPOSITION OF CRANBERRY by MARGARET ANN LESCHYSON B.Sc, University of British Columbia, 1967 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department of Plant Science We accept this thesis as conforming to the required standard THE UNIVERSITY OF April, BRITISH COLUMBIA 1969 i ABSTRACT Two similar greenhouse experiments wore carried out to observe the effects of 2 forms of nitrogen f e r t i l i z e r , (nitrate and urea), each at .5 rates, ( 0 , 30, 60, 90, and 120 lb N per acre), on cranberry plants, (Vaccinium macrocarpon Ait. cv. McFarlin). In the f i r s t experiment, treatments were applied to cuttings which had been rooted for a short time whereas in the second experiment, cuttings which had been rooted for 7 months as well as cuttings which had just been rooted were used. Growth measurements and foli a r mineral analyses were carried out on shoots collected from actively growing plants 2 and 16 weeks after differential treatment in the f i r s t experiment and 14 weeks after treatment in the second experiment. In both experiments, N f e r t i l i z a t i o n soon after rooting enhanced vegetative growth. Greater vegetative growth was obtained with nitrate in the f i r s t experiment but with urea in the second. Analysis of variance indicated that treatments which increased growth also increased f o l i a r N and decreased foli a r P, Fe, and Ca. In the f i r s t experiment, growth measurements were linearly correlated with foli a r Mg or Mn or both in the f i r s t harvest, but in the second harvest, cor-relations were with foli a r N, P, and K content. i i In the second experiment, cuttings which had been rooted for some time initiated more shoots and leaves, but did not respond to N treatments. i i i In presenting th i s thes is in p a r t i a l fu l f i lment of the requirements for an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree that the L i b r a r y s h a l l make i t f r e e l y ava i lab le for reference and study. I further agree that permission for extensive copying of t h i s thes i s for scho lar ly purposes may be granted by the head of my Department or by h is representat ives . It i s understood that copying or publ i ca t ion of th i s thes i s for f i n a n c i a l gain s h a l l not be allowed without my wri t ten permiss ion. Department of Plant Science U n i v e r s i t y of B r i t i s h Columbia Vancouver 8, Canada A p r i l , 1969 ACKNOWLEDGMENTS I wish to thank Dr. G.W. Eaton, A s s o c i a t e P r o f e s s o r Department o f P l a n t S c i e n c e , U n i v e r s i t y of B r i t i s h Columbi under whose s u p e r v i s i o n t h i s p r o j e c t was undertaken, f o r h i s t e c h n i c a l a d v i c e d u r i n g the r e s e a r c h , and f o r h i s guidance i n the p r e p a r a t i o n o f t h i s t h e s i s . G r a t e f u l acknowledgment i s a l s o extended to the ot h e r members of my t h e s i s committee: Dr. V.C. B r i n k , Department of P l a n t Science Dr. L.E. Lowe, Department of S o i l S c i e n c e Dr. D.P. Ormrod, Department of P l a n t Science Dr. A. S t o r r , Department of Chemistry I a l s o wish to thank the B i g Red Cranberry Company, and the Northern Peat Company f o r p r o v i d i n g the p l a n t s used i n t h i s p r o j e c t . The r e s e a r c h was supported i n part by the CD.A. Op e r a t i n g Grant No. 48 awarded to Dr. G.W. Eaton, and a U n i v e r s i t y o f B r i t i s h Columbia Graduate F e l l o w s h i p . V TABLE OF CONTENTS Page I. INTRODUCTION 1 I I . LITERATURE REVIEW 2 A. N i t r o g e n F e r t i l i z e r s 2 B. P l a n t T i s s u e A n a l y s i s 10 I I I . MATERIALS AND METHODS 14 IV. RESULTS 21 A. E f f e c t s o f Treatments on Growth 21 1. Fresh and Dry Weights 21 2. Shoot Number and Shoot Length 25 3. Leaf Number 28 4. Leaf Area 28 B. E f f e c t s of Treatments on Peat Leachate 31 1. pH 31 2. Ammonium .31 3. N i t r o g e n 31 C. E f f e c t s of Treatments on F o l i a r N u t r i e n t Composition 34 1. N i t r o g e n 36 2. Phosphorus 36 3. Potassium 40 4. Calcium 40 5. Magnesium 42 6. Iron 42 7. Manganese 42 D . M u l t i p l e Regression Analyses 45 DISCUSSION A. Growth 51 B . Minera l Analyses 53 1. Nitrogen 53 2. Phosphorus 55 3. Potassium 55 4. Calcium 56 5. Magnesium 57 6. Iron 58 7. Manganese 58 8. General 59 SUMMARY . . BIBLIOGRAPHY v i i LIST OF TABLES Page Tabl e 1. A n a l y s i s of v a r i a n c e models f o r growth r e c o r d s and chemical analyses 20 Tabl e 2. E f f e c t s of N source on growth i n experiment 1 22 Ta b l e 3. E f f e c t o f r a t e o f a p p l i e d N on s o i l parameters i n experiment 1 34 Tabl e 4. Grand means f o r f o l i a r m i n e r a l l e v e l s i n the three h a r v e s t s (ppm) 35 T a b l e 5 . E f f e c t s o f r a t e s of a p p l i e d N on f o l i a r m i n e r a l composition (ppm) 38 T a b l e 6. E f f e c t s of sources of a p p l i e d N on f o l i a r m i n e r a l composition (ppm) 39 Ta b l e 7. L i s t o f v a r i a b l e s used f o r m u l t i p l e r e g r e s s i o n a n a l y s i s f o r experiment 1 46 T a b l e 8. S i g n i f i c a n t c o r r e l a t i o n s between growth measurements and m i n e r a l a n a l y s e s at f i r s t h a r v e s t , (experiment l ) 46 T a b l e 9. S i g n i f i c a n t c o r r e l a t i o n s among m i n e r a l a n a l y s e s i n the second harvest and growth and m i n e r a l a n a l y s e s i n the f i r s t h a r v e s t , (experiment l ) 47 T a b l e 10. S i g n i f i c a n t c o r r e l a t i o n s between growth measurements be f o r e second harvest and a l l o ther measurements made i n the f i r s t experiment 47 T a b l e 11* S i g n i f i c a n t c o r r e l a t i o n s from m u l t i p l e r e g r e s s i o n i n experiment 2 49 T a b l e 12> S i g n i f i c a n t l i n e a r c o r r e l a t i o n s i n .\ V - - - experiment 2 49 v i i i LIST OF FIGURES F i g u r e 1. F i g u r e 2. F i g u r e 3. F i g u r e 4. F i g u r e 5. F i g u r e 6. F i g u r e 7. F i g u r e 8. F i g u r e 9 . F i g u r e 10. F i g u r e 11. F i g u r e 12. Page E f f e c t of source and r a t e of N and age of c u t t i n g s on f r e s h weight 23 E f f e c t of source and r a t e of N and age of c u t t i n g s on dry weight 24 E f f e c t of age of c u t t i n g s and N source on shoot l e n g t h 26 E f f e c t of age of c u t t i n g s and r a t e of a p p l i e d N on shoot l e n g t h 27 E f f e c t of age of c u t t i n g s and r a t e of a p p l i e d N on shoot number 29 E f f e c t of age of c u t t i n g s and N source on l e a f number 30 E f f e c t o f source and r a t e o f a p p l i e d N on peat e f f l u e n t pH 32 E f f e c t o f source and r a t e of a p p l i e d N on peat e f f l u e n t N 33 E f f e c t of source and r a t e o f a p p l i e d N on f o l i a r N 37 E f f e c t o f source and r a t e of a p p l i e d N on f o l i a r Ca 41 E f f e c t of source and r a t e o f a p p l i e d N on f o l i a r Mg 43 E f f e c t of source and r a t e of a p p l i e d N on f o l i a r Mn 44 I. INTRODUCTION The c r a n b e r r y , (Vaccinium macrocarpon A i t . ) , i s an E r i c a c e o u s v i n e whose crop i s borne on u p r i g h t s , ( v e r t i c a l s h o o t s ) , a r i s i n g from long v e g e t a t i v e runners which t r a i l along the ground. The p l a n t i s propagated from runner c u t t i n g s and grown i n a c i d i c s o i l . In B r i t i s h Columbia, commercial c r a n b e r r y p l a n t i n g s are e s t a b l i s h e d on peat bogs, but much o f the l i t e r a t u r e d e a l s w i t h p l a n t s grown i n s o i l s low i n or g a n i c matter. The v a r i e t y M c F a r l i n i s d e s c r i b e d by Crowley (14) as a high y e i l d i n g commercial v a r i e t y with b e r r i e s of good keeping q u a l i t y though l e s s a t t r a c t i v e i n c o l o r than some other v a r i e t i e s . V e g e t a t i v e growth i s a s s o c i a t e d with N metabolism, and i n e s t a b l i s h i n g a new p l a n t i n g i t i s d e s i r a b l e to o b t a i n maximum v e g e t a t i v e growth f o r the f i r s t 2 to 3 y e a r s . The o b j e c t o f the present study was to apply N as urea and n i t r a t e , each at d i f f e r e n t l e v e l s , to Vaccinium  macrocarpon A i t . cv. M c F a r l i n i n peat s o i l i n a greenhouse experiment. V a r i a b i l i t y i n growth and m i n e r a l composition of the t i s s u e were then e v a l u a t e d and c o r r e l a t e d to pr o v i d e c r i t e r i a d i a g n o s t i c of the balance among t i s s u e c o n d i t i o n s which i s optimum f o r v e g e t a t i v e growth. 