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UBC Theses and Dissertations

The absorption of mineral nutrients and their effect upon the metabolism of the plant with special reference.. DesBrisay, Eileen 1934-12-31

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THE ABSORPTION OF MINERAL NUTRIENTS AND THEIR EFFECT UPON THE METABOLISM OF THE PLANT WITH SPECIAL REFERENCE TO THE TOMATO* by E i l e e n DesBrisay, A Thesis submitted f o r the Degree of MASTER OF SCIENCE IN AGRICULTURE i n the Department of HORTICULTURE. THE UNIVERSITY OF BRITISH COLUMBIA. A p r i l 1934. CONTENTS. Page I n t r o d u c t i o n . 1 Purpose of the experiment » . 1 - 2 Review of L i t e r a t u r e 2 -19 Calcium 3 - " 5 Magnesium 5 ~ 6 Sulphate .6 - 7 N i t r a t e 7 -10 Potassium ........ ..11 -14 Phosphate .14 -17 General l 8 -19 P r e l i m i n a r y Experiments 20 -2o M a t e r i a l and Methods .........20 -24 Res u l t s t a b u l a t e d .... 25 -26 Main Experiment 27 - 5 1 S e c t i o n I , Absorption Experiment ....... 28 -32 M a t e r i a l s and Methods 28 -30 R e s u l t s 30 -32 S e c t i o n I I , N u t r i t i o n Experiment ....... 40 - £ 1 M a t e r i a l s and Methods ........40 -44 Re s u l t s 44 -51 S u b s i d i a r y Experiments 52 -54 D i s c u s s i o n 55 -77 Calcium ...................... 56 - Magnesium 57 Sulphate ..................... 58 N i t r a t e 59" 62 Potassium . 6 3 - 68 Phosphate ....................69- 72 Double Concentration 73" 75 Complete S t a r v a t i o n ..........76- 77 Recommendations «... 78 Summary , 79 ""82 Acknowledgements ......................... 83 L i t e r a t u r e C i t e d ......................... 84 -89 Appendix THE ABSORPTION OF MINERAL NUTRIENTS AND THEIR EFFECT UPON THE .METABOLISM OF THE • PLANT -"WITH SPECIAL REFERENCE TO THE TOMATO INTRODUCTION. A backward glance over one hundred years shows that S i r Humphrey Davy (9) i n e a r l y work i n A g r i c u l t u r a l Chemistry found m i n e r a l c o n s t i t u e n t s to be e s s e n t i a l f o r p l a n t development. L i e b i g , i n I85O, presented h i s c l a s s i c a l Law of the Minimum, showing t h a t c e r t a i n p a r t i c u l a r minerals were necessary and that the absence of one of them retarded or prevented pl a n t growth even though a l l others should be present i n abundance. Lawes, at Sotharnsted, f o l l o w e d L i e b i g , c o n t r a d i c t i n g on some p o i n t s and progressing to f u r t h e r f i n d i n g s on others. Wolff (53) i n 1871? and again i n 1880, showed that the q u a n t i t a t i v e composi t i o n of one and the same p l a n t would vary according t o the s o i l i n which i t was grown.. Many - l a t e r workers have denied, or sub s t i t u t e d f o r , or elaborated upon the t h e o r i e s presented by these e a r l i e r ones. At the end of the century Loew (29) s t a t e d t h a t p l a n t s a b s o l u t e l y r e q u i r e a c e r t a i n minimum of each m i n e r a l nut r i e n t ; and i n most cases they take up not only an excess of these v a r i o u s compounds but a l s o substances which are perhaps useful-, but not a b s o l u t e l y necessary to the p l a n t . On these t h e o r i e s of Wolff and Loew the i n t e n t i o n of the present inves t i g a t i o n i s based. PURPOSE OF THE DIVEST IC-AT ION. The tomato, Lycopersicum esculentum M i l l . , being a pla n t w e l l adapted to n u t r i e n t c u l t u r e s , has been chosen f o r the i n v e s t i g a t i o n . I t i s purposed to f i n d when c e r t a i n ions enter the tomato pla n t and to l e a r n whether a withh o l d i n g of these ions at c e r t a i n times w i l l have an e f f e c t upon the growth and f r u c t i f i c a t i o n of the p l a n t . I t was felly, t hat f e r t i l i z e r s may o f t e n be a p p l i e d when they are unnecessary? that i s , as suggested by Loew, while the p l a n t absorbed these n u t r i e n t s , they might not be e s s e n t i a l and normal development might be obtained without them. These elements might, there f o r e , be w a s t e f u l l y a p p l i e d , or conversely, might not be s u p p l i e d at the c r u c i a l time. These n u t r i e n t m a t e r i a l s , as has been i n d i c a t e d , enter the p l a n t i n the i o n i c forms f o r instance when potassium n i t r a t e i s used as a f e r t i l i z e r the K and NO^ ions are absorbed independently. The ions to be concentrated upon are Ca, Mg, SO4, NO^, K, PO4 The ab s o r p t i o n of these s i x ions has been f o l l o w e d throughout the l i f e of the p l a n t i n the Absorption Experiment + of S e c t i o n I , and NO-^ , K, and PO^ have been e s p e c i a l l y stud i e d i n the N u t r i t i o n Experiment of S e c t i o n I I . REVIEW OP LITERATURE. P e t r i e (38) has r e c e n t l y presented a summary of the inta k e of ions by the pla n t and i t s r e l a t i o n to the r e s p i r a  t i o n of the r o o t , and l i s t s as causes amongst others f o r i o n absorption.: 1. Chemical f i x a t i o n . 