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Responses of the Cuthbert raspberry to mineral treatments Hornby, Cedrick Albert 1940

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RESPONSES Off THE CUTHBSET, RASPBERRY TO MIHERAL TB1A.TKEHTS by Cedrie Albert Hornby A Thesis submitted i n P a r t i a l Eulfilment The Requirements for the Degree of MASTER. j^.SCTHOB- IK AGRICULTURE-in the Department of HORTICULTURE. THE OTIWERSITT 01 BRITISH COLUMBIA. A p r i l , 1940 Table of Contents. Page Introduction ...., 1 Review of Literature 2 Materials and Methods .................. 4 Results 10 (a) Observations on Sand Cultures.. 10 (b) Laboratory Data ............... 15 Discussion .v. 25 Summary 31 f Literature Cited ....................... 32 Ac kn owle dg me nt s 34 KBSP01BES Off- THE, GWMBM&T KfiSEEEBEY TO TSSkTmSTS. Introduction In the. Eraser Valley of. Br i t i s h . Columbia many plan-tations once producing vigorous raspberry plants have suffered a gradual, weakening and. decline i n health* Thus plants are now smaller and less produc.ti.Ye than they should he according to past, standards. Usually no pathological symptoms are. noted, although the land has been planted to raspberries con-tinuously i n many caaes from ten to f i f t y years. From these facts and e a r l i e r work on this problem (9,10), i t seems very probable that the trouble i s a n u t r i t i o n a l one; therefore, i t was decided to study some factors affecting the n u t r i t i o n of the. raspberry plant. It was decided to grow. the. plants i n well washed sand and supply food elements i n solution*, because the s o i l i s such a complex body and provides so many unknown and variable factors that i t i s d i f f i c u l t to study responses of the plant to treatments in this medium, (27). Now, i t i s established that the following elements are necessary for plant growth: nitrogen, phosphorus, potash, calcium, sulphur, magnesium, Iron, boron, manganese, and zinc, and probably others such as copper. Hence, deficiencies of the elements nitrogen, phosphorus, potassium, calcium, and sulphur, were studied and compared with plants which received a l l the necessary nutrients. For vigorous plant growth i t i s important to have a "balance i n -the necessary food elements, because a large quanti-ty of one nutrient i n r e l a t i o n to the others may prove harmful to the plant. Also, the; lack of one: nutrient may cause a v i r t u a l excess of some other element and produce poor plants. There!ore? studies were made on the effects of heavy appli-cations of one element i n r e l a t i o n to other nutrients, and "excess" quantities of nitrogen, phosphorus, potassium, and calcium were used i n treating some, plants. In recent years many of. the so-called deficiency diseases have been found to be caused by lack of the micro or trace elements. Thais, the use of boron, manganes.e, and. zinc was i n -cluded' i n these studies.• Occasionally s o i l substances may be toxic or i n h i b i t plant growth. Aluminum i s known to be. harmful i n acid s o i l s , and since the s o i l s of the raspberry growing areas i n the Fraser Valley are acid, and frequently very acid (.9), i t was decided to study the effects of aluminum on the growth of the raspberry plant. By the use of sand cultures to grow, raspberries with various nutrient treatments, i t was expected to f u l f i l l two objects:: f i r s t l y , to study the effects of essential mineral elements on the growth of the raspberry plant; secondly, to produce under known and controlled conditions, a plant con-d i t i o n similar to that found i n the unthrifty f i e l d plantings. Review of. Literature In the. l a t t e r half of the nineteenth century, Sachs, Khop, U.obbe, P f e f f e r , and others did a great deal of work to determine what elements were essential to. plant growth. Their work showed that carbon, hydrogen, oxygen, nitrogen, phosphor-us, potassium, calcium, magnesium, sulphur, and iron are es-sent i a l . to plant l i f e . . Since, then, workers l i k e Breachley ( l ) , and McHargue (17) have demonstrated the essential nature of the micro elements, boron, manganese, zinc, and copper. This work was accomplished or proved by the use of sand and water cultures. The present day technique of using sand cultures and nutrient solutions has been developed by Hoagland, Shive, and others (13, 87). The symptoms of deficiencies and excesses of these plant food elements as ahown by various parts of the plant have, been studied and recorded for many species (3, 4, 6 12,, 19, 28). Much l i t e r a t u r e lias been published in an, attempt to make laboratory determinations give an expression of the re-sponses noted i n the f i e l d . One of the most favoured has been the so-called carbohydrate nitrogen r a t i o as o r i g i n a l l y formu-lated by Kr.aus. and K r a y b i l l (14), when working with tomatoes, showing the r e l a t i v e proportion of carbohydrates to nitrogen t be a s i g n i f i c a n t factor i n determining vegetative. and repro-ductive a c t i v i t i e s . This work has been developed by many workers, and evidently a narrow r a t i o , or low: carbohydates in proportion to nitrogen is, conducive to vegetative; growth, whereas a wide r a t i o , high carbohydrate content i n r e l a t i o n to nitrogen, i s conducive, to f r u i t i n g . Apparently there i s no literature; i n the. above works dealing with the raspberry. Publications on this, crop deal _^ 4« largely with, the c u l t u r a l aspects. In 1937, Harris (9) named eight factors as being the causes of raspberry f a i l u r e s on some plantings in the Eraser Valley. n These, were: ( l ) a. nitrogen deficiency, (2) a lime de-fic i e n c y , (3) a phosphorus deficiency, (4.) a potassium deficiency, (5) s o i l s being too acid, [6) an unbalanced supply of plant nutrients, (7) the exhaustive; f