2 I I . LITERATURE REVIEW A. N i t r o g e n F e r t i l i z e r s N a t u r a l i n c r e a s e s i n s o i l N from the breakdown of s o i l o r g a n i c matter by m i n e r a l i z a t i o n to n i t r a t e , d i r e c t f i x a t i o n by organisms, and capture o f N compounds by r a i n f a l l are o f f s e t by c o n d i t i o n s u n f a v o r a b l e to the f u n c t i o n of a e r o b i c organisms such as low temperatures and c o n s t a n t l y h i g h water t a b l e , and by l o s s e s due to d e n i t r i f i c a t i b n by anaerobic organisms, l e a c h i n g , e r o s i o n , and weed c o m p e t i t i o n (15). Dana (15) t h e r e f o r e recommends N a p p l i c a t i o n s e a r l y i n the season before m i n e r a l i z a t i o n can c o n t r i b u t e s i g n i f i c a n t l y to a v a i l a b l e s o i l N, and d u r i n g the summer to ensure a h i g h l e v e l o f a v a i l a b i l i t y to c r a n b e r r y p l a n t s . I t has been advised (5) that a complete NPK f e r t i l i z e r may cause a severe weed problem, e s p e c i a l l y c l o v e r , i n some c r a n b e r r y bogs, but that N alone gave e x c e l l e n t v i n e growth without accompanying i n c r e a s e i n the c l o v e r p o p u l a t i o n . Most of the l i t e r a t u r e (41) d e a l s w i t h b e a r i n g p l a n t i n g s and f o r t h i s reason a d v i s e s moderation i n the use of N f e r t i l i z e r s s i n c e abundant and r e a d i l y a v a i l a b l e N i n c r e a s e s y i e l d but impairs keeping q u a l i t y o f the f r u i t . Addoms and Mounce ( l ) r e p o r t e d a s y m b i o t i c , endophytic fungus, Phoma r a d i c i s . a s s o c i a t e d with the c r a n b e r r y p l a n t . The mycorhiza were more abundant i n s o i l s of h i g h N and o r g a n i c matter content, and t h e i r presence appeared to enable d i r e c t a b s o r p t i o n by the c r a n b e r r y p l a n t of N i n the NH^ form. In a greenhouse sand c u l t u r e experiment ( l ) , c r a n b e r r y p l a n t s were s u p p l i e d with (NH^^SO^, CafNO^^, and no N. The Nrf£ form induced the best v e g e t a t i v e response i n terms of runner growth though both NH^ and low r a t e s of NOg produced b e t t e r growth than the c o n t r o l , In two a d d i t i o n a l experiments, NO^, NH^, d r i e d blood, and amino a c i d s were added to pots p r e v i o u s l y r e c e i v i n g no N. W i t h i n a week, p l a n t s s u p p l i e d w i t h d r i e d blood, NH^, and amino a c i d s responded f a v o r a b l y i n terms of v i n e growth. The sand was t e s t e d f o r amino a c i d s , NH*, and NO^ i n the d r i e d blood treatment. No NO^ was found, but t h e r e was an abundance of amino a c i d s , and a s m a l l amount of NH^. I t was thought that perhaps NO^ may have been absorbed but not a s s i m i l a t e d at low temperatures, but a chemical a n a l y s i s c a r r i e d out on the c r a n b e r r y t i s s u e showed no NO^ or NO^ present even i n p l a n t s p l a c e d i n a low temperature environment. 4 The question was posed whether the mycorhiza were capable of N f i x a t i o n , but i t was concluded that i f so, the amount was inadequate to provide s u f f i c i e n t N to the host since the control plants showed very poor growth. In a further experiment, Addoms and Mounce (2) found that, in sand culture, NO^ gave the most favorable vine growth under acid conditions, and NH^ under neutral or a l k a l i n e conditions, and that in bog s o i l , (considered a buffered condition), NH* may be used at higher pH's than in sand. Of Ca(N0 3) 2, (NH 4) 2S0 4, and glycine in sand medium at pH 4, 6 , and 8, plants supplied with glycine and NOg at pH 8, and NH^ at pH 4 were unsuccessful and ultimately died. Treatments of glycine at pH 4, and NH* at pH 6 and 8 were the most successful, giving the greatest amount of stem growth and best general appearance. Glycine at pH 8 also produced s a t i s f a c t o r y vine growth. NO^ at pH 4 and 6 produced plants which were apparently healthy but produced less growth than the successful NH* and glycine cultures. In s o i l culture, NH^ with a small amount of lime produced the best vegetative growth. Both NH^ 5 with no lime and NO^  with a small amount of lime were satisfactory. However, NH^  and NO^  with a large amount of lime and NO^  with a small amount of lime showed less vigorous growth. In these experiments, i t was found also that amounts of ether soluble o i l s in the leaf blades were inversely related to the vigor of vegetative growth. No NO^  was found in any of the plant material. NH^  was found in plants supplied with glycine. Larger amounts of <<-amino N were found in plants supplied with NH^  than NO^  indicating rapid "conversion" within the plant. Furthermore no NH^  was found in the s o i l solution of NO^  or glycine treated pots. The NH* ion applied as urea is a readily available source of N, but according to DeLong (17) is a highly leachable form especially in acid soils since i t cannot replace the H + ion which is more strongly bound by s o i l colloids. Kender and Childers (29) investigated the effect on the growth of cranberry vines of urea-formaldehyde, (UF), a more slowly available source of N less susceptible to leaching and compared i t 6 w i t h urea, ( N H 4 ) 2 S 0 4 and C a ( N 0 3 ) 2 i n a greenhouse experiment i n bog s o i l (of which the o r g a n i c content was .not r e p o r t e d ) . UF produced f a v o r a b l e v e g e t a t i v e growth throughout the experiment and a f t e r 3 months t h i s growth was s i g n i f i c a n t l y g r e a t e r than i n the c o n t r o l , urea, or ( N H 4 ) 2 S 0 4 treatments, the c r i t e r i a b e i n g t o t a l l e n g t h of runners, dry weight of a e r i a l p a r t s , and l e n g t h of u p r i g h t s . Runner growth of UF t r e a t e d p l a n t s was not s i g n i f i c a n t l y g r e a t e r than that o f p l a n t s t r e a t e d w i t h C a ( N 0 3 ) 2 » but f r e s h weight of the p l a n t m a t e r i a l was s i g n i f i c a n t l y g r e a t e r than the c o n t r o l . P l a n t s t r e a t e d w i t h ( N H 4 ) 2 S 0 4 d i d not d i f f e r s i g n i f i c a n t l y front the c o n t r o l s i n producing v e g e t a t i v e growth. I t was thought that s i n c e UF a p p l i c a t i o n s produced, f a v o r a b l e v e g e t a t i v e growth and was s l o w l y a v a i l a b l e to the p l a n t , one a p p l i c a t i o n per season would be adequate to f u l f i l the N requirement without c a u s i n g an o v e r s u p p l y of N to which the c r a n b e r r y p l a n t i s s e n s i t i v e (29). Somogyi et a l . (39), i n a 3-year greenhouse experiment i n v e s t i g a t e d the growth produced by 4 sources o f N: urea, UF, ( N H 4 ) 2 S Q 4 » and a mixture of equal p r o p o r t i o n s of urea and UF, i n s o i l s of 0, 2, and 4% o r g a n i c matter. 7 The results of the experiment showed that (NH^^SO^ produced greater vegetative growth in the sand substrate than did organic N sources. In the media containing organic matter, the organic sources of N produced more favorable vegetative growth. In particular, the combination of urea and UF was more effective than either urea or UF applied alone. In fact, applied alone urea or UF were less effective than the (NH^^SO^ treatment in terms of total runner growth and dry weight in the 2% organic so i l and UF was less effective in the sand culture. There were no significant treatment differences in runner growth in the 4% organic soils. The conclusion drawn was that the presence of microbial activity in media containing organic matter made organic sources of N available to the cranberry plant. In order to place these results in proper perspective, i t should be mentioned that total runner growth, dry weight, and number of uprights was greater for the urea + UF treatment in media containing organic matter than for (Nr^^SC^ in sand culture. Leaf N content was higher in plants receiving organic N because of i t s sustained availability. The urea and urea + UF treatment produced higher leaf N in plants grown in soils containing organic matter. 