2. Donnan e q u i l i b r i a i n the cytoplasm 3. The accumulation of f r e e ions i n the vacuole at the expense of energy rendered a v a i l a b l e i n r e s p i r a t i o n . 4. A d s o r p t i o n "These mechanisms operate i n each l i v i n g c e l l of the p l a n t and the r a t e of supply of ions to the shoot w i l l he determin ed "within l i m i t s by the concentrations t h a t are maintained i n the c e l l - s a p of the root c e l l s l y i n g i n the path of the trans p i r a t i o n stream. This w i l l , according to the hypothesis, depend on the rate of r e s p i r a t i o n of these c e l l s , so that w i t h i n l i m i t s the rate of r e s p i r a t i o n of the root c e l l s w i l l determine the rat e of supply of ions to the r o o t s . Some experiments w i t h oat p l a n t s i n which the roots were made to r e s p i r e at d i f f e r e n t r a t e s , and the r a t e of i o n i n t a k e there upon measured, were performed, and were found to conform w i t h the above hyp o t h e s i s . " This i s i n t e r e s t i n g i n view of the present i n v e s t i g a t i o n . A review of the t h e o r i e s and c o n s i d e r a t i o n s of c e r t a i n i n v e s t i g a t o r s on the p a r t i c u l a r ions i s i l l u m i n a t i n g . ~Qa^di;umv Calcium has.long been regarded as an e s s e n t i a l element. I t s d e f i c i e n c y w i l l cause a y e l l o w i n g and dwarfing of a l l p a r t s of the p l a n t . I t has been s a i d to be necessary f o r the formation of c e l l y/alls and thus f o r normal l e a f and root development. Some have considered i t to be necessary f o r the sy n t h e s i s of f a t s and probably of p r o t e i n s , due to i t s formation of plant soaps. N i g h t i n g a l e (35) experimenting w i t h ample calcium and calcium d e f i c i e n t tomato p l a n t s , found that i n the calcium d e f i c i e n t p l a n t s , n i t r a t e s were not ab sorbed? there was th e r e f o r e a d e f i n i t e i n t e r f e r e n c e w i t h the formation of p r o t e i n s e s s e n t i a l to the p r o t o p l a s t s of l i v i n g c e l l s * The y e l l o w i n g and dwarfing of calcium starved p l a n t s was thus accounted f o r . Since the a b s o r p t i o n of n i t r a t e s d i d not take p l a c e , there was an accumulation of carbohydrates. He suggests t h a t calcuim i s present i n the p l a n t as "combined" and "uncombined" calcium, and t h a t the l a t t e r type i s d i r e c t l y r e s p o n s i b l e f o r the calcium a c t i v i t i e s i n the p l a n t . He r e  e s t a b l i s h e s f a i r l y c o n c l u s i v e l y the f a c t of the presence of a middle l a m e l l a of calcium p i c t a t e . Calcium d e f i c i e n t p l a n t s , he p o i n t s , o u t , are s h o r t - l i v e d , the calcium present being confined to the lower p a r t s of the p l a n t . I t i s thus a non- migratory element and necessary at a l l times of growth. Colby (5) corroborates N i g h t i n g a l e , f i n d i n g that low calcium p l a n t s absorbed very l i t t l e n i t r a t e , that calcium s t a r v a t i o n prevented ab s o r p t i o n of any considerable q u a n t i t y of any i o n , and t h a t i t prevented root growth e n t i r e l y , the root t i p s i n v a r i a b l y dying. The e f f e c t of calcium i n antagonism and on c e l l p e r m e a b i l i t y would seem to be apparent here. Dorothy Day (8) f i n d s calcium d e f i c i e n t p l a n t s to be s h o r t e r , the lower leaves c h l o r o t i c , and the youngest tough and c u r l e d , but the d i f f e r e n c e i s i n a v a r i a t i o n i n the amount of elonga t i o n , r a t h e r than i n the anatomical s t r u c t u r e . She t h i n k s that many s o l u t i o n s i n use may have more than the "optimum amount of calcium", but undoubtedly c e r t a i n p l a n t s are c a l - c i p f c L i l e s and others c a l c i p h o i e s . Wallace (47) has reported on a "lime-induced c h l o r o s i s " of f r u i t t r ees growing i n s o i l s c o n t a i n i n g l a r g e amounts of calcium carbonatej since the t r o u b l e was c o r r e c t e d by a p p l i c a t i o n s of f e r r o u s sulphate sprays to the f o l i a g e , 'It would seem that t h i s c h l o r o t i c con d i t i o n i s due to the u n a v a i l a b i l i t y of i r o n " , due i n t u r n to an oversupply of calcium. Again, calcium has been found to be a co-enzyme, g r e a t l y f a c i l i t a t i n g pectase i n i t s f u n c t i o n s , Jiagne.B.iugu Magnesium i s