i r s t year effects of cover cropping on poor or. impoverished soils., and (8) "wet feet" or root injury due to an excess of water in the s o i l . Other work (9) stated that the decline was i n i t i a l l y due to loss, of organic matter with i t s resultant effects of. allowing leaching and undesirable s o i l conditions to develop. Thus starvation produces weakened plants. Materials and Methods Raspberry plants of the Guthbert variety were obtain-ed" and passed by Government o f f i c i a l s , as healthy stock. These were grown in fresh water, pure quartz sand, held i n 12-inch clay flower, pots,. Sixty such pots were divided into fifteen, groups of four, and each, group received a different nutrient solution. The fifteen, treatments were; 1. Complete, nutrient solution 2. Nitrogen deficient nutrient solution 3. Phosphorus deficient nutrient solution 4. Potassium deficient nutrient solution 5. Calcium deficient nutrient solution 6. Sulphur deficient nutrient solution 7. Complete nutrient solution plus excess nitrogen 8. Comple.te nutrient solution plus excess phosphorus 9. Complete nutrient solution plus excess potassium 10. Complete, nutrient solution plus excess calcium 11. Complete; nutrient solution plus 5 p.p.m.. boron, 2 p.p.m. manganese, 1 p.p.m. zinc 12. . Complete, nutrient solution plus 5 p.p.m. boron 13. Complete nutrient solution plus 2 p.p.m. manganese 14. Complete nutrient solution plus 10 p.p.m. aluminum The above f i f t e e n nutrient solutions were based on Hoagland* s Complete Nutrient Solution (13.) which has a concen-tration of approximately 2000 p.p.m. In preparing the nutrient solutions, stock solutions, were made up by dissolving the. following weights of chemicals, in d i v i d u a l l y and making the volume, up to 2 l i t r e s s 1. Complete Nutrients •(a)' (13.4. g. KN.Q3 (200; g. MgS04.7 K a0 (b) 416 g. Ca(N0 3) 2. 4 H20 (e) 100 g. KH2P0-4 2. Nitrogen Def.icie.nt Solution (d) (1.00 g. KC1 (2:00 g. -MgS04. 7H20 (a) 176 g. CaC.l2 (c) 100; g. KH2P04 3. Phosphorus Deficient Solution (a) . (1.34 g. KN02 (200 g. Mgs04.7 H20 (b) 416 g. Ca{N0 3) 2.4 I%0 (f) 55 g. K01, 4. Potassium Be£iciant Solution. Cg:> U l 4 g • • (2.OO; g-. MgS04.7 H20 (b) 416 g. Ga.(lQ 3) 2. 4. H2G (h) 100 g'. HaH2P04 5. Calcium Deficient: Solution (a) (134 g. KHO3 (2.Q0 g. HgS04.7 H20; ( i ) 150 g. ~SaM0z (c) 100 g. KH 2P0 4 6. Sulphate: Deficient Solution (j.) (1.34 g. KKQg >• • (164.9 g. MgCl.2. 4 H20 (b) 416; g. Ca(]J03)2.. 4 H20 (a). 10.0 g. KH 2P0 4 (A bracket indicates that the chemicals grouped together are. included i n the same 2 - l i t r e stock.) Those series receiving an excess or luxury feeding of nitrogen, phosphorus, potassium, and calcium received fi v e times the normal amount of each of. these i n d i v i d u a l nutrients in t h e i r respective diets. The same stocks as f o r f u l l n u t r i -ents i n Solution 1 were used plus these atocks: 7. Excess. Nitrogen. (k) 1.6.49 g. NaN.03 made up to 2 l i t r e s . 8. Excess Phosphorus. (l) 406 g. HaH2P04.H20 made up to 2 l i t r e s . 9. Excess Potassium (m) 613 g. KG1 made up to 2: l i t r e s . ce-^ est 10. Excess: Calcium: (n) 782.5 g-. GaCLg made up to 2.. l i t r e s . Those, groups receiving the micro elements, boron, manganese and zinc, were, given f u l l nutrient solution plus these elements i n the respective: concentrations of 5: p.p.m., and 1 p.p.m«> The. stock solutions were made using the follow-ing quantities of materials made up to 2 l i t r e s : . 11. Complete Nutrients, plus Boron, Manganese, and Zinc (o) 18.32 g. H3BO3 (Boron) made up to 2 l i t r e s . (p) 4.6 g. MhGl.g. 4H2.0-- (Manganese) made up to 2 l i t r e s . (q) 1.34 g. ZnGlg (Zinc), made, up to 2 l i t r e s . 12. Complete Nutrient Solution plus Boron Stock. 13. Complete Nutrient Solution plus Manganese Stock. 14. Complete, nutrient Solution plus Zinc Stock.. 15. 'Clomplata Nutrient Solution plus, 1.0 p.p.m. Aluminum. (r). 56.2 g. A12(S0:4)3.K2S04.2.4 H20- made up to 2 l i t r e s . Thesa stocks were, diluted before, being fed to the plants. The following quantities of stock solutions were, diluted to 4. l i t r e s i n accordance, with the preceding l i 3 t of stocks used to prepare each of the f i f t e e n nutrient treatments. 22. c.c. of stock solutions (a), (d), (g), (j) 26 c.c. of stock solutions (b), (e), ( i ) 12 c.c. of stock solutions (e), ( f ) , (h), ( l ) 10Q c.c. of stock solution (k) 18.5 c.c. of. stock solution (m). 2.6.0 c.c. of stock solution (n). «*8«» 12:. 5 e.a> of stock solutions (o) , (p)',, (q)« When. the. dilution, was made just "before feeding, iron was added at the rate, of 1 c. c. of 1 per cent f e r r i c tar-trate solution per l i t r e of nutrient solution. The raspberries were set i n the sand with one young sucker- cane per pot. Each group of. four, pots w.aa placed i n a square hole in the. earth so that about an. inch of. each pot was above ground l e v e l . The hole was cribbed to keep s o i l away from the pots, and a heavy layer of excelsior, had been put on the. bottom of. the hole to pre.vent any contact between the pots: and s o i l . The spaces, between pots , and pots and cribbing, were f i l l e d with, shavings. Thus evaporating surface was cut down and no s o i l touched the.. p.ata. Plants were set In the pots of sand on May 12, 1.93.7, and le f t , f o r one week to s t a r t root development. Seeding was started May 1.8, 1937. The plants were fed twice, weekly u n t i l June 21, 1937. At that time, the, plants w:ere making vigorous growth and i t was decided to feed them three times each week. They were, watered whenever the feeding did not supply adequate, water. Daily observations were made on the development of the plants.» The.