8 There i s disagreement in the l i t e r a t u r e regarding the use of (NH 4) 2S0 4 as a source of N for cranberries. Fisher ( 2 4 ) reported that (NI-^^SO^ i s the most widely used f e r t i l i z e r in correcting poor vine growth, and Crowley (14) noted that N deficiency was quickly a l l e v i a t e d by (NH 4) 2S0 4 applications. On the other hand, Eaton (21) regarded (NH 4) 2S0 4 as injurious and Beckwith (7) stated that (NH 4) 2S0 4 applied alone was an undesirable source of N. No experimental data were presented in support of the above statements. In the greenhouse experiment in bog medium, discussed above, Kender and Childers (29) found that (NH 4) 2S0 4 produced growth response not s i g n i f i c a n t l y d i f f e r e n t from that of the control cranberry plants. There i s also disagreement regarding the use of NO^ as a source of N for cranberries. Eaton (21) recommended NaNO^ on hard bottom bogs. Beckwith (7), in a f i e l d experiment on savannah s o i l (hard bottom or sand mixed with dark s i l t ) , comparing NaNO^ and dried blood alone and in combination at 20 and 30 lb N per acre found that NaN03 at 30 lb N per acre induced the best vine growth in the f i r s t season. Vine growth was stimulated within 1. week of application of NaNO-. Co n t r a r i l y , Addoms and Mounce (l) found that 9 NOg was an acceptable form of N for use in sand culture at low pH's only, but was s t i l l inferior to the NH^  form, with regard to vine growth. Herath and Eaton (28), in a greenhouse experiment found that, in peat medium, highbush blueberry preferred NH^ over NO^  as a source of N, based on leaf N content and growth measurements, although NO^  may produce satis-factory growth on acidic peat soils since i t may be converted to the NH^  form by s o i l microorganisms. Furthermore, Eaton (21) and Morse (33) reported that no f e r t i l i z e r was required on peat or muck, and Morse (33) recommended that vine growth be obtained by holding winter flood late rather than f e r t i l i z i n g for vine growth. It must be assumed that the above recom-mendations were based on experience, and observation of practical situations since no experimental procedures were implied or data presented. Contradictory recommendations regarding the form of N to be applied to cranberry plants may arise from different s o i l types, s o i l pH's, types of plantings (bearing and non-bearing), or perhaps unsuitable experimental designs. With regard to rates and timing of cranberry f e r t i l i z e r applications, Sorensen (40) recommended 10 20 lb N per acre (form not specified) applied after 1 week of new growth. Chandler and Colby ( l l ) found that 40 lb N per acre for McFarlin was satisfactory and that 80 lb annually was too great. Dana (15) recommended 48 lb N per acre applied as NH^ NOg as a split application for vines in poor condition, 35 lb on medium vigor vines, and 16 lb N per acre in a single application on f a i r l y vigorous vines. On new plantings, 20 lb N per acre every 2 to 3 weeks in the summer months was recommended, and on peat soils, 2 applications, (in May and June), would suffice. Again, no experimental data were presented in the above articles in support of the recommendations. B. Plant Tissue Analysis Leaf tissue analysis is supported by Reuther and Smith (36) and Bould (8), as a method of nutritional evaluation for orchard trees, based on studies with citrus. It is compared to other methods such as visual diagnosis and s o i l analysis. It is reasoned that leaves act as a reservoir for carbohydrates and minerals which influence the efficiency of C assimilation through photosynthesis, and therefore best reflect, though in a general way, the nutrient status of the entire plant. Tissue analysis delineates nutritional problems distinct from environmental ones since i t allows evaluation of 11 the nature and causes of certain responses. Tissue analysis also allows results to be extrapolated from experimental to practical situations. Although leaf analysis is admittedly only a guide in nutrient evaluation, certain limitations exist, such as the inability to establish absolute standards of excess, optimum, and deficient ranges of concentration in the plant. A visual analysis is limited in that symptoms often must be severe before a nutrient deficiency is detected, and s o i l analysis is considered by Reuther and Smith (36) as of limited value since available nutrients differ from those which are chemically extracted from the s o i l samples. Ion antagonism, so i l pH, moisture, aeration, microorganisms, and chemical form of elements in the s o i l a l l affect availability of nutrients to the plant. Medappa (31) in an experiment using cranberry plants in solution cultures of pH 3, 4, 5, 6, 7, and 8 with different levels of P and Ca, and 2 sources of Fe (FeSO^ and FeEDTA), found differences in growth and mineral composition of the plant tissue. Low levels of Ca, P and Fe and high levels of Mg were found at pH 3. P content decreased above pH 6 and the decrease was marked at pH 8. Ca content increased with pH to pH 7 then dropped at pH 8 in the plants supplied with FeEDTA. Mg levels decreased after pH 3 and again 12 increased at pH 7 and 8. At pH levels 3, 4, and 5, there was no difference in vine growth between plants supplied with FeS04 or FeEDTA though above pH 6 FeEDTA produced better growth. Chlorosis was evident in plants supplied with FeSO^ above pH 6, Variation in P (15, 30, and 60 ppm)'and Ca (80, 200, and 300 ppm) induced l i t t l e difference in plant growth compared with those induced by different pH levels. Growth appeared to have improved however, as a result of high P supply, and chelated Fe at pH 4. At pH 5, FeSO^ treated plants showed increased growth with increased Ca supply. No differences in root growth were detected, as a result of varied P and Ca supply. In the same experiment, Medappa (31) showed that optimum growth, in terms of fresh and dry weight of tops, occurred in cultures of pH 5 and 6. Kender and Childers (29) in a greenhouse experiment with bog s o i l of undefined organic content reported that (NI-^^SG^ and urea did not cause s o i l pH to rise above pH 6. Hall et al (26) in a greenhouse study of blue-berries found that growth, pH levels of media and tissue N, P, K, Mn, and Mg levels could be correlated significantly. In experiments with citrus, Reuther and Smith (36) considered N the most important and overriding of. a l l 13 elements in nutritional evaluation and analytical interpretation since i t strongly influences the levels of other elements, especially P, S, K, and Mg. High N levels in general are indicated by dark green foliage and indicate low levels of P, S, and K, but high levels of Mg. Assuming optimum levels of N, high f o l i a r K is generally accompanied by low Mg, Ca, and one or more heavy metals. High Ca levels may reflect low levels of heavy metals and major bases. Low Ca, on the other hand may be accompanied by toxic proportions of Al, Fe, and Mn and usually reflects acid s o i l unfavorable for growth. Such conditions are commonly accompanied by low n i t r i f i -cation rates. Somogyi et al. (39) reported typical levels of leaf N as 0.64 to 1.05% of dry weight in greenhouse cultures, and 0.8 to 1.0% in a commercial survey, those having less than 0.8% showing visual N deficiency symptoms described as light green to reddish colored leaves. It was also found that the cranberry leaves contained approximately twice as much N as stems. 14 III. MATERIALS AND METHODS Runners of the variety McFarlin were collected from a small area of a commercial cranberry planting on January 15, 1968. On January 16 and 17, approximately 720 6-inch cuttings were made and 120 stuck about 2.5 inches into each of six 12 X 18 inch flats of peat obtained from the same bale, then placed under mist in the greenhouse for 4 weeks. The misting system was set to operate for 1 minute every 20 minutes. Peat from several bales was mixed in a wheel-barrow, and as 2 to 3 pots were f i l l e d , more peat was added to the wheelbarrow from each of the different bales, and the contents mixed thoroughly. The f i r s t "Beacon" plastic pot 10 inches deep and 7.5 inches in diameter was f i l l e d with peat and placed on a double pan balance, the other 59 pots were subsequently f i l l e d with peat to balance the f i r s t . It was found later that the depth of peat in the pots varied. Each pot was provided with 1 hole 1 cm in diameter at the bottom of the side. The pots were placed in 4 rows of 15 on a greenhouse bench. On February 13, 1968, 600 of the rooted cuttings were transplanted into the prepared plastic pots. Ten holes in rows of 3, 4, and 3 respectively were made in 15 the peat and the cuttings transferred into these holes by scooping each out of the f l a t , row by row and with-out shaking the peat from the roots. On March 4, 4 banks of cool white fluorescent lights consisting of 8 tubes were set 11 inches above the pots and timed to give a 12 hour photoperiod, extended to 21 hours on March 13. Differential f e r t i l i z e r treatments were applied on March 15. Two f e r t i l i z e r s , (Nr^)2^0 and NaNO^ were applied in solution each at 5 levels, consisting of 0, 30, 60, 90, and 120 lb actual N per acre on a surface area basis. The solution was delivered by means of a 25 ml pipette, one aliquot being equivalent to 30 lb NaNO^-N or (Nr^^CO-N per acre. Two, 3, and 4 aliquots were delivered to provide 60, 90, and 120 lb N per acre. Each of the 10 treatments was replicated 5 times. I n i t i a l measurements of shoot number and total shoot length were taken on March 14 for each plant. Subsequent similar measurements were taken on March 21 and 28. On March 21, and every 7 to 10 days thereafter, the 50 pots were rearranged completely at random on the bench top. The plants were watered at f i r s t with tap water. 