the only mineral element i n the c h l o r o  p h y l l molecule, but as such i t holds the c e n t r a l p o s i t i o n and i s a b s o l u t e l y e s s e n t i a l i n i t s s y n t h e s i s . Being a migratory element, that i s , being capable of t r a n s l o c a t i o n , i t s l a c k may not show f o r some l i t t l e timef but f i n a l l y a spotted c h l o r o  s i s w i l l appear, beginning from the v e i n s outwards. I t i s more abundant, according to Raber (40), i n p a r t s undergoing development, and i s t h e r e f o r e thought to be necessary f o r the formation of n u c l e o - p r o t e i n s . I t seems a l s o to be a s s o c i a t e d w i t h p l a n t o i l s and i n some way to a f f e c t t h e i r s y n t h e s i s . The c a t i o n s , calcium and magnesium, perhaps because of t h e i r s t r o n g l y a n t a g o n i s t i c e f f e c t s , are often l i n k e d together. N i g h t i n g a l e (35> ) found that there was 110 marked excess nor d e f i c i e n c y of magnesium i n calcium d e f i c i e n t p l a n t s ; that i s , the absence of a b s o r p t i o n of calcium d i d not a f f e c t that of magnesium, Tyson(46) s t a t e s that i t appears t h a t "magnesium has an i n j u r i o u s e f f e c t on the plant when i t i s taken up i n amounts gqual to or greater than the amounts of calcium pre sent, even though there are f a v o r a b l e concentrations and r a t i o s • of the other elements present;" though t h i s w i l l depend on the nature of the s o i l and the f e r t i l i z e r treatments. But P f e i f - f e r and R i p p e l (39) have shown that i n oat p l a n t s , the r a t i o of magnesium to calcium may vary w i t h i n r a t h e r wide l i m i t s , w i t h no n o t i c e a b l e e f f e c t upon the p l a n t . Tyson (46) has noted the percentage of calcium i n leaves of l a r g e beets to be higher than the percentage of magnesium; whereas the per centage of magnesium i n the leaves of small beets i s higher than that of calcium. Raber (40) considers magnesium to be necessary f o r the t r a n s p o r t a t i o n of phosphorus and suggests t h a t i t i s only i n d i r e c t l y i n t h i s way a s s o c i a t e d w i t h the s y n t h e s i s of f a t s . I n t h i s connection Colby (5) has found phosphate ab s o r p t i o n to be more badly a f f e c t e d by magnesium s t a r v a t i o n than by potassium or sulphate s t a r v a t i o n . Wallace (471 i n s i s t s t h a t a proper potassium : magnesium r a t i o i s necessary i n p l a n t n u t r i t i o n . S_ulphate» The sulphate i o n i s the slowest moving of the ions according to Hoagland (19). The n e c e s s i t y of the element sulphur to p l a n t n u t r i t i o n , l i e s i n the f a c t that i t enters i n t o the composition of p r o t e i n s . Wallace (47) experimenting w i t h apip.es o and s m a l l f r u i t s , found the omission of sulphate to produce i n these p l a n t s a c o n d i t i o n resembling i n general t h a t caused by p a r t i a l n i t r a t e s t a r v a t i o n . There was a r e s  t r i c t e d shoot growth, a p a l i n g and y e l l o w i n g of the l e a v e s , f o l l o w e d by b r i l l i a n t c o l o r i n g and f i n a l l y e a r l y d e f o l i a t i o n . Colby (5) found that a sulphate omission had a depressing e f f e c t on the absorption of a l l the other elements, but that i t was l e a s t depressing on n i t r a t e a b s o r p t i o n . The c o n d i t i o n e x i s t i n g , however, might conform to 'Wallace's " p a r t i a l n i t r a t e s t a r v a t i o n " and t h e r e f o r e account f o r i t . fficllurtry (32) des c r i b e s the v e i n s as being a l i g h t e r green than the t i s s u e s between i n sulphate c h l o r o s i s . Tomato seeds c o n t a i n a consid erable amount of sulphur, but i n s u f f i c i e n t t o c a r r y the new s e e d l i n g to f u l l development. I t s d e f i c i e n c y , i t i s b e l i e v e d r e s u l t s i n a r e t a r d a t i o n of c e l l d i v i s i o n and a hindrance or suppression of f r u i t i n g . The s l o w l y moving sulphate i o n has an e f f e c t on the other ions i n slowing down t h e i r a b s o r p t i o n as Colby has shown. M..txatjag,<r The e s s e n t i a l nature of n i t r a t e i n the synthesis of p r o t e i n s and t h e r e f o r e of protoplasm, i s w e l l recognized and the y e l l o w i n g and s t u n t i n g due to i t s l a c k i s a f a m i l i a r f e a  t u r e . Kraus and I t r a y b i l l (27) have demonstrated that a l a r g e y i e l d of tomatoes i s a s s o c i a t e d w i t h an abundant supply of both n i t r o g e n and carbohydrates, and that they must be i n the c o r r e c t r a t i o . -Basing h i s work on t h e i r s , Watts (51) f i n d s that