- plants were fed u n t i l the middle, of October, and as the season was very mild and some growth was. s t i l l being made, they were l e f t u n t i l November 1, 1937, before samples fo r anal.ysi3= were taken.. Two representative plants from each group were taken into, the laboratory f o r analyaia except in. the case of the nitr.ogSn and phosphorus deficient groups. These- Latter groups produced: such small, plants that four had to be. used 'to provide, s u f f i c i e n t material for the laboratory determinations. P i n a l observations on the plants including examination of .the roots, were: made, and then the: roots, and aerial, parts were separated. The. roots were washed free of sand, and the fresh weights of roots aoid a e r i a l parts were, measured. Leaves and stems were separated and respective fresh weights determined. Then dry weights, of root3, stems and leaves were obtained by drying to constant waight i n an e l e c t r i c oven at 80° C. Since the. sterna and roots were, thick, after 24 hours in the oven, each l o t was. ground, re-weighed, and put back to dry to constant weight. A l l material brought into the labora-tory was dried, and after grinding waa used for further analy-ses • The root samples were found to contain considerable-fine grains of. sand, hence this, material was not used f o r further a n a l y t i c a l work. The ash weights or mineral content was determined on leaves and canes by heating 10-gram aliquots in a Muffle furnace: at 6.00° G. for one hour. Aliquots of sterna and. leaves were analyzed for reduc-ing sugars, sucrose, starch, and t o t a l nitrogen. Reducing sugars and sucrose were determined by the Lane and Eynon method, starch by the hydrochloric acid method, and nitrogen by the Kjeldahl method (22.). -10-Resulta A.. OBSERVATIONS OH" SAND CULTURES, There was a. marked, difference i n the response of. the raspberry plants according to their nutrient treatment. The. following deacriptions of each group are. condensed from the season's observationa. 1. Complete Nutrient Solution..- This group produced normal? vigorous growth a l l season. At the. end of the season, the canes were, about f i v e and one half feet, t a l l and heavily branched. The leaves were a healthy, dark, green colour a l l through the season. When the roots were examineds they were; found to be very w e l l developed with abundant fibrous roots and many root hairs. In every regard, this group could be. considered healthy, normal, vigorous, raspberry plants, and i s used, aa the. check in. eompariaon3 made. in. following observations. 2. Nitrogen Deficient Solution.- This group was the poorest i n the. whole series, a l l season. These plants made no perceptible gain i n growth aft e r the end of June. In July, the whole leaf surface, was d i s t i n c t l y yellow, and the bright red of the. anthocyanin pigments was very marked. By August almost the entire leaf surface was bright red with the mid-r i b and large veins showing yellow. In September defoliation began and continued rapidly, although remaining groups re-tained t h e i r leavea and many continued making new growth. The largest leaves only attained a size: about one-quarter of that of average leaves on the complete nutrient solution plant. -11-At the. end of the season* the: canes were from 6. to 12 inches high. Also, the. roots were very scanty and consisted of a few long: stringy, thin ones. 3. Phosphorus Deficient Solution.- Next to the nitrogen deficient plants, this group was the. poorest i n the series. Early i n the- season, f a i r l y r a p i d growth took place, hut not as much as made "by the check. By July the leaves were showing a marked dark colour of a bronzed, purple hue. Growth ceased in August and the purpling of the. leaves be-came more intense, and began to show on the canes. At the end of the season the plants were stunted compared to the check, and the height of the canes averaged, about 2. feet. The; canes were very th i n , weak-looking, and. an extremely dark reddish purple, colour. The leaves were a very dark green overlayed with a dark bronze-purple colour, and only about one-half or less the size. of. those of the check plants. Also, the roots were small* stunted, and generally poorly developed compared to those of. the check. 4. Potassium Deficient Solution.- This group made slower growth than the check, but the general appearance was normal. By July some, of the lower leaves showed a yellowing and dying around leaf margins and these leaves were the only ones which showed potassium deficiency symptoms. The tops continued making apparently normal growth although i t was not as vigorous as that made by checks. At the end of the season the plants were smaller than the checks, as the canes were about 4 feet high, and 1ower leaves showed de-fic i e n c y symptoms, but otherwise the plants appeared normal. -12-T.he roats were r e l a t i v e l y well developed, compared to those of the checks, and showed exceptional development of root hairs. 5:. Calcium Deficient Solution.- This group made normal vigorous growth u n t i l August and then did not continue, quite so rapidly or vigorously as the. checks. In July some lower leaves "began to show yellow streaking and. mottling, and this, condition developed over more leaves as the season ad-vanced. At the end of the season these plants were almost as large as the checks, and leaves were the same s i z e as those on the checks, but the foliage of the whole plant was a l i g h t e r green than the checks. Lower leaves were d i s t i n c t l y mottled, and streaked with yellow. The root system was excel-lent with a strong development of. fibrous roots. 6". Sulphate Deficient Solution.- In the early part of the. season, these plants presented a normal appearance but made, weaker and slower growth than the checks. In August the plants were growing very slowly, and the. foliage began to turn a l i g h t green. At the end of the; season very marked de-ficiency symptoms were, showing. Leaves were smaller than those of the checks, and were mottled yellow and green. The mottling was very f i n e , thus i t gave a general appearance, of a uniform l i g h t yellow green colour. The canes were also a very l i g h t yellow-green and averaged about 4 feet i n height. The root systems, were, w e l l developed with abundant fibrous roots. 