16 but later i t was decided to use d i s t i l l e d water because of possible effects due to the chloride ion in tap water. A l l pots were given approximately equal quantities of water until the end of May when i t was necessary to supply more water to the larger plants since i t was clear that the s o i l was very much drier and the pots lighter in weight. On March 29. a l l new shoots were cut back leaving 3 fu l l y expanded leaves on each. The shoots from each pot were weighed and placed in a drying oven at 105°C for 48 hours, and the dry weights determined. The dried leaves were separated from the stems and ground to a fine powder with a porcelain mortar and pestle for 10 to 15 minutes. The powdered sample was stored in glass jars with bakelite lids which were opened in a drying oven at 105°C for 48 hours before use in chemical analyses; no moisture corrections were considered necessary. Total N was determined using the semi-micro kjeldahl method described by Chapman and Pratt (12). No replicate determinations were done since previous analyses indicated negligible variance. Mineral extracts were made using the wet ash 17 method described by Chapman and Pratt (12). The extracts were stored in "Nalgene" containers and used to determine Ca, Mg, Fe, and Mn by atomic absorption flame photometry using an Evans Electro-selenium Model 140 Atomic Absorption flame spectro-photometer. The extract was also used to determine K by emission using the Model 140 EEL photometer supplied with an emission attachment, and P by the colorimetric method described by Dickman and Bray (18). Replicate analyses indicated that analytical variation was negligible. On May 8, s o i l effluent solution was collected from each pot in a beaker placed beneath the drain hole. pH was determined on each sample using a Beckman Zeromatic pH meter. Ammonia content was determined by d i s t i l l a t i o n of 10 ml of solution, and total N was determined by the semi-micro kjeldahl method (12). On May 17, length and width measurements were made on 2 leaves chosen from any midshoot or midshoots of 1 plant chosen at random from each pot. Area for each leaf was calculated using the following formula: AREA = 0.3054 + 0.7708 (LENGTH X WIDTH). . This formula was obtained using linear regression techniques on previous data for 18 randomly chosen 18 leaves whose length and width were measured and whose areas were found by cutting out and weighing the areas of developed ozalid paper left unexposed by leaves. On May 21, leaves were counted on each plant, not including the rosette of leaves at the tip of each shoot. Analysis of variance was carried out on a l l data. Duncan's New Multiple Range Test (30) was applied at the 5% level to results which were s i g n i f i -cant according to the analysis of variance. A second experiment was carried out using materials and procedures similar to those in the f i r s t experiment. The cuttings Used, however, were of 2 kinds. The "old" cuttings were collected from the f i e l d on November 11, 1967, kept in a refrigerator u n t i l November 14, and cut into 6-inch lengths and stuck in fl a t s of peat in a misting bed. The "new" cuttings were collected, cut into 6-inch lengths, and stuck into flats of peat in a misting bed on June 17, 1968. Two "old" and 4 "new" rooted cuttings were transferred to plastic pots on July 12, this time shaking the peat from the roots as much as possible. Wet peat for these pots was prepared by adding water directly to the pots while the holes were plugged 19 with rubber stoppers. D i f f e r e n t i a l N treatments were appl ied as before, on J u l y 19, 1968. Again, pots were randomized every 7 to 10 days and watering was done when necessary using d i s t i l l e d water. S o i l e f f luent samples were obtained on November 12 and pH and t o t a l N content determined. On November 4, a l l new shoots were cut o f f next to the o r i g i n a l c u t t i n g . Fresh weight, shoot length, shoot number and leaf number were determined separately for o ld and new plants in each pot and l a t e r averaged for each kind of p lan t . A l l o ld cut t ings survived, but about 6 new cutt ings d i ed , (never more than one per pot ) . F o l i a r N, P, K, Ca, Mg, Fe, and Mn were determined as before, pooling the new and o ld leaves from each pot in order to obtain s u f f i c i e n t mater ia l for a n a l y s i s . Analyses of variance were carr i ed out on a l l data , and the models for these are given in Table 1. Duncan's New M u l t i p l e Range Test was applied at the 5% l e v e l to r e s u l t s . s ign i f i cant in the analys i s of var iance . M u l t i p l e regress ion was c a r r i e d out on the data for f o l i a r mineral content and growth in both experiments. 20 Table 1. Analysis of variance models for growth records and chemical analyses SOURCE OF VARIATION Source Rate SxR Error TOTAL DF 1 4 4 40 49 DATA TESTED A l l s o i l data ( N , NH3, pH) Experiments 1 and 2 A l l mineral analysis data Experiments 1 and 2 Shoot number and length Experiment 1 Fresh and dry weights Experiment 1 Source 1 Rate 4 SxR 4 Pots 40 Leaf area Plants 450 Experiment 1 Error 500 TOTAL 999 Source Rate SxR Pot/SR Age AxR AxS AxRxS Error 1 4 4 45 1 4 1 4 35 Shoot number and length Experiment 2 Fresh and dry weights Experiment 2 TOTAL. 99 S - source R - rate A - age Pot/SR - pots within sources and rates IV. RESULTS A. E f f e c t s of Treatments on Growth 1. Fresh and Dry Weights In the f i r s t experiment there were no s i g n i f i -cant differences in t o t a l fresh and dry weights of shoots two weeks after application of the f e r t i l i z e r treatments. However, 14 weeks l a t e r , at a second harvest, both fresh and dry weights of shoots were s i g n i f i c a n t l y higher for plants receiving n i t r a t e than for those receiving urea, (Table 2). In the second experiment, average fresh and dry weights of shoots per plant were affected by an age X source X rate i n t e r a c t i o n , (Figures 1 and 2). Although there were no s i g n i f i c a n t differences in fresh weights of shoots from old cuttings supplemented with N, plants supplied with 120 lb N per acre as urea had higher fresh weight of shoots than those supplied with 30, 60, and 120 lb N per acre as n i t r a t e and 90 lb N per acre as urea. The l a t t e r 4 did not d i f f e r from each other with regard to fresh weights of shoots. Duncan's New Multiple Range Test showed that the "new" control plants d i f f e r e d from the "old" control plants in fresh weights of shoots. Sixty, 90, and 120 lb N per acre as urea gave s i g n i f i c a n t l y higher fresh weight of new shoots than the controls. At 90 lb N per 22 Table 2. E f f e c t s of N source on growth in experiment 1 Treatment M e a n s Standard Urea Nitrate Error Leaf area (mm ) 55 61 (May 17, 1968) Leaves per plant 220 279 12 (May 22, 1968) Fresh weight (gm) 42 58 (June 20, 1968) Dry weight (gm) (June 20, 1968) 15 20 1 23 FIG. I E F F E C T OF SOURCE AND RATE OF N AND AGE OF CUTTINGS ON F R E S H W E I G H T OLD NEW 0 30 60 90 120 0 30 60 90 120 RATE APPLIED N (LB/ACRE) Means not sharing the same l e t t e r d i f f e r s i g n i f i c a n t l y according to D.N.M.R.T. - 5% 24 F I G . 2 EFFECT OF SOURCE AND RATE OF N AMD AGE OF CUTTINGS ON DRY WEIGHT 3 . 5 3 . 0 X o UJ > o 2 . 5 2 0 OLD n - ' 1 0 r a n LiDiDidn "O T3 T3 NEW o o U R E A N I T R A T E •8 3 0 6 0 9 0 1 2 0 0 3 0 6 0 9 0 1 2 0 R A T E A P P L I E D N ( L B / A C R E ) Means not sharing the same letter differ significantly according to D.N.M.R.T. - 5% acre as nitrate, fresh weight of shoots was significantly less than in the controls or any other treatment. None of the treatments significantly affected shoot dry weight in comparison with the controls, but 90 lb per acre N applied as nitrate resulted in signif-icantly less dry weight than 30 and 60 lb per acre N as nitrate. 2. Shoot Number and Shoot Length An analysis of covariance was carried out on shoot lengths and numbers measured on March 14, (before f e r t i l i z e r applications), and on March 28, (2 weeks after treatments); no significant differences were detected. However, increases in shoot length and number and f i n a l shoot number were significantly correlated with leaf Mg at f i r s t harvest. There was no significant difference between urea and nitrate in shoot length of old cuttings, but there were significant age X rate and age X source interactions with regard to average total shoot length per plant. New cuttings showed significantly greater shoot length under the urea regime, though under either N source, total shoot length of new cuttings was s i g n i f i -cantly greater than that of old cuttings, (Figures 3 and 4). Young cuttings did not differ significantly 2 6 . 