i t i s the amino-nitrate f r a c t i o n w i t h i n the plant which i s of importance i n f r u i t i n g . The percentage of n i t r o g e n w i t h s p e c i a l reference to the amino-acid f r a c t i o n , decreases when temperature or i n t e n s i t y of l i g h t or d u r a t i o n of the photosyn t h e t i c p e r i o d i s increased, thereby i n c r e a s i n g the carbohydrate content. A h i g h carbohydrate content, e s p e c i a l l y s t a r c h , i s u s u a l l y accompanied by a low amino-nitrate content, while a low carbohydrate content i s accompanied by a h i g h to moderate amino-nitrate content. P r u i t f u l n e s s i s a s s o c i a t e d w i t h a balanced carbohydrate r a t i o , g i v i n g a moderate succulence to - s - the p l a n t • He t h e r e f o r e advises the growers th a t to o b t a i n the greatest f r u i t f u l n e s s , much n i t r a t e may be used i n an abundance of l i g h t c o n d i t i o n s , such as i n the f i e l d ; whereas under l e s s l i g h t c o n d i t i o n s , such as i n the greenhouse, l e s s n i t r a t e i s needed. N i g h t i n g a l e (34) corroborates t h i s work,emphasizing that i t i s the a s s i m i l a t e d s o l u b l e compounds, that i s the amino- a c i d s , which are of importance not the n i t r a t e n i t r o g e n as such, Murneek (33) b e l i e v e s that l a r g e q u a n t i t i e s of n i t  rogenous m a t e r i a l are necessary f o r the development of a l l p a r t s of the tomato p l a n t , i n c l u d i n g the f r u i t . He goes so f a r as to s t a t e that i n the tomato "vegetative growth i s c o n t r o l l e d by the f r u i t " . The plant "apparently does not store much n i t  rogen" and "a c l o s e and d e l i c a t e balance between absorption and a s s i m i l a t i o n i s probably e s t a b l i s h e d " . The f r u i t i n some way i s able to monopolize p r a c t i c a l l y a l l the incoming or elabora ted n i t r o g e n , thus causing an evident shortage i n the s t r i c t l y v e g e t a t i v e p a r t s of the p l a n t . He considers carbohydrates may a l s o be s i m i l a r l y monopolized by the f r u i t , again causing a shortage. This shortage of carbohydrates and nitrogenous m a t e r i a l s r e s u l t s i n c e s s a t i o n of v e g e t a t i v e growth. Emmert (10) shows there i s a close c o r r e l a t i o n bet ween the n i t r a t e present i n the s o i l and i n the tomato pla n t and that t h i s i s , i n t u r n , very c l o s e l y c o r r e l a t e d w i t h y i e l d s . He a l s o s t a t e s t hat an a l k a l i n e r e a c t i o n not only stimulates n i t r i f i c a t i o n i n the s o i l , but a l s o s t i m u l a t e s both absorption and a s s i m i l a t i o n by the growing p l a n t s . He t t o o , f i n d s toma toes to be heavy n i t r a t e feeders. In an a l k a l i n e medium, as - 9 - - contrasted w i t h an a c i d one, while there i s more n i t r a t e i n the stems there i s l e s s i n the growing p o i n t s , since i t i s so r a p i d l y u t i l i z e d "by the growing t i p s . In c o n t r a s t to Eraraert, Ho ag land (18) claims t h a t n i t r a t e , u s u a l l y one of the s w i f t e r moving io n s , enters the pl a n t more slowly i n an a l k a l i n e med ium. Also working w i t h tomatoes, M a c G i l l i v a r y (3'1) found a predominant storage of s t a r c h i n p l a n t s from which n i t r o g e n had "been w i t h h e l d . He a l s o obtained higher sugars i n the f r u i t s * , McCool and Cook (30) obtained a r a p i d r a t e of t r a n s  formation of n i t r a t e n i t r o g e n i n sm a l l g r a i n s and Kentucky blue grass. This was i n d i c a t e d by a considerable decrease i n n i t r a t e content of the expressed sap three hours a f t e r the source of supply was removed. Harrison:tl|)•shows' i t : to be p o s s i b l e to k i l l Ken tucky blue grass p l a n t s , f r e q u e n t l y f e r t i l i z e d w i t h n i t r a t e s by repeated d e f o l i a t i o n s . The reason f o r t h i s was t h a t r a p i d growth exhausted the reserve carbohydrate supply and d e f o l i a  t i o n prevented a r e p l e n i s h i n g and t h e r e f o r e growth between d e f o l i a t i o n s became l e s s and l e s s . In "reference to l i g h t r e l a t i o n s h i p s / l o t t i n g h a m and Stephens (45) found w i t h young wheat p l a n t s that the n i t r a t e r a d i c l e was absorbed more f r e e l y from K M)^ i n n u t r i e n t s o l u  t i o n than from n i t r a t e s of other common metals and t h i s absorp t i o n was promoted by increases i n blue to longer u l t r a v i o l e t r a d i a t i o n s . The form of r a d i a t i o n used, they point p u t , i s d e f i c i e n t as regards the s h o r t e r r a d i a t i o n s as compared w i t h s u n l i g h t . 