7. Excess Nitrogen Solution.- These plants grew, at the same. rate, as the checks, and the leaves and stems were -13» about the same; size, as those of the checks. The only apparent difference -was the very r i c h , darker green colour of the leaves compared, to those of check plants, and this increased colour was noted during the whole growing period. The roots were very succulent and. fleshy, also not very extensive. There: were, p r a c t i c a l l y no, r o o t l e t s . 8. Excess Phosphorus Solution." This group was out-standingly the best i n the. series during the whole season. Prom June t i l l the end of the season the plants were t a l l e r and made more growth than any other group. Growth was normal. The size and colour of, leaves was. the same as for the checks. The canes were about 7 feet high, thus being t a l l e r than the checks. There was more, heavy branching on the canes and growth from the crown than f o r any other group. The root system was exceptionally w e l l developed, each system being larger than those i n any other group. PIbrous roots were so abundant that the root system appeared the most dense i n the entire series. 9. Excess Potassium Solution..- A l l through the grow-ing season, the plants i n this group were apparently the same as the checks i n size and colour of. canes and leaves. The roots were, well developed and about the same size: as those of the: checks but more fleshy and succulent than a l l other groups except that receiving excess nitrogen. 10. Excess Calcium Solution.- The plants i n this group grew, normally i n the early part of the season. In. August growth slowed down materially, and lower leaves, showed brown, dead margins giving the appearance of a. "scorch". The plants made very small increases i n growth, l o r the. rest of. the season and at the l a s t examination, they appeared stunted in comparison to the checks. The foliage was a l i g h t e r green than that of the checks, and more leaves, had dead, brown margins. The root systems were r e l a t i v e l y well developed, and had many fleshy, succulent roots similar to those of the excess potassium plants. 11. Complete Nutrient Solution, plus Boron, Manganese, and Z i n c - This group of plants was p r a c t i c a l l y i d e n t i c a l with the check plants. The plants grew vigorously a l l season. The roots, however, did not appear as dense and well developed as those of the check, plants. 12. Complete Nutrient Solution, plus Boron.- This group was also very similar to the checks. In July and August, the. plants made very rapid, vigorous growth and were second only to the excess phosphorous plants. However, by September increase in growth was slow., and at the f i n a l examination they were apparently the same as the check plants i n regard to height, amount and colour of fol i a g e . The. roots were dense and well developed, being similar to those of the cheeks. 13. Complete Nutrient Solution, .plus Manganese.,- This group was apparently the same as the checks with regard to size and colour of. canes and leaves. The roots, however, were scanty with more fleshy, succulent roots and fewer fibrous ones than the cheek, plants. 14. Complete Nutrient Solution, plus. Z i n c - This group was. also the same as the checks i n appearance of canes, -15-leaves and roots. 15. ' Gomplete Nutrient S'olution, plus Aluminum.- This group was also the same as the checks i n appearance of canes, leaves and roots. The deficiency symptoms exhibited by raspberry plants lacking nitrogen, phosphorus, potassium, calcium, and sulphur are. t y p i c a l of symptoms of such deficiencies as described by other workers (3, 4, 6, 19):. B". LABORATORY DATA The fresh, weights of roots and a e r i a l parts, and the root shoot r a t i o are given i n Table I. The root shoot r a t i o was determined by dividing the weight of shoots by that of the roots. The most noticeable point i n Table I i s that the excess phosphorus plants had the largest weight of. cane and l e a f , and i s an excellent t h i r d i n regard to weight of roots. The root, shoot r a t i o however does not: i l l u s t r a t e , this d i f -ference... ~16~ TABLE. I. . Fresh Weights of Hoots and Shoots, and the Root Shoot Ratio. GROUP No. of Plants Ereah. weight of. ahoata Presh weight of roots Roo t Shoot -Ratio weight of 2 Ave-rage weight of 2 Awe-rage 1. Complete 2 gm. 356-. Q gm. 178.0 gm. 279.8 gm. 139.9 1. s>l • 2 V 2. -N 4 2 2 «2. 5.5 14. Q 3-8 5 1:1.57 .,3... ~P 4 111.0 27.7 50.7 1:2.18 4. -K 2 3X34.0 152.0 188.6 94.3 1*1.61 5. -Ca 2 3,52, Q 176.0 127.0 63.5 1i2... 78 6. -S0 4 2. 138.. 0 79.0 100.7 50.4 lei.46. 7. +E 2 186.0 93.0 200.5 100.2 1:0.92 8. +-P 2 446. Q 223.0 255.5 127.8 l t l . 7 4 9. +K 2 338.0 1.69.0 210.0 105.0 Is1.61 10. •fCa •. 2 228»0 114.0 175.0 87.5 1- £ 1. • 30 11. Complete; +B,Hh.,2n 2 218.0 1Q9.0 283.7 .141.9 1:0.77 12,» it; +B: 2 352.0 176.0 216.0 10.8,0 JL s^ l. • 63 13. It •flrr 2 217.0 108.5 150.3 ? 5 e 2 It 1.44 14/.. It. •hZn 2 30 5.0 152. 5 175.4 a7.7 l t l . 7 4 15. It. +A1. :• 2/ 327.0 163.5 13.3 • 3 66.16; 1*2:. 45 The dry weight, determinations f o r the canes, leaves and roots are shown in Table I I . TABLE. I I . Percentage Dry Weight of Leaves, Cane.s and Roots. CROUP ' Percentage dry weight - Leaves^ Canea^ Roots^ 1. Complete. 29.80 45.89 45.2. z. -N — 42.. 64 51.8 3.. -P 38.29 47.83 44.5 4. -K 37.87 47.05 38.6 5. -G'a 38 • 13 47.19 46.8 6. -SO4 47.07 44.8 7. 35.80 42.88 53 ..4 8. +p 31.99 41.68 45.5 9. +K 35.09 43.39 40.. .3-10'. +Ca 32,.S9 41.29 46.2. 