3 E F F E C T O F A G E O F C U T T I N G S A N D N S O U R C E O N S H O O T L E N G T H Means not sharing the same l e t t e r d i f f e r s i g n i f i c a n t l y according to D . N . M . R . T . -27 FIQ. 4 E F F E C T O F A G E O F C U T T I N G S A N D R A T E O F A P P L I E D N O N S H O O T L E N G T H 50 4 5 3E An z Id O o X co 35 3 0 25 20 • OLD I | N E W n Ml 0 30 o a S-y = 3 60 9 0 120 RATE APPLIED N ( L B / A C R E ) Means not sharing the same l e t t e r d i f f e r s i g n i f i c a n t l y according to D.N.M.R.T. - 5% 28 from one another, but produced more vine growth than the old cuttings. Vine length of old cuttings receiving supplementary N were not significantly different from the controls. Shoot number per plant in the second experiment, (Figure 5), was significantly higher for old cuttings than for new. Shoot numbers for new cuttings did not differ significantly from the controls. For old cuttings, shoot numbers were higher at 120 lb applied N than at 0, 60, and 90 lb per acre. 3. Leaf Number In the f i r s t experiment, a leaf count made 9 weeks after f e r t i l i z e r treatment showed significantly more leaves for plants receiving nitrate than for those receiving urea, (Table 2). In the second experiment, average leaf numbers were calculated from total leaf counts made on new and old plants. There were significantly fewer leaves per plant from new cuttings receiving nitrate N than those receiving urea or than for old cuttings, (Figure 6). 4. Leaf Area Leaf area was measured in the f i r s t experiment 2 months after treatment applications. Leaf areas of plants receiving nitrate were significantly greater 29 F I G . 5 E F F E C T O F A G E O F C U T T I N G S A N D R A T E O F A P P L I E D N O N S H O O T N U M B E R • O L D I I N E W R A T E A P P L I E D N ( L B / A C R E ) Means not sharing the same l e t t e r d i f f e r s i g n i f i c a n t l y according to D.N.M.R.T. - 5% 30 FIG. 6 EFFECT OF AGE OF CUTTINGS AND N SOURCE ON LEAF NUMBER Means not sharing the same letter differ significant according to D.N.M.R.T. - 5% 31 than those receiving urea, (Table 2). B. Effects of Treatments on Peat Leachate 1. pH In the f i r s t experiment, pH of the peat leach-ate was not significantly affected by the treatments, but in the second experiment, there was a highly significant source X rate interaction, (Figure 7). Sixty and 120 lb N per acre as nitrate caused pH to be significantly higher than the controls or any of the other treatments. 2. Ammonium Only in the f i r s t experiment was the s o i l leachate analyzed for NH^ content which was found to be significantly higher than the controls at 90 and 120 lb N per acre in either the nitrate or urea form, (Table 3). 3. Nitrogen In the f i r s t experiment, total N content of the peat effluent was significantly higher in treated pots, (Table 3). In general, N content of the s o i l leachate rose with the amount of applied N. In the second experiment, a significant source X rate interaction on so i l effluent N content occurred, (Figure 8). Pots treated with urea at 60 and 120 lb 32 F I G . 7 EFFECT OF SOURCE AND RATE OF APPLIED N ON PEAT EFFLUENT pH • U R E A • N I T R A T E 4.7 4 . 6 -4.3 UJ d 4 .4 ui 4.3 -4.2 = 0.04 3 0 6 0 9 0 1 2 0 R A T E A P P L I E D N ( L B / A C R E ) Means not s h a r i n g the same l e t t e r d i f f e r s i g n i f i c a n t l y a c c o r d i n g to D.N.M.R.T. - ^% 33 F I G . 8 EFFECT OF SOURCE AND RATE OF APPLIED N ON PEAT EFFLUENT N .3 R A T E A P P L I E D N ( L B / A C R E ) Means not sharing the same letter differ significantly according to D.N.M.R.T. - t% 34 N per a c r e showed a s i g n i f i c a n t l y h i g h e r N content o f peat l e a c h a t e than the c o n t r o l s . Except f o r 60 l b per ac re N , n i t r a t e a p p l i c a -t i o n s showed s i g n i f i c a n t l y h i g h e r e f f l u e n t N content than the c o n t r o l s . O n l y at 30 l b . N per ac re were n i t r a t e t r e a t e d po t s o f s i g n i f i c a n t l y h i g h e r e f f l u e n t N conten t than u r e a at the same r a t e . T a b l e 3 . E f f e c t o f r a t e o f a p p l i e d N on s o i l parameters i n experiment I f Rate o f A p p l i e d N ( l b per ac re ) ; : s t a n d a r d 0 30 60 90 120 E r r o r N i n e f f l u e n t (ppm) 5c 10b 12ab 19a 17a 2 (Mean per t r e a t . ) N H 3 i n e f f l u e n t (ppm) 4b 7a 8b 11a 11a 2 (Mean per t r e a t . ) f Means not s h a r i n g the same l e t t e r d i f f e r s i g n i f i c a n t l y w i t h i n a row a c c o r d i n g to D . N . M . R . T . - 5% C . E f f e c t s o f Trea tments on F o l i a r N u t r i e n t C o m p o s i t i o n Average m i n e r a l l e v e l s f o r the 3 a n a l y s e s were not compared. The grand means f o r each element i n each exper iment are p re sen ted i n T a b l e 4 , r e p r e s e n t i n g average f o l i a r content over a l l s o u r c e s , r a t e s and r e p l i c a t e s . 35 Table 4. Grand means f o r f o l i a r mineral l e v e l s i n the three harvests (ppm) Element Experiment 1 1 2 1st harvest 2nd harvest r e s i d u a l r e s i d u a l r e s i d u a l ppm v a r i a b i l i t y f ppm v a r i a b i l i t y ! ppm v a r i a b i l i t y ! N 23850 3023000 11335 1620100 8733 1852300 P 1456 103910 556 15147 171 7758 K 11263 416980 4540 362170 7536 162930 Ca 992 5851 1056 17823 2478 603590 Mg 231 578 3927 119646 1804 22389 Fe 113 549 86 370 96 346 Mn 201 1220 442 4539 39 68 f R e s i d u a l v a r i a b i l i t y (s ) f o r each element represents the measure of v a r i a b i l i t y among pots of the same source and r a t e of a p p l i e d N . \ 36 1. N i t r o g e n In the f i r s t experiment, ( f i r s t h a r v e s t ) , t r e a t -ments with urea at 60 l b N per acre gave s i g n i f i c a n t l y h i g h e r f o l i a r N l e v e l s than d i d the c o n t r o l s or those at 30 and 120 l b N per a c r e . With n i t r a t e at 90 and 120 l b N per acre, f o l i a r N was s i g n i f i c a n t l y g r e a t e r than i n c o n t r o l s but d i d not d i f f e r s i g n i f i c a n t l y from the 30 l b r a t e , ( F i g u r e 9 ) . F o l i a r N was hig h e r i n p l a n t s r e c e i v i n g 90 or 120 l b N per acre as n i t r a t e than i n the c o n t r o l s or p l a n t s r e c e i v i n g 30 or 120 l b N as u r e a . The second harvest of the same p l a n t s , (14 weeks a f t e r the f i r s t h a r v e s t ) , showed a response of f o l i a r N to r a t e of N a p p l i e d to the s o i l . P l a n t s r e c e i v i n g 90 or 120 l b N per acre had s i g n i f i c a n t l y h i g h e r f o l i a r N than those r e c e i v i n g 0 or 30 l b N per acre, (Table 5 ) . In the second experiment, f o l i a r N was found to be hig h e r i n p l a n t s s u p p l i e d with urea than i n p l a n t s s u p p l i e d with n i t r a t e , (Table 6 ) . 2. Phosphorus P content of the l e a f t i s s u e was higher i n the f i r s t h a rvest i n p l a n t s s u p p l i e d with urea. In the second harvest there was no s i g n i f i c a n t response to d i f f e r e n t i a l treatments. In the second experiment, 37 F I G . 9 EFFECT OF SOURCE AND RATE OF APPLIED N ON FOLIAR N = 7 7 8 0 3 0 6 0 9 0 1 2 0 R A T E A P P L I E D N ( L B / A C R E ) Means not sharing the same letter differ significantly according to D.N.M.R.T. - b% 38 Tabl e 5. E f f e c t s of r a t e s of a p p l i e d N on f o l i a r m i n e r a l composition (ppm) t Element 0 Rate ( l b N per acre) 30 60 90 120 Standard E r r o r (Experiment 1 - F i r s t h a r v e s t ) Mg 249a 229ab 231ab 232ab 212b 8 (Experiment 1 - Second h a r v e s t ) N 10299b 10729b 11431ab 11993a 12220a 403 K 5166a 4570b 4300b 4313b 4350b 190 Ca 1192a 1096ab 990b 1010b 986b 42 Mn 509a 468ab 435bc 402c 396c 22 (Experiment 2) P 885a 604b 691b 684b 660b t t t Means not s h a r i n g the same l e t t e r d i f f e r s i g n i f i c a n t l y a c c o r d i n g to D.N.M.R.T. - 5% t t S e v e r a l standard e r r o r s were used i n comparing the means of unequal sample s i z e s . 39 able 6. E f f e c t s of sources of applied N on f o l i a r mineral composition (ppm) Element S o u r c e Standard Urea Nitrate Error (Experiment 1 - F i r s t harvest) K 10579 11949 129 M g 224 238 5 P 1270 1642 64 (Experiment 1 - Second harvest) K 4791 4289 170 M g (Experiment 2) 75 79 2 N 9313 8514 t K 7369 7703 t Several standard errors were used in comparing the means of unequal sample s i z e s . 40 appl ied N caused a decrease in f o l i a r P l e v e l s , (Tables 5 and 6) . 3. Potassium F o l i a r K was found to be higher in plants supplied with n i t r a t e in the f i r s t harvest of the f i r s t experiment and in the second experiment. In the second harvest of the f i r s t experiment, however, f o l i a r K was higher in plants supplied with urea, and app l i ca t ion of N e i ther as n i t r a t e , or urea increased leaf K content, (Tables 5 and 6) . 4. Calcium In the f i r s t experiment, in the f i r s t harvest urea app l i ca t ions d id not s i g n i f i c a n t l y affect f o l i a r C a ' l e v e l s . P lants r ece iv ing n i t r a t e app l i ca t ions at 60, 90, and 120 lb N per acre had s i g n i f i c a n t l y less f o l i a r Ca than e i ther contro ls or urea treated plants , (Figure 10). In general , f o l i a r Ca l eve l s decreased with increased app l i ca t ions of n i t r a t e . At the second harvest , Ca l eve l s were also decreased in plants supplied with N except at 30 lb N per acre . In the second experiment, f o l i a r Ca l eve l s were not s i g n i f i c a n t l y affected by d i f f e r e n t i a l f e r t i l i z e r a p p l i c a t i o n s . 