'i'ottingham and Lowsna (44) a l s o found r a d i a t i o n s of the h i g h e s t frequency i n the v i s i b l e spectrum and the lowest i n the u l t r a v i o l e t to promote absorption of n i t r a t e s by young p l a n t s , but t h a t y e l l o w to v i o l e t rays increased the synt h e s i s of p r o t e i n , while long u l t r a v i o l e t rays do not. Brazeale (3) considers there i s a d i r e c t r e l a t i o n between the absorption of potassium and n i t r a t e s : t h a t potas sium i s probably necessary i n the process of sy n t h e s i s of p r o t e i n - l i k e compounds, although i t does not enter i n t o t h e i r chemical composition. i f much n i t r o g e n i s a v a i l a b l e much potassium w i l l be demanded; i f l i t t l e n i t r o g e n i s a v a i l a b l e l i t t l e potassium w i l l be demanded. Davidsen (6) found i n c e r t a i n f i e l d experiments that potassium was higher i n p l a n t s from p l o t s t r e a t e d w i t h Ha NO^ than were those from the check p l o t s . Here again i s a l i n k a g e between the two* F i n a l l y i t i s i n t e r e s t i n g to note t h a t Kraus and Kray- b e l l (27) t h i n k that n i t r a t e s "may a i d i n r a p i d growth and formation of new c e l l s which have r e l a t i v e l y t h i n n e r and l e s s . l i q u i f i e d w a l l s and a gre a t e r percentage of amphoteric sub stances whose water h o l d i n g c a p a c i t y i s r e l a t i v e l y h i g h . " This would account f o r the f a c t that a high n i t r a t e supply gives a h i g h degree of succulence. - 11 - g.o:t.as.sl.urru.. Considerable research, upon the e f f e c t of potassium on tomato p l a n t s has been c a r r i e d on by Jansen and B a r t h o l o  mew and they have issued s e v e r a l b u l l e t i n s and papers thereon. They f i n d s there i s evidence that potassium i s necessary f o r c e l l d i v i s i o n s that i n a potassium d e f i c i e n c y , i t i s t r a n s  l o c a t e d and r e u t i l i z e d by the growing part of the p l a n t ; t o t a l and s o l u b l e n i t r o g e n i s much hi g h e r i n potassium d e f i  c i e n t p l a n t s ; that there i s an optimum potassium concentra t i o n which i s conducive to the normal a s s i m i l a t i o n of carbo hydrate compounds above and below which a s s i m i l a t i o n i s red uced; and they suggest that i n as much as h i g h n i t r o g e n and h i g h sugars are c o r r e l a t e d i n the blooming stage of low pot assium plants« the absence of good growth may have been due to a l a c k of condensation or p o l y m e r i z a t i o n of these com pounds to more complex forms due to the potassium l a c k (23)• Again (24) they f u r t h e r suggest "that r a p i d absorption of l a r g e amounts of potassium by p l a n t s during t h e i r e a r l y stages of growth w i t h a subsequent t r a n s l o c a t i o n and r e u t i l i - z a t i o n during the l a t e r stages of development i s , under some co n d i t i o n s , . t h e process,by which p l a n t s absorb potassium". There then f o l l o w e d a report on experiments w i t h corn, cow- peas, soybeans, oats and Soudan Grass. I t i s noteworthy t h a t i n one of these experiments w i t h one l o t of pla n t s , potassium was absorbed very slowly f o r a time and then r a p i d l y whereas the next group absorbed i t more s t e a d i l y . This same i r r e g u  l a r i t y was found i n the present i n v e s t i g a t i o n . The nature - 12 - of the c u l t u r e medium, whether water or s o i l c u l t u r e , d i d not a f f e c t the general r a t e of a b s o r p t i o n . This i s again note worthy suggesting the p r a c t i c a b i l i t y of water and sand c u l t u r e work to s o i l c o n d i t i o n s . They maintain that p l a n t s take up potassium r a p i d l y at a l l stages of growth i f a good supply i s a v a i l a b l e ; and that when the c o n c e n t r a t i o n of the n u t r i e n t i s reduced p l a n t s q u i c k l y r e a d j u s t themselves to feed on the lower c o n c e n t r a t i o n . They p o i n t out t h a t when f e r t i l i z i n g w i t h potassium, there i s no r e s i d u a l e f f e c t , since the p l a n t w i l l absorb potassium i n such l a r g e concentrations i f i t i s present i n the s o i l s o l u t i o n , but i f i t i s no longer s u p p l i e d there w i l l be t r a n s l o c a t i o n and r e u t i l i z a t i o n . They found no d e f i n i t e r e l a t i o n s h i p s between the percentage of potassium i n the p l a n t s and the simple carbohydrates and nitrogenous com pounds (2p). Potassium i s a r a p i d l y moving i o n . I t i s known to be r a d i o - a c t i v e : as such i t i s an energy