11. Comple.te -f-B., Mn, Zn 315.5.4 46.21 46.5 12. Complete +B. 37.09 45.12:, 42.5 13.. Complete +Hh 36«65 45.40 41.8 14. Complete ^ Zn 33.33 44.75 41. 5 15. Complete. +A1 3,7.3.8 44.2.0 36.2. The above figures appear high f o r percentages dry weight on material from raspberry plants; however, the samples were, gathered when carbohydrate reserves would, be high after a season of active growth. -18= Rapid healthy growth l a associated with low total s o l i d and high water contents (7). In Table I I , the percent-age dry weight of leaves i a low for the plants treated with complete, nutrient solution, and. excess phosphorus, with excess calcium and complete nutrient solution plus sine, also being low. Since the calcium excess plants were so obviously un-healthy, and. probably had the entrance of many nutrients inh i b i t e d , i t does not compare with others i n this group. The percentage, dry walght for canes shows plants treated with excess phosphorus, excess calcium, and nitrogen deficient to be the lowest. Of the normal vigorous plants, desired by growers, the excess phosphorus i s the. only one entering this low percentage dry weight group. The values for roots cannot be considered accurate because of the d i f f i c u l t y of removing sand especially from the. w e l l developed root systems. The percentages of ash f o r the canes and leaves are given i n Table I I I . TABLE I I I . Percentage Ash. of. Raspberry Leaves- and Canes /GROUP Canes Leaves. % ash. of dry w-eight % ash, of green •weight % ash;, of dry •weight % ash of green weight > S : % 1., Complete 3.00 1.37 5.69 1.62 2.. -It 4.60 1.96 3. ™p ' 1.08 2.11 4. 2.99 1.41 5,13: 1.94 5. ~Ca 1.83 Q.86 4.75 1.81. 6. -SO4 3.76 1.77 6.09 2.34 : ?. 2. 38 1.02 5.29 ,: 1.88 s. 2.43 1.01 6.26 2:. 00: 9. 2.87 ; 1.2.5 6.62 2_« 32. 10. +Ca 2.95 1.22.. 6.86 2.25 11. Complete +B, Mn, Zn 2.10 0.98 4.94 1.76 12. Complete +-B 2.16 0.97 5. 59 2.07 13:. Complete fMh 2.60 1»18' 5.24. 1.92 14. Complete +Zn 2.30 1.03; 6.13 &.04 15'. Complete +A1. I. 81 a. 60 6.40 2.39 Apparently no consistent correlations between growth and percentage aah can be made from the above table. The analyses of carbohydrates considered as available for the raspberry plant's l i f e processes, axe shown i n Tables W. and "V. Value3 f o r reducing sugars, sucrose, end starch are expressed i n percentage of both green and dry weights. (insert Table m.) There are no apparently outstandingly uniform, d i f f e r -ences, i n these analyses for reducing sugars, sucrose and starch. The best growing plants, were neither high nor low i n d e f o l i a t i o n , i t i s hardly to be- expected that they would give, indications of behaviour during rapid growing season. (Insert Table V.) The. excess phosphorus plants have the. highest, per-centage of reducing sugars. The poorest, groups including the potassium deficient and other deficiency groups show the low-est percentages. It i s noted that most groups of plants having high reducing sugar content have a low sucrose content. The starch figures show no consistency i n that poor and desirable plants have sim i l a r values i n these data. The t o t a l carbohydrates including reducing sugars, sucrose, and starch, t o t a l nitrogen, and the carbohydrate-nitrogen [C/s) r a t i o f o r leaves and canes are shown i n Tables VI. and: VII., respectively. TABLE IV. Percentages of Reduoing Sugars, Sucrose, and Star oh i n Raspberry Leaves. Reducing Sugars; Sucrose : Starch, • GROUP %,Of grei en weight dry . weight fo Of green weight To of dry weight •• fo Of •green weight fo Of dry weight 1. Complete 7.85 26.33 3.86 12.96 3*34 11.20 • 2 « -H 3. -P 6.10 16.30 1.83 4. 90 5.16 13.79 4. -K 4.46 11.85 1*88 5.00 . 4.60 11.01 5. -Ga 6.26 16.50 3.48 9,13 5.00 13*09 6. -so 4 6*48 16*90 4.07 10*50 4*54 11.80 7 * >I 4*98 13.86 3.18 8.86 6.16 17*03 8* +P 6.41 19.92 1.27 3.86 4.85 15.10 9. +K 8*45 24.06 1,28 3.53 7.09 20*09 10, +Ca 8.85 26.89 1.71 5.20 7.17 21.80 11. Complete4-B,Mnk2n 9*84 27.71 2.24 6.33 5.72 16.18 IE. Complete +B 6.28 16.92 1.24 3.33 5.76 15.49 13. Complete +Mn 7.87 21.42 1.08 2.93 6.79 18.56 14. Complete VAn 7.04 21.13 0,24 0.73 6,42 19,26 15. Complete +A1 8.78 23.51 1.07 4.30 6.08 16.28 TABLE V. Percentages of Reducing S ugars, Suorose, and Staroh i n Raspberry Canes. GROUP Reduoin g Sugars Sucrose v-..Staroh;;.,.. %:of v green weight % of dry weight :- % of, green weight ' °/o Of : dry ,,weight , % of green ...weight of: dry weight, -1. Complete 7*07 15.40 0.64 1.40 13,21 28.81 ~' ; Zf - I 8.53 80.00 5.56 13.85 3. -P . 6*31 13.20 5.64 1*18 10.42 21.84 4* -K , • 6.17 13.90 . '. •.—- . --- 10.68 : 22*62 5. -Ca 6.56 13.86 1.67 3«33 10*12 : 21.20 6. -SO 4 7.38 15.65 1.48 3.13 10.60 22*23/ \ 7, +B 8.26 19*00 —. . 12.50 . 29.01 8. +P 8.56 22.50 9.31 22.25 9 * +K 8.83 21*70 . • _ . — — • 8.66 ; 19*97 10 * +Ca 6*74 16.2 7 . 9.15 : 22*21 11. Complete+B, -Mn,-Zn 7*06 15*52 1.38 3.00 9.34 21.12 12. Complete *B 8.55. : 18*92 1*99 4.40 11.30 25.00 13. Complete +Mh 7.73 17.01 1.95 4.20 9.60 20.06 14. Complete +Zn 7.54 16.80 • 2.936 .5,26 9.14 20.29 15* Complete 4-A1 8.32 18*80 1.56 3.53 9,06 20.24 TABLE "O. Percentages of total. Carbohydrate, and t o t a l Nitrogen, and The C/N Ratio of Raspberry Leaves on the. Fresh "Weight Basis, SAMPLE Total earho-hydratea Total, nitrogen Ratio Complete 15.05 4.12 3 ,66 13.09 5.09 2.58 -K 1Q-.94 4.06, 2.70 • -era \ ' 14.7 & 4.0.9 5. 60 15.09 •4..S6. 3.»25 14.32. 4.67 . 3,06 +P . 12..53 3.44 4.31 : +K 16.82: 4.29 3.91 •MJa : isi; .'?a. 4.01 4.4:0 Complete +B,. Mh, Zn. 17.80 : a.ia. * 5.65 Complete. +B 1.3.28 3.74, 4.95 Complete +Mh 15.74 5»21. 3.10 Complete. +Zn 13.70 3.86 3-. .54. Complete 4A1. 