41 no. 10 EFFECT OF SOURCE AND RATE OF APPLIED N ON FOLIAR Ca MOO -a. 1000 o 900 -800 • I UREA • NITRATE = 34 30 60 90 120 RATE APPLIED N (LB/ACRE) Means not sharing the same letter differ significantly according to D.N.M.R.T. - 5% 42 5 . Magnesium In the f i r s t experiment, fol i a r Mg levels were higher in plants supplied with nitrate compared with those supplied with urea. In the f i r s t harvest, 120 lb N per acre decreased leaf Mg levels, (Tables 5 and 6). In the second experiment, 60 and 90 lb N per acre as nitrate resulted in lower leaf Mg content than in any other treatment, (Figure l l ) . 6. Iron Foliar Fe levels did not differ significantly as a result of differential N treatments. 7. Manganese In the f i r s t experiment, the f i r s t harvest showed a significant source X rate interaction with respect to f o l i a r Mn, (Figure 12). For this analysis the controls were significantly different, and none of the f o l i a r Mn levels were significantly different from the controls. However, plants receiving nitrate at 120 lb N per acre were significantly lower in leaf Mn than those at 30 and 60 lb N per acre as nitrate, or 120 lb N per acre as urea. In the second harvest, the controls had higher Mn levels in the foliage than plants receiving more than 30 lb N per acre. In general, fol i a r Mn decreased with increased rates of N application, (Table 5 ) . 43 FIG. Jl EFFECT OF SOURCE AND RATE OF APPLIED N ON FOLIAR Mg • UREA o • N I T R A T E 2200 I _ 2000 cc 1800 < 3 o U. 1600 1400 -o xt o J Q O XI "8 o XI u n 90 60 90 120 RATE APPLIED N (LB/ACRE) Means n o t s h a r i n g t h e same l e t t e r d i f f e r s i g n i f i c a n t l y a c c o r d i n g t o D . N . M . R . T . - 5% 44 Fie. 12 400 EFFECT OF SOURCE AND RATE OF APPLIED N ON FOLIAR Mn • UREA • NITRATE 300 200 o U . 100 2 1 S ^ = 16 30 60 90 120 RATE APPLIED N (LB/ ACRE) Means not sharing the same l e t t e r d i f f e r s i g n i f i c a n t l y according to D.N.M.R.T. - 5% 45 D. Multiple Regression Analysis Multiple regression analysis was carried out on the leaf mineral content and growth measurements made in the f i r s t experiment, (Table 7), and unless specified, a l l r e s u l t s discussed re f e r to those s i g n i f i c a n t at the 1% l e v e l . Using the f i r s t growth measurements as dependent variables, f o l i a r mineral l e v e l s at f i r s t harvest were correlated. It was found that increase in shoot length and shoot number, and f i n a l shoot number corresponded to an increase in f o l i a r Mg. F i n a l shoot length, fresh weight and dry weight corresponded to increases in both f o l i a r Mg and Mn l e v e l s , (Table 8 ) . In a second set of multiple correlations, f o l i a r mineral contents at second harvest were related to f i r s t harvest growth measurements and f o l i a r mineral l e v e l s . High f o l i a r K and Ca both corresponded to low f o l i a r N at f i r s t harvest. F o l i a r P was multiply correlated with N (negatively) and P ( p o s i t i v e l y ) at f i r s t harvest. F o l i a r Mn lev e l s corresponded in a positive way to f o l i a r N and K l e v e l s in the f i r s t harvest. Leaf Mg in the second analysis was higher in plants showing a larger shoot number at the f i r s t harvest, (Table 9 ) . With f i n a l growth measurements as dependent variables, and both mineral analyses and f i r s t growth 46 Table 7. List of variables used for multiple regression analysis for experiment 1 First Growth First Mineral Second Growth Second Mineral Measurements Analysis Measurements Analysis Increase in 1-N Second fresh 2-N shoot length weight 1-P 2-P Increase in Second dry shoot number 1-K weight 2-K First fresh 1-Ca Leaf area 2-Ca weight 1-Mg Leaf number 2-Mg First dry weight 1-Fe 2-Fe Final shoot 1-Mn 2-Mn length Final shoot number Table 8. Significant correlations between growth measurements and mineral analyses at f i r s t harvest, (experiment l) Dependent Variable Independent Variable b lOOR2^ Pr o b a b i l i t y Increase in 1--Mg -5 .5530 0. 1415 39.63 0. 0001 shoot length Increase in 1--Mg -11 .5100 0. 1432 33.48 0. 0 0 0 1 shoot number Final shoot 1--Mg -3 .0289 0. 1550 40.51 0. 0 0 0 1 length 1--Mn 0. 0505 Final shoot 1--Mg 3 .1068 0. 1587 32.49 0. 0 0 0 1 number First fresh 1--Mn -1 .6104 0. 0139 41.54 0 . 0 0 0 1 weight 1--Mg 0. 0291 First dry 1--Mn 0 . 0947 0. 0035 33.48 0 . 0 0 0 1 weight 1--Mg 0 . 0053 47 Table 9. S i g n i f i c a n t c o r r e l a t i o n s among m i n e r a l analyses i n the second harvest and growth and mi n e r a l a n a l y s e s i n the f i r s t h a r v e s t , (experiment 1) Dependent V a r i a b l e Independent V a r i a b l e b 100R^ P r o b a b i l i t y 2-P 2-K 2-Ca 2-Mg 2-Mn 1-N 1-P 1-N 1-N 825.9 -0.0249 25.85 0.2221 9435.0 -0.2052 44.34 2534.6 -0.0399 17.74 F i n a l shoot 2882.9 26.4419 27.62 number 1-N 1-K -488.7 0.0230 41.05 0.0339 0.0010 0.0001 0.0001 0.0001 0.0002 Tabl e 10. S i g n i f i c a n t c o r r e l a t i o n s between growth measurements before second harvest and a l l other measurements made i n the f i r s t experiment. Dependent V a r i a b l e Independent V a r i a b l e b lOOR 2^ P r o b a b i l i t y Second f r e s h F i r s t f r e s h -23.5403 3.0941 82.94 0.0001 weight Second dry weight Leaf area Leaf number weight 1-P 1-N 1-K F i r s t f r e s h weight 1-P 1-N 1-K 1-P 1-K Increase i n shoot l e n g t h F i n a l shoot number 0.0283 0.0038 -0.0077 -1.1248 0.9764 83.37 0.0096 0.0009 -0.0029 66.7110 0.0097 82.94 -0.0051 -0.7395 5.0428 30.83 -2.6577 0.0001 0.0001 0.0002 48 measurements as independent variables, a third set of correlations was carried out. Increases in both fresh and dry weights were significantly correlated with increases in fresh weights, and fo l i a r N and P, and decreases in f o l i a r K at f i r s t harvest. Increase in leaf area was found to reflect high fol i a r P at f i r s t harvest and low f o l i a r K at second harvest. Leaf number was positively correlated with increase in shoot length and negatively with f i n a l shoot number, (Table 10). Simple correlations were many and i t was found that except for leaf number, which could be correlated only with f i n a l shoot length, a l l other growth measure-ments could be correlated significantly with one another. In the second experiment, growth measurements were correlated as dependent variables, with s o i l pH and N content, and f o l i a r mineral levels, and in a second consideration, plant mineral analysis was cor-related with the s o i l analysis data. Significant results are presented in Table 11. Fresh weight was multiply correlated with s o i l pH (negatively), and soil N content (positively). Shoot number and dry weight both increased with decrease in fo l i a r P. High levels of fo l i a r K reflected low soil pH. Simple correlations were also carried out, (Table 12). 49 Table 11. Significant correlations from multiple regression in experiment 2 Dependent Variable Independent Variable Fresh weight Soil pH Soil N Shoot number Dry weight Foliar K P P Soil pH a b 100FT% Probability 1.1130 -0.1721 27.46 0.0095 0.0594 4.8116 -0.0020 14.46 0.0303 0.3702 -0.0002 30.17 0.0012 12750. -119.2868 27.75 0.0020 Table 12. Significant linear correlations in experiment 2 Dependent Independent Variable Variable • Fe Mn Mn Fresh weight Fresh weight Dry weight Dry weight Ca Ca Fe Soil N P P Soil N 0.3491* 0.3656* 0.9985** 0.4068* 0.3807* 0.5492** 0.4042* * Significant at the 5% level ** Significant at the 1% level or lower 50 Foliar Fe was significantly correlated with f o l i a r Ca, f o l i a r Mn with Ca and Fe, fresh weight with so i l N and •foliar P, and dry weight of shoots with so i l N and f o l i a r P. It was also found that a l l growth measure-ments could be correlated significantly with a l l others. 