producer and i t s uses t h e r e f o r e manifold. While sodium has been shown to r e  place potassium to some extent (14) i t cannot do so i n t h i s f i e l d . o f r a d i o - a c t i v i t y . Hoagland (19) f i n d s the absorption of potassium to be depressed by sodium salts© James and Penston (22) found potassium to be needed f o r greenhouse tomatoes i n d u l l weather. Haas and H i l l (15) i n c o n s i d e r i n g the work of S t o k l a s a , s t a t e the r o l e of potassium to be p a r t l y i n d i r e c t and only i n darkness does i t p l a y an important part i n p r o t e i n development. I t s r a d i o  a c t i v e q u a l i t y i s of i n t e r e s t i n t h i s connection and w i l l be - 13 - discussed l a t e r . Jansen and Bartholomew^conversely found potassium to "be taken up "by p l a n t s as r e a d i l y by night as by day. Wallace (49) and Tincker and Darbyshire (42) and numerous other workers have n o t i c e d an abnormal requirement f o r water i n potassium starved p l a n t s . This i s considered i n the d i s c u s s i o n and t h e o r i e s of such i n v e s t i g a t o r s as James (21) Weevers (52) and Warne(50) are presented t h e r e . James and Penston (22) found potassium i n a l l reg ions of the potato p l a n t , and consider that i t may be a h i g h percentage of the dry matter. I t i s e s p e c i a l l y h i g h i n a l l a c t i v e l y growing t i s s u e s , I.e., i n stem, r o o t , sprout and repr o d u c t i v e p a r t s . They can d i s c o v e r no evidence that potassium precedes and provokes growth but i t i s r e l a t e d to i t . That p l a n t s may c o l l e c t potassium they r e q u i r e f o r f u r  ther growth they s t a t e , i s a l l o w a b l e . This would be i n ac cordance w i t h Jansen and Bartholomew (24) who a l s o q u a l i f y the statement u s i n g the expression "under some c o n d i t i o n s . " James and Penston t h i n k t r a n s l o c a t i o n of potassium i n the form o f s a l t s of amino-acids or p r o t e i n s i s not u n l i k e l y . This i s i n c o n t r a s t to Brazeale (3) who t h i n k s n i t r a t e s and potassium are not chemically bound up but merely a s s o c i a t e d . The former workers base t h e i r o p i n i o n on the f a c t that potas sium and p r o t e i n s have a s i m i l a r d i s t r i b u t i o n i n the meris- tematic t i s s u e s and appear to be abundant i n sieve tubes. "The older leaves c o n t i n u a l l y l o s e potassium w h i l e at the same time theyjunger ones gain i t , and the work of Ruhland and ' - 14' - Wetzel quoted by Onslow ( 3 6 ) w i t h begonia suggests a s i m i l a r movement f o r amino-acids. K i s t o c h e m i c a l and a n a l y t i c a l evidence considered together suggest a continuous c i r c u l a t i n g . " James (21) has found potassium to be p l e n t i f u l at the surface of the p l a s t i d s , i n the c e l l vacuoles and c y t o  plasm and suggests that i n view of the marked e f f e c t of potas sium n u t r i t i o n on carbohydrate metabolism, the presence of the metal at the c h l o r o p l a s t surface i s i n t e r e s t i n g . Tincker and Darbyshire (43) agree t h a t potassium i s p o s s i b l y a c a t y l i s t f a c i l i t a t i n g the condensation and h y d r o l y s i s of s t a r c h . In working w i t h corn s t a l k s , H o f f e r (20) r e p o r t s that i n the absence of potassium, there w i l l be an accumula t i o n of i n s o l u b l e i r o n which k i l l s the p l a n t t i s s u e s . In a d d i t i o n to the n e c r o t i c e f f e c t upon the t i s s u e s "would be the suggestion that i r o n would not be present i n as l a r g e quan t i t i e s i n i t s r o l e of c a t y l i s t i n photosynthesis, i f i t were i n the i n s o l u b l e form. Phosphate.. — • ' I III III lltlfll-l || A t y p i c a l phosphorus d e f i c i e n c y w i l l show a p u r p l i n g and s t u n t i n g of growth. Wallace (47) has l a t e l y found t h i s to be true of such v a r i e d p l a n t s as s t r a w b e r r i e s , gooseberries, r a s p b e r r i e s , apples-- and many workers have found i t to be true of tomatoes and other crops. I t has been considered to be necessary i n the storage of carbohydrates, f o r the syn t h e s i s of n u c l e o - p r o t e i n s and also i t i s of consequence i n the a s s i m i l a t i o n of f a t s through the formation of phospho- l i p o r d s . M a c G i l l i v a r y (31) f i n d s phosphorus to be e s s e n t i a l • - 1 5 - . at a l l stages of growth. I f there i s a shortage, there w i l l "be a m i g r a t i o n of phosphorus to the growing t i p , and a r e - u t i l i z a t i o n of i t . But, M a c G i l l i v a r y s t a t e s , "insomuch as blossoms r e q u i r e c e l l d i v