15.93 4.32 3t. 6.9 In Table VI., the. excess potassium, excess, calcium and. complete nutrient boron, manganese and zinc plants have the highest percentages, of t o t a l carbohydrates. The pot-assium deficient plants have the lowest t o t a l carbohydrate (2.4). The phosphorus deficient plants, excess nitrogen, and complete nutrients plus manganese, have the highest percentages of. n i t r o -gen. Besides highest t o t a l carbohydrates, the complete nutrients plus boron, manganese, and zinc, plants have the lowest t o t a l nitrogen content, thus, giving this group the highest. C/N r a t i o . -24-A l l deficient plants and the excess nitrogen, plants have a. low ratio::. ; TAEDE VH. Percentages of Total Carbohydrate, and Total Nitrogen, and The C/N Ratio of • Raspberry Canes. = " SAMPEE: v Total carbo-hydrates Total, nitrogen C/N Ratio Complete. 2.0;.. .92, - 3.1'29-' 6*14 -N 14.09 ~£ 2&.37' 2.4FT 9 ® 32 • -K. 16.85 2® 2.3 ^»55 13.36'-' 2.06 9.09 I9.4B: . 2.44 7.95 +N 20.76 3.17 6. 54 :' *p 1.7,87 2 • 53- 6.97 •+-K' 1.7.49 .a. 79 6.28 +Ca 15.89 3128 4.84 Complete +E., Mn, Zn 17.78 3.02, 5.87 Complete *B 21.84 3.48 6.28 Complete +Mn 19.28 3.95 4.88 Complete. +Zh 19.-04 3. • 29 5.78 Complete +A1 1.8. 90 3-43: 5. 50 & Sample too small f o r analysis.. In Table VII., the complete nutrient, phosphorus de-f i c i e n t , and complete nutrient plus boron, have the highest per-centages of t o t a l carbohydrate. ' The nitrogen and potassium de-f i c i e n t plants have the lowest. The complete nutrient plus manganese plants have the highest percentage of t o t a l nitrogen, and the calcium deficient plants show the lowest. T.he excess calcium and the complete nutrient plus manganese plants have -25-the. lowest G/N r a t i o , and the deficiency groups have the highest. DISCUSS ION Observations on the raspberry planta grown i n sand cultures proved that they w i l l grow s a t i s f a c t o r i l y i f they can obtain the necessary nutrients. Aiao these same observations show, the falseness, of. an idea prevalent among growers that the Cuthbert variety has. degenerated. When able to obtain the es-sen t i a l food, elements in adequate, quantities, the Guthbert variety w i l l produce, healthy plants. The plants, grown i n the nitrogens* phosphorus, potas-sium, and sulphur deficient groups produced foli a g e symptoms similar to those found i n many f i e l d plantings, presenting further evidence that decline of. raspberry vigour i s due to n u t r i t i o n a l causes. Of those, plants grown i n the excess groups, the phosphorus plants demonstrated the. need for an excellent supply of phosphorus. Evidently many farms do not supply this nutrient i n adequate quantities. It is. reported that. Eraser "Valley s o i l s are. acid, and many are extremely acid (9). The raspberry thrives best i n a s o i l of pH. 5.2. to 6.8, according to Morgan. Thua a s l i g h t l y acid s o i l i s desirable. However, high a c i d i t y or low pH causes f i x a t i o n of phosphorus (26, 30). Thus phosphatic f e r t i l i z e r s do not prove of much benefit. On d i s t i n c t l y acid s o i l s i n Quebec, i t was found that surface applications of phosphatic f e r t i l i z e r s merely increased the readily soluble, phosphorus i n the surface one-half inch layer of. s o i l (2.9). Hence, since Eraser Valley s o i l s are acid, I t i s very possible that the plants do not have access to s u f f i c i e n t quantities of available phosphorus. The. poor growth i n the. excess calcium series was possibly due to lack of adequate: phosphorus, because this element is. f ixed as insoluble tricalcic-phosphate. Thus lim-ing to. correct acute s o i l acidity may reduce the. a v a i l a b i l i t y of -phosphorus as found i n studies by Pierre and Browning (23). There was no apparent benefit, from the use of. boron, manganese, and zinc, in the. one., season's work. Since these, elements, are needed i n such minute quantities, i t i s l i k e l y that the. plants obtained enough from the sand or pot, and thus the growth of these p a r t i c u l a r groupa this season does not preclude the possible need of. such elements on s o i l s cropped f o r a. long period of time. The. group receiving aluminum showed no toxic effects. Since McLean and Gilbert reported plant, growth, to be depressed by concentrations of 3.5 p.p.m. of aluminum (.18), i t i s evident that the t o x i c i t y of this element had been reduced. Since aluminum i s precipitated by calcium and phosphorus (18, 3.0), the aluminum content must, have been reduced from the toxic, l e v e l of. 15 p.p.m. to a non-Injurious concentration by phos-phorus and calcium i n the nutrient solution:. Since the excess phosphorus plants produced the most desirable plants i n this, experiment, I t was expected that, this superiority would, be shown i n the. laboratory work. These plants did produce the largest w.eight of leaves and. canes by a, considerable, amount. The weight of roots was a l i t t l e less, than that, for plants receiving a complete nutrient solution, and the. same solution plus, boron, manganese, and zinc. How-ever, since, phosphorus promotes root formation, especially fibrous roots (21), (25), these plants probably had a more, e f f i c i e n t root system than any o>the-r group. The. weight i s l i k e l y to be too low i n comparison to other plants because a greater proportion of. fibrous^ roots. w.oul.d be. l o s t when washing The root shoot ratios apparently show, no consistently favorable data for the best series, but i t i s interesting to note that the. phosphorus deficient plants have a smaller, amount of shoot growth, i n r e l a t i o n to the. roots, than the: excess phosphorus plants.. As noted bef ore the most actively growing plants re-ceiving adequate nutrients have a higher percentage of water or lower dry weights (.7). The data show that the plants re-ceiving complete nutrient solution, excess, phosphorus, and excess calcium, had the: lowest dry weight for leaves. Davis (4) found excess calcium to increase dry matter i n comparison to checks, hence the above, data may be. a result of a sampling error. However, since the type, of plant produced, with