51 V. DISCUSSION A. Growth It was found, in the f i r s t experiment, that after 2 weeks there were no measurable differences in growth as a result of N applications, though foli a r mineral composition reflected differences in regime even at this early stage. Both growth and compositional differences were apparent after 14 additional weeks of growth. The cuttings used in the f i r s t experiment were rooted in the winter. There was more growth in plants receiving nitrate N; leaves were larger and more numerous, and consequently, fresh and dry weights were much higher. Since s o i l pH was acidic, (3.7) the results agree with those of Addoms and Mounce (2) who found that nitrate gave the most favorable growth under acid conditions. Indeed, in the f i r s t analysis i t was found that more N was taken up in plants supplied with nitrate N, whereas urea treated plants did not differ significantly in f o l i a r N from the controls. Application as nitrate would appear to be the better choice in that abundant vegetative growth is promoted and furthermore is reflected in tissue N content. Later, however, in the second analysis, only a main effect of rate was evident, and in the multiple 52 regression analysis, early N content was partially correlated with fresh and dry weights. Foliar N was higher in plants treated with higher rates of N. Total N content of peat effluent samples increased as rates of applied N increased. Sixty lb per acre N would appear most practical since leachate N content did not increase with higher rates, yet high N rates increased growth. In the second experiment, i t was found that cuttings rooted in the spring gave more favorable growth than those which had been rooted since the previous November. Furthermore, new cuttings gave a greater response to f e r t i l i z e r treatments, even though the old cuttings had developed an extensive root system. This may indicate the need to f e r t i l i z e the cuttings early or that cuttings made from actively growing runners give better growth response with or without N applications than those made from physiolog-i c a l l y inactive runners in the winter. Fresh and dry weights and average total shoot length were significantly lower for old cuttings which did not respond to f e r t i l i z e r applications. Unfortunately, i t was necessary to pool shoots from the new and old cuttings in order to obtain sufficient dry material for f o l i a r mineral analysis and conclusions 53 could not be drawn regarding N uptake in the 2 kinds of cuttings. Higher N content would probably have been found in the vegetatively proliferating new cuttings. Since average leaf number and shoot numbers per plant were found to be lower in new cuttings while fresh and dry weights and shoot length were higher, i t must be concluded that the leaves were larger, and/or the stem length produced the additional shoot weight. There was more net vegetative growth in new cuttings whereas in old cuttings, shoot and leaf i n i t i a t i o n predominated over increase in size. New cuttings produced greater fresh and dry weights than the old cuttings, and furthermore showed a response in increase in fresh weight, to urea applic-ations at 60 lb per acre and greater, though applications of more than 60 lb per acre did not further increase fresh weight. There were also more leaves on plants receiving urea, a contradiction to the f i r s t experiment. Since dry weights of treated cuttings were not significan different from the controls, plants supplied with N f e r t i l i z e r may have been more succulent, a characteristic of vigorous vegetative growth. B. Mineral Analyses 1. Nitrogen Foliar N content in the f i r s t experiment was 54 found to be higher in plants supplied with higher N f e r t i l i z e r rates, and was an important indicator for fresh and dry weight at second harvest. This result appears reasonable as vegetative growth is generally stimulated by N which is used in the production of proteins and other"building" materials. It is d i f f i c u l t to relate growth and f o l i a r N content in the second experiment since growth measure-ments were significantly different in new and old plants and a l l of the material was pooled for the mineral analysis. It was found, however, that urea applications caused greater N uptake by the plants than did nitrate. This contradiction of the results of the f i r s t experiment may possibly be the result of pooling the plant material. It might also be explained by s o i l pH and N status since fresh weight of shoots was correlated with both s o i l pH and N content, and dry weight with s o i l N. In the f i r s t experiment there were no significant differences in s o i l pH, and nitrate at moderate and high levels gave the most favorable growth. In the second experiment, however, at moderate and high levels, nitrate caused s o i l pH to rise significantly which may have inhibited i t s uptake, although s o i l N content was not significantly different in pots supplied with N. 55 2. Phosphorus In the f i r s t experiment, fo l i a r P was higher under the nitrate regime after 2 weeks, whereas after 14 more w e e k s , there was no significant difference in P levels. The resluts of the f i r s t analysis did not agree with the findings of Reuther and Smith (36) who generalized that high N content, (found here in the nitrate treated plants), led to a correspondingly low level of fol i a r P. Neither did this appear to hold at the second harvest where fo l i a r N was higher at higher rates of N application, but P was not affected, although the second P analysis was correlated significantly with that of the f i r s t analysis and could provide c r i t e r i a for vegetative growth. In the second experiment, foli a r P was significantly greater in the controls and in the regression analysis was negatively correlated with growth measurements. 3. Potassium In the f i r s t harvest of the f i r s t experiment, fo l i a r K was higher in plants treated with nitrate, but in the second harvest was higher in plants treated with urea. In the second experiment, foli a r K was again significantly higher under the nitrate regime and in controls compared with treated pots. It was found that K uptake was greater in more acidic s o i l . 56 Plants with more vegetative growth contained a lower concentration of K in the leaves and this may have been a dilution effect. It may be concluded that no K need be added to peat soils for cranberry growing. 4. Calcium In the f i r s t analysis of the f i r s t experiment, f o l i a r Ca was lower in plants which had more growth and which had received higher rates of nitrate. In the second analysis, similarly, plants receiving higher rates of N and having more vegetative growth were low in Ca. This may be interpreted as a dilution effect, vand may indicate a lack of available Ca in the s o i l . Ca is known to be low in acidic soils such as those used in cranberry culture, and experiments involving liming might be carried out, though Herath and Eaton (28) found that in blueberry, also an Ericaceous species, that liming did not significantly affect f o l i a r Ca levels though i t tended to increase N and K uptake. Furthermore, i t was found that high rates of nitrate and corresponding Ca deficiency resulted in leaf burning. In the second experiment, there did not appear to be a significant effect of f e r t i l i z e r treatments on fo l i a r Ca, and again a separate analysis might have been useful in order to compare uptake for rapidly 57 p r o l i f e r a t i n g shoots of newly rooted cuttings and slower growing shoots of older rooted cuttings. 5. Magnesium Mg content of the leaves was found in both analyses of the f i r s t experiment to be higher under the n i t r a t e regime. This agrees with the findings of Reuther and Smith (36) who found that in c i t r u s , f o l i a r Mg was higher in fo l i a g e containing higher l e v e l s of N. Higher l e v e l s of K and Ca were also associated with low Mg l e v e l s and both were demonstrated in the second analysis, though the negative r e l a t i o n s h i p with K was not demonstrated in the f i r s t analysis. Every growth measurement was s i g n i f i c a n t l y correlated with Mg alone or Mg and Mn after 2 weeks of growth under the d i f f e r e n t i a l f e r t i l i z e r regimes, ( i t should be noted that a l l growth measurements could be correlated with one another). That Mg l e v e l s were l a t e r not correlated with growth may indicate an i n i t i a l requirement for Mg. Results of the second experiment were contra-dictory to those in the f i r s t with regard to Mg l e v e l s . F o l i a r Mg was lower in plants treated with n i t r a t e which tended to have less growth than the plants receiving urea and which had higher f o l i a r N. There were no s i g n i f i -cant correlations involving Mg in this experiment, though 58 the negative relationship to K was again demonstrated. 6. Iron In none of the analyses was fo l i a r Fe s i g n i f i -cantly affected by differential N treatments. However, i t is recommended by this investigator that a more concentrated mineral extract be made in further exper-iments of this kind. Atomic absorption readings for this element were very low, the range of standards used was only from 0.5 to 2.5 ppm. In analysing for Fe, i t must also be remembered that the ferrous ion is metabolically active and that plants containing ferric ions may appear Fe deficient (42). Medappa (31) found that there was no difference in growth when Fe was applied in solution culture to cranberries at pH 3, 4, and 5. 7. Manganese In the f i r s t experiment, Mn showed a significant interaction as a result of source and rate of applied N, but conclusions are d i f f i c u l t since controls differed significantly in fo l i a r Mn content, and none of the treated pots differed from the controls. In the second harvest, f o l i a r Mn was higher in the controls than in plants supplied with N. This too may have been explained as a dilution effect since plants were larger at this harvest and growth differences were more pronounced 59 than at any other harvest. Even so, Mn was an indicator of fresh and dry weight and f i n a l shoot length after f i r s t harvest and may indicate an i n i t i a l requirement. 