i s i o n f o r development" and f r e s h n u c l e a r m a t e r i a l s " i t would seem the l i m i t e d supply was not s u f f i c i e n t f o r a l l the organic and i n o r g a n i c phosphorus necessary". He r e p o r t s that i n a l l p l a n t s , whether amply s u p p l i e d w i t h phosphorus or not there i s a gradient of more and more phosphorus from the base of the stem upward u n t i l the f r u i t i s reached, where there i s as much as i n the r e s t of the p l a n t . He c o n t r a s t s t h i s f u n c t i o n of phosphorus wi t h t h a t of n i t r o g e n , where the highest c o n c e n t r a t i o n i s found i n the growing t i p s ; and again, whereas n i t r a t e s move r a p i d l y the movement of phosphates i s slow. I t has been contended that phosphorus i s necessary f o r seed formation. M a c G i l l i  vary p r o t e s t s t h a t t h i s seems to be true of the f r u i t as a whole and that i t i s j u s t as e s s e n t i a l to the formation of pulp as of the seeds. He p o i n t s out, however, the d i f f i c u l t y of s e p a r a t i n g pulp from seed i n the tomato. Statements that i t i s necessary to seed production are based on a n a l y s i s of such p l a n t s as wheat and corn; but the outer coats of these i s a p o r t i o n of the c a r p e l w a l l and t h e r e f o r e they are f r u i t s . He p o i n t s out that pulp i s decreased i n amount both i n the c a r p e l w a l l s and i n the c e n t r a l placenta region and that the d i f f e r e n c e s are so great that one would expect a d i f f e r e n c e i n v a r i e t y . F r u i t s grown wi t h a phosphorus d e f i c i e n c y had fewer and smaller seeds than the ample phosphorus p l a n t s , but the seeds from both had almost the same percentage of phosphorus. They gave s i m i l a r g e r m i n a t i o n t e s t s and produced s i m i l a r p l a n t s . Phosphorus d e f i c i e n t p l a n t s showed an i n  crease i n the percentage of dry weight of s t a r c h e s and t o t a l n i t r o g e n and a decrase of coaguable n i t r o g e n , and of growth, p o l l i n a t i o n , s i z e , weight and q u a n t i t y of f r u i t . G e r i c k e (11) working w i t h wheat, f e l t t h a t m a t u r a t i o n was b e n e f i t t e d by a c e s s a t i o n of phosphorus but M a c G i l l i v a r y p o i n t s out t h a t the • tomato d i f f e r s from wheat i n that t h e r e i s v e g e t a t i v e growth and f r u i t i n g at the same time and that " i t would seem essen t i a l to supply the p l a n t w i t h a continuous supply of phos phate. T h i s i s n e c e s s a r y d u r i n g f r u i t f o r m a t i o n i f good s i z e d f r u i t s are d e s i r e d " s i n c e such a l a r g e p r o p o r t i o n i s present i n the f r u i t . Andre (2 ) f i n d s that a r e u t i l i z a t i o n of phosphorus i s g r e a t e r than w i t h the other elements and t h a t t h i s may be the r e a s o n why p l a n t s have such a s m a l l t o t a l amount of phosphorus. Emmert (10) r e p o r t s the t o t a l phosphorus content of tomato leaves v a r i e d w i t h the phosphate phosphorus content of the s o i l ; i t was h i g h at a s t r o n g l y a c i d c o n c e n t r a t i o n , low i n a limed s o i l and medium i n a sodium carbonate t r e a t e d s o i l . The t o t a l phosphorus content of the f r u i t was not v e r y c o n s i s t e n t , lime a p p l i e d to the s o i l decreased i t somewhat, whi l e phosphoric a c i d d i d not. Phosphate a b s o r p t i o n was more badly depressed by magnesium s t a r v a t i o n than by potassium or sulphate s t a r v a t i o n , and a phosphate' s t a r v a t i o n s e r i o u s l y depressed n i t r a t e a b s o r p t i o n even c a u s i n g a l o s s from the - 17 - r o o t s l a t e i n the season, The Kentucky workers consider an a n a l y s i s of the p l a n t t i s s u e s f o r phosphorus, n i t r o g e n and potassium to he of more value as an i n d i c a t o r of a v a i l a b l e s o i l n u t r i e n t s than i s an a n a l y s i s of the s o i l i t s e l f . They have developed a speedy technique a p p l i c a b l e to use i n the f i e l d . S i m i l a r work has a l s o been c a r r i e d out i n the f i e l d a n a l y s i s of c o r n , s t a l k s at Purdue U n i v e r s i t y , Of p a r t i c u l a r i n t e r e s t i n t h i s i n v e s t i g a t i o n are Brazeale's f i n d i n g s that the absorption of both phosphorus and potassium increased up to a c e r t a i n c o n c e n t r a t i o n a f t e r which there was a sharp d e c l i n e . - 18 - In c o n c l u s i o n are quoted a few a d d i t i o n a l i n v e s t i g a  t i o n s of i n t e r e s t . Wallace (47) whose experiments w i t h apples, and sm9.ll f r u i t s i n sand c u l t