excess calcium i s not desired, the favorable dry weight data are. of no benefit. . Since the percentage dry weight of the excess phosphorus and complete nutrient plants i s so much lower than for other, treatments, i t appears a s i g n i f i c a n t point i n show-ing that these plants were superior to the others. In com-paring data on the dry weights of canes, the. excess phosphorus excess calcium, and nitrogen deficient plants, show the lowest percentage. Thus for the desired type of plants, once more the excess phosphorus plants are shown, up favorably. Wo- correlation between growth and percentage, ash could be. found. The worst plants, appeared almost at extremes -28-of high, and low. aah contents. The varying degrees of maturity as caused by the nutrient treatments may cause these data to be in s i g n i f i c a n t , since the amounts of minerals taken up by plants varies- according to the. stage of growth (2, 16). Also It i s possible that there i s an optimum percentage of minerals such as the: value for the excess, phosphorus plants which w i l l , pro-duce the mos t desirable, plants. Since carbohydrates are the. means of storing energy In a plant, i t i s expected that plants, with. the. greatest carbo-hydrate, reserves w i l l have the largest store of energy, and only vigorously growing plants can bu i l d up such a supply. The data i n Table I\T. give the percentages of the. carbohydrate fractions in the leaves sampled i n this work. Some, facts such as lack of potassium producing low carbohydrate content are shown (24). Since, however, the carbohydrate content of the leaf i s so variable, depending not only on n u t r i t i o n a l d i f f e r -ences, i n metabolism but also on time of. day of sampling (.20), stage of maturity, and the fact that these samples could not be collected simultaneously, the data cannot be judged on a com-parative, basis.. The results i n Table V. showing the carbohydrate frac-tions of the canes may be s i m i l a r l y regarded as not comparative for judgment. However., the carbohydrate content i s more l i k e l y to be stable i n this part of the plant according to Hicks ( l l . ) • In that l i g h t , the excess phosphorus plants showing the highest percentage of reducing sugars may be a si g n i f i c a n t point in favour of this, group of. plants. The figures for. starch seem to be too si m i l a r to allow much interpretation, and evidently the values for t o t a l available carbohydrate show, more significance, Total carbohydrate available to: plant l i f e processes and total nitrogen content of plant tissues have bean used i n an attempt to evaluate, growth responses since the I n i t i a l work of Pischer (5). I t i s customary to express, these values in the C/iT r a t i o as. done, by Kraus. and KraybilL i n their work with the tomato (14"). K r a y b l l l et a l (15) found that i t made no s i g n i f i c a n t difference to the picture whether the ratio i s expressed on a fresh, or dry weight has!3. The: data on the C/N r a t i o of. raspberry leaves and canes are presented so as to compare with, this previous work. It has been noted 'which, groups of plants had the highest and lowest G/U r a t i o s . It is interesting to note, that i n the nitrogens, phosphorus, potassium, calcium, and sulphur deficient groups, that the. leaves have the lowest ratios and the canes- have the highest ratios i n com-parison to the other plants. Hicks (.11) i n studying the c/N" r a t i o found considerable variations of the r a t i o in different organs of the plant. He found that the carbohydrate content did not vary in the stem9 however, the nitrogen did vary and more was present in the younger tissue. Also, the r a t i o varied a great deal within the leaves. Thus i t i s to be expected that the canes of the raspberry plant would be less, l i k e l y to be influenced and varied by such factors as differences i n stage of growth as the leaves. In considering the G/E ratios, i n the. canes, the deficiency group3 show high r a t i o s , and this i s to be expected (14) as Kraus, and K r a y b l l l found high G|/ir ratios accompanied by a weak vegetative condition. Also, an exceedingly low G/lT r a t i o -•30-accompanies a weakly vegetative condition (.14), and this probably accounts f o r the excess calcium plants having the lowest: r a t i o as they were adequately supplied with nutrients, yet were poor plants i n comparison to the checks. The com-plete nutrient plus manganese, plants show the second lowest r a t i o , and as these plants appeared equally as good as the checks, i t i s possible that an experimental error in sampling for analyses may account for this situation, or i t may be at the extreme end of optimum C/ft ratios for raspberry plants for making satisfactory growth. Gurjar (8) found that the C/N r a t i o of the tomato varied from extremes of 19 to 2, but that the f r u i t was borne when the. ratios were between 4. and 6.. Thus i t seems l i k e l y that the raspberry has a similar optimum r a t i o , The excess phosphorus plants did not appear l u x u r i -antly vegetative, and from the f i r s t year appearances, i t was judged that such plants would produce, the desired type of crop. The C/lT r a t i o of 6.97 is. well between the extremes of 4.84 f o r the excess calcium: plants and 9.31 f o r the phosphorus deficient plants. Thus i t seems probable that this rat i o of 6.97 may be i n the optimum C/U r a t i o range f o r raspberry plant A l l series which made good growth—complete, nutrient, excess nitrogen, phosphorus, potassium, complete nutrients plus boron manganese and zinc, and complete nutrient plus boron and zinc alone, respectively—have ratios between 5,78 and 6.97, which may be within or close to what would be the optimum range of Q/S r a t i o s f o r raspberry plants. The laboratory work does not apparently show the