8. General Several factors may have been responsible for the fact that levels of elements were not comparable in the three analyses. Cuttings were gathered from different fields which may have had different environ-mental conditions and different histories. They were also collected at different times of year. Age of the rooted cuttings before f e r t i l i z e r application was quite clearly a factor influencing growth, and possibly mineral uptake. Pooling of samples may also have affected mineral evaluation. In some cases a dilution effect was thought to have taken place in rapidly growing material. 60 VI. SUMMARY Rooted cuttings in the f i r s t experiment and "new" rooted cuttings in the second experiment showed growth responses to differential N applications, but the "old" cuttings were negligibly affected, hence immediate f e r t i l i z e r application appeared beneficial. There were definite responses to the form of N applied but the results were contradictory in the 2 experiments. It was found, however, that f o l i a r mineral constituents bore relationships to the amount of growth made by the plants. N was higher in rapidly growing plants whereas P, Fe, and Ca were lower and ascribed to dilution effects or differences in physiol-ogical age. Growth in the f i r s t experiment was greater under the nitrate regime though shoot length was greater at low N rates. In the second experiment, 60 lb N per acre as urea was sufficient to produce maximum vegetative growth. Only shoot and leaf number were greater for old rooted cuttings but fresh and dry weights and shoot lengths were lower indicating an expenditure in old cuttings for in i t i a t i o n rather than increase in size. Soil pH was not affected in the f i r s t experiment but in the second i t was found that nitrate treatments 6 1 had caused a r i s e in s o i l pH which p a r a l l e l e d an increase i n f o l i a r K, and a decrease in fresh weight (accompanied by low f o l i a r N) . It may be concluded that n i t r a t e was the better N source only i f the pH was maintained at a low l e v e l . Herath and Eaton (28) concluded that high ra tes of n i t r a t e were harmful or toxic to blueberry plants when s o i l pH was h igh . Peat leachate N was higher for increas ing rates of appl ied N as was leachate ammonia content. Growth may be predicted ear ly by Mg and Mn ana lys i s of the f o l i a g e , but both growth and mineral content at a l a t e r time may be indicated ear ly by N, P, and K f o l i a r l e v e l s . Source and time of root ing of cutt ings and length of time before f e r t i l i z e r appl ica t ions may have inf luenced f u l l y , or in part , growth and mineral uptake and were sources of v a r i a t i o n in t h i s experiment. 62 V I I . BIBLIOGRAPHY 1. Addoms, R . M . , and F . C . Mounce 1931. Notes on the nutr ient requirements and the h i s to logy of the cranberry (Vaccinium  macrocarpon A i t . ) with spec ia l reference to mycorrhiza. P I . Phys. 6:653-668. 2. 3. and 1932. Further notes on the nutr ient requirements and the h i s to logy of the cranberry, with spec ia l reference to the sources of n i t rogen . P I . Phys. 7:643-656. Anon. 1953-4.The cranberry s t a t i o n . East Wareham, Mass . , Ann. Rep. Mass. Agr . Expt . S ta . B u i . 472:41-45. 4. 1955, E f f e c t s of f e r t i l i z e r on cranberry upr ight s . Cranberr ies 19(11):9. 5. 1956, Report of the Canada Min i s ter of A g r i c u l t u r e for year ended March 31, 1956. p. 101. 6. 7. 1960. N . J . f e r t i l i z e r and lime recommendations, N . J . Agr . Expt . S t a . C i r c . 589. Beckwith, C . S . 1919. Ef fec t of c e r t a i n nitrogenous and phosphatic f e r t i l i z e r s on the y i e l d of c r a n b e r r i e s . S o i l S c i . 8:483-490. 8. 9. Bould, C . 1966. C a i n , C . J . 1954. Leaf ana lys i s of deciduous f r u i t s . In Temperate to t r o p i c a l f r u i t n u t r i t i o n . Edited by N . F . C h i l d e r s , Somerset Press , Somervi l l e , New Jersey , p.651-684. and G . J . Galletta Blueberry and cranberry n u t r i t i o n . In F r u i t n u t r i t i o n . Edited by N . F . C h i l d e r s , H o r t i c u l t u r a l P u b l i c a t i o n s , Rutgers U n i v e r s i t y , New Jersey , p .121 -152. 1 0 . Chandler, F.B. 1956. Timely facts on f e r t i l i z a t i o n . Cranberr ies 2 l ( l ) : 12 -13 . 63 11. , and W.G. Colby 1949. F e r t i l i z e r requirements of c r a n b e r r y . Ann. Rep. Mass. Agr. Expt. S t a . B u i . 453:36. 12. Chapman, H.D., and P. P r a t t 1961. Methods of a n a l y s i s f o r s o i l s , p l a n t s and waters. U n i v e r s i t y of C a l i f o r n i a , D i v i s i o n of A g r i c u l t u r a l S c i e n c e s . 13. Colby, W.G. 1945. The use of commercial f e r t i l i z e r s on c r a n b e r r i e s . C r a n b e r r i e s 10(6):6-7. 14. Crowley, D.J. 1954. Cranberry growing i n Washington. Wash. Expt. S t a . B u i . 554. 15. Dana, M.N. 1968. N i t r o g e n f e r t i l i z a t i o n and c r a n b e r r i e s , P a r t I . C r a n b e r r i e s 32(12):10-11. 16. 20. 1968. N i t r o g e n f e r t i l i z a t i o n and c r a n b e r r i e s , P a r t I I . C r a n b e r r i e s 33(1)-.10-11,15. 17. DeLong, W.A. 1965. The n i t r o g e n n u t r i t i o n of woody f r u i t c r o p s . Unpublished mimeographed review. Can. Dept. Agr. Res. S t a . , K e n t v i l l e Nova S c o t i a . 18. Dickman, S.R., and R.H. Bray 1940. C o l o r i m e t r i c d e t e r m i n a t i o n of phosphate. Ind. Eng. Chem. A n a l . Ed. 12:665-668. 19. D o e h l e r t , C A . 1954. Composition and t i m i n g of c r a n b e r r y f e r t i l i z i n g . P r o c . Amer. Cranberry Growers Assoc. p. 9-17. 1955. Cranberry f e r t i l i z e r r e s e a r c h i n New J e r s e y . C r a n b e r r i e s 1 9 ( l l ) : 1 0 - 1 2 . 21. Eaton, E.L. 1948. Cranberry c u l t u r e . CD.A. Farmer's B u i . 151:5-26. 22. Eck, P. 1962. Cranberry f e r t i l i z e r s t u d i e s . P r o c . Amer. Cranberry Growers Assoc. p.27-33, 64 23. Edmond, J . B . , T . L . Senn, and F . S . Andrews 1964. Fundamentals of h o r t i c u l t u r e . McGraw-H i l l Book Company, New York. 24. F i s h e r , R . A . 1951. S o i l Data on n u t r i t i o n on Washington state bogs. Cranberr ies 16(2):8-10. 25. F r a n k l i n , H . J . 1948. Cranberry growing in Massachusetts. Mass. Agr . Expt . S ta . B u i . 447:1-44. 26. H a l l , I . V . , L . E . Alders and L . R . Townsend 1964. The effect of s o i l pH on the mineral composition and growth of lowbush b lueberry . Can. J . Plant S c i . 44: 143-144. 27. H a r r i s . G . H . 1955. Small f r u i t s . Cranberr i e s . Can. Hort . Counc i l Rep. on Hort . Res. p.143-144. 28. Herath, H . M . E . , and G.W. Eaton 1968. Some ef fects of water tab le , pH and nitrogen f e r t i l i z a t i o n upon growth and nutr ient element content of highbush blueberry p l a n t s . Proc . Amer. Soc. H o r t . S c i . 92:274-283. 29. Kender, W . J . , and N . F . C h i l d e r s 1959. Growth of cranberry plants (Vaccinium  macrocarpon) with various sources of n i t rogen . Proc . Amer. Soc. Hort . S c i . 74:407-413. 30. L i , J . C . R . 1965. S t a t i s t i c a l inference I . Edwards Brothers Inc . Ann Arbor, Michigan. 31. Medappa, K . C . 1966. Growth and composition of the cranberry plant in r e l a t i o n to nutrient medium r e a c t i o n . PhD. Thes i s . Univ . Wisconsin. 32. Meyer, B . S . , D .B . Anderson and R . H . Bohning 1966. Plant physiology. D. VanNostrand Co. L t d . , Toronto. 65 33. Morse, F.W. 1930. A chemical study of cranberries. Mass Agr. Expt. Sta. Bui. 265:87-102. 34. Mulder, E.G., R. Boxma and W.L. Van Veen 1959. The effect of Mo and N deficiencies on nitrate reduction in plant tissues. Plant and Soil 10:335-355. 35. Peltier, G.L. 1955. Wisconsin f e r t i l i z e r . Cranberries 9(11):14-15. 36. Reuther, W. and P.F. Smith, 195H. Leaf analysis of citrus. In Fruit nutrition. Fir» t edition. Edited by N.F. Childers, Somerset Press, Somerville, New Jersey, p. 257-294. 37. Schlatter, F.P. 1917. Report on f e r t i l i z e r experiments with cranberries. Proc. Amer. Cranberry Growers Assoc. 48:9-12. 38. Somogyi, L.P. 1962. Nitrogen studies with apple and cranberry. Part II - Growth, flower-ing and fruiting of cranberry as influenced by N sources and substrates with different organic matter content. PhD. Thesis, Rutgers Univ. 39. , N.F. Childers and P. Eck 1964. Influence of nitrogen source and organic matter on the cranberry. Proc. Amer. Soc. Hort. Sci. 84: 280-288. 40. Sorensen, L.A. 1955. F e r t i l i z e r s on Wisconsin marshes. Cranberries 19(11):7-8. 41. Thiens, J.R. 1955. F e r t i l i z i n g in Oregon. Cranberries 19(ll):8, 42. Tisdale, S.L. and W.L. Nelson 1966. Soil f e r t i l i t y and f e r t i l i z e r s . Second Edition. The MacMillan Company, New York. 

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