ex-cess phosphorus plants to be abnormal i n any way. but rather makes them appear superior. Hence., the red raspberry needs a p l e n t i f u l , supply of. phosphorus, and some, If. not most of the trouble with Eraser. Valley raspberry plantings can be overcome when a means of. supplying the plants, with, plenty of. phosphorus Is- found. Raspberry plants of. the. Cuthbert variety were grown i n sand cul.turea with f i f t e e n different nutrient treatments* Observations on these plants were: recorded. Plants showing symptoms of nitrogen, phosphorus, pot-assium, and sulphur deficiencies were sim i l a r to plants i n many commercial plantings. A l i b e r a l supply of phosphorus produced excellent plants showing that the raspberry w i l l grow i f given s u f f i c i e n t nutrients and also. the. Cuthbert variety has not degenerated. Laboratory work, further showed that the. excess phos-phorus plants were superior. Hence the problem resolves i t -self into finding a means of supplying adequate phosphorus to th.e raspberry plant. -32-Literature Cited 1. Brenchley, W. E.. On the action of. certain compounds of zinc, arsenic, and boron on the growth of plants. Ann. Bet. XXVIII: 283-302. 1914. 2. Burd, J. S;... Bate, of absorption of s o i l constituents at successive stages of plant growth. Jour. Agr. Res* XVIII.:. 51-72. 1919. 3. Davis, H. B. and H. H i l l . N u t r i t i o n a l studies with f r a -garia. I. S c i . Agr. ¥111: 631-692. 1928. 4. Davis, M. B. and H. H i l l and E. B. Johnson. Nutritional studies with fragaria. I I . S c i . Agr. XIV: 411-432. 1934. ~ 5. Elscher, H. Zur Erage der kohlensaure ernahung der pflan-zen. Gartenflora LXIV: 232-237. 1916.. 6. Eisher, P. L. Responses of the tomato i n solution cultures with deficiencies and excesses of certain essential elements. Maryland Agr. Exp. St a.. B u l l . 37 5. 1935. 7. Gruzit, On. M. and R. P. Hibbard. The influence, of an In-complete culture solution on photosynthesis. Report of Michigan Acad. S c i . XVIII: 50.-52:. 1916.~ 8. Gur j a r , A. II. Carbon nitrogen r a t i o i n r e l a t i o n to plant metabolism. Science L I ; 351-352. 1920. 9. Harris, G. H. Raspberry n u t r i t i o n . I I . Causes of rasp-berry f a i l u r e s In the. coastal area of B r i t i s h Columbia. Sci . Agr. Xyit 353-357. 1936. 10. Harris, G. H. Raspberry n u t r i t i o n . I I I . Are sulphates deficient in B r i t i s h Columbia coastal s o i l s ? S c i. Agr. XVII t. 707-711. 1937 . 11. Hicks, P. A.. Distribution of carbon/nitrogen r a t i o i n the various organs of the wheat plant at different periods of i t s l i f e , history. New. Phytol. XXVII: 108-116. 1928. 12. H i l l , H. Some fundamentals of n u t r i t i o n of h o r t i c u l t u r a l crop plants. S c i . Agr. XVI t 21-26. 1935. 13. Hoagland, D:. E. Optimum nutrient solutions for plants. Science L i l t . 562-564. 1920. 14. Kraus, E. J . and H. R. K r a y b l l l . Vegetation and reproduc-tion with special reference, to the tomato. Oregon Agr. Exp. Sta. B u l l . 1.4.9. 1918. 15. K r a y b i l l , H. R., G. R. Potter, S. W. Wentworth, P. T. Blood, and J. T. Sullivan. Some chemical constituents of f r u i t spurs associated with blossom bud formation i n the. Baldwin apple. New Hampshire. Agr. Exp. Sta. Tech. B u l l . 2.9. 1925, -33-16. Mc.Call, A. G-. and P. E. Richards. Mineral food requirements of the wheat plant at different stages of its. develop-ment. Jour. Amer. Soc. Agron. X; 12.7-134. 1.918. 1.7. McHargue., J. S. The. r o l e of manganese, i n plants. Jour. Amer. Cliem. Soc. XL IT: 1592-1598. 1922. 1.8. Mclean, For man T., and B a s i l E,. Gilbert. Aluminum t o x i c i t y I I I : 293-3,02. 1928. 19. McMurtrey, J. E. Symptoms on field-grown tobacco character-i s t i c of. the deficient supply of each of several es-s e n t i a l chemical elements. U.S.D.A. Tech. B u l l . 612. 1938. 20. M i l l e r , E. C. Daily variation of the carbohydrates i n the leaves of corn and the sorghums. Jour. Agr. Res. XXVII: 785-808. 1917. 21. IToll, Charles 3P. The effects of phosphate on early growth and maturity. Jour, Amer. Soc. Agron. XV: 87-99. 1923. 22. O f f i c i a l and tentative methods of analysis of the Associa-tion of O f f i c i a l A g r i c u l t u r a l Chemists, p. 2.4. 1935. 23. Pierre, W. H., and G. M. Browning. The temporary injurious effect of liming acid s o i l s and i t s relation to the phosphate n u t r i t i o n of plants. Jour. Amer. Soc. Agron. XXVII: 742-759. 1935. 24. Reed, H. S. The value of certain n u t r i t i v e elements to the. plant c e l l . An. Bot. XXIs 501-543. 1907. 25'. Russell, E;. J. S o i l conditions and plant growth, p. 3,5. Longmans, Green & Go. 1915. 26. Salter, Robert M. and E. E. Barnes. The efficiency of s o i l and f e r t i l i z e r phosphorus as affected by s o i l reaction. Ohio- Agr. Exp. Sta. B u l l . 553. 193,5. 27. Shive, J. W. The influence of sand upon the concentration and reaction of a nutrient solution for' plants. S o i l Science IX: 169-179. 192.0. 28. Wallace, T. Manuring of raspberries. Jour, of Pom. XVI: 3-13. 193.8. 29. Wrenshall, C. L. and R. R. McKIbbin. The u t i l i z a t i o n of native and applied phosphorus by pasture, crops. Sci. A'gr. XVIII: 60:6.-618. 1938. 30. Wright, K. E. Effects of phosphorus, and calcium carbonate. in reducing aluminum t o x i c i t y of acid s o i l s . Plant Phys . X l l t 1937. -3.4-Aokn owledgmenta The write.?: to thank Dr. @. H. Harris, Associ ate Professor of: HoatticuLture--, under whose direction this work was carried on, for his, kind supervision and valuable assistance in. outlining the Investigations, carrying- out the laboratory work, and preparing this thesis. Acknowledgment i s also given to Dr. A. 1. Bars a, Professor of Horticulture, for the kindly interest and helpful advice, given during the progress of th i s work. I 


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