u r e s have already been r e f e r r e d t o , shows th a t a d e f i c i e n c y of n i t r a t e , potassium, phosphorus, calcium, magnesium and sulphur produces c h a r a c t e r i s t i c e f f e c t s i n the v a r i o u s p l a n t s , a.nd the view i s expressed that some of these may be of use f o r d i a g n o s t i c purposes i n the f i e l d . Davis (7), c o n t i n u i n g Wallace's type of experiment w i t h apples, found the omission of an element was r e f l e c t e d by a low per centage of that element i n ash and dry matter, and demonst ra t e s a h i g h degree of c o r r e l a t i o n between symptoms e x h i b i t e d and the amount of the r e l a t e d element i n ash and dry matter. Gregory and Richardson (12) c l a i m r e s p i r a t i o n to be subnormal when n i t r o g e n i s w i t h h e l d , normal when phosphorus i s w i t h h e l d , and supernormal when potassium i s w i t h h e l d . They a l s o show the r a t e of a s s i m i l a t i o n to be d i s t u r b e d through the omission of these elements. A number of i n t e r e s t i n g observations are presented by Tyson (46) as a r e s u l t of h i s experiments with, sugar beets• He has found that i n the e a r l y stages of growth the r a t e of a b s o r p t i o n of elements i s greater than that of a s s i m i l a t i o n . When the p l a n t i s making r a p i d growth there i s a smaller percentage of the v a r i o u s n u t r i e n t s i n the dry matter, i n d i c a t i n g t h a t now a s s i m i l a t i o n i s g r e a t e r than ab s o r p t i o n . The l i f e processes are more i n f l u e n c e d by the r a t i o and c o n c e n t r a t i o n than by a supply of any one element other - 1 9 > than n i t r o g e n . He f i n d s l i g h t i n t e n s i t y to be important i n the u t i l i z a t i o n of m i n e r a l n u t r i e n t s but not i n t h e i r absorp t i o n : p l a n t s absorb j u s t as much i f they are shaded as unsha ded, N i t r a t e s i n c r e a s e d the absorption o f phosphorus and a l l p l a n t foods increased the absorption of potassium, The intake of calcium was not increased by potassium, phosphates or n i t r a t e s , Hoagland (18) f i n d s t h a t w i t h b a r l e y there w i l l be a b s o r p t i o n of n u t r i e n t m a t e r i a l s at a l l stages, i f a s u i t a b l e c o n c e n t r a t i o n of ions be maintained. But intense absorption d u r i n g l a t e r stages of growth l e a d to no important increases i n crop y i e l d , which seem r a t h e r to be conditioned i n a l a r g e measure by a fa.Yorg.ble supply and c o n c e n t r a t i o n i n the e a r l y stages, Heydemann, however, (16) found w i t h tomatoes t h a t the a s s i m i l a t i o n of n i t r o g e n , calcium, potassium, and phos phate proceeded at an equal r a t e f o r a time, a f t e r which potassium and n i t r a t e were used more r a p i d l y . "° 20 ** PRELIMINARY EXPERIMENT S The purpose of these experiments has already "been pointed outs that i s , an endeavor was made to l e a r n the exact period of absorption of the ions Ca, Mg, SO4., N0^, K and PO4 i n the development of the tomato, w i t h the object of l e a r n i n g the most s u i t a b l e time f o r f e r t i l i z e r a p p l i c a t i o n s . P r e l i m i  nary a b s o r p t i o n experiments were c a r r i e d out i n the greenhouse dur ing 1931"32» M a t e r i a l s ' a n d Methods» Three d i f f e r e n t s e t s of p l a n t i n g s were worked w i t h i n these experiments. The f i r s t ran from November 18, 193 1 to January 7* 1932* On November 13 twelve tomato p l a n t s of an average height of 5» 5 cm. were set up i n 2 quart g l a s s j a r s and grown i n a n u t r i e n t s o l u t i o n . These were f i r s t thoroughly washed and r i n s e d w i t h d i s t i l l e d water and covered w i t h dark paper to prevent the growth of algae. The metal tops were cut and a cork w i t h a 4 cm. bore f i t t e d i n t o the openings* The roots of the p l a n t s were r i n s e d f r e e of any s o i l from the seed bed, the stems were wrapped w i t h a small piece of absor bent cotton, and the p l a n t s were f i t t e d through the corks i n t o the j a r s . Hoagland's N u t r i e n t s o l u t i o n of the normal concen t r a t i o n was made up and the j a r s almost f i l l e d w i t h i t . A few drops of i r o n c i t r a t e wixt* added every few days and the s o l u t i o n s kept up to volume w i t h d i s t i l l e d water. A sample of the n u t r i e n t s o l u t i o n was kept f o r the subsequent comparative a n a l y s i s . - 21 - The p l a n t s were allowed to grow f o r one week, when the s o l u t i o n s were poured o f f , measured, and thoroughly mixed, and a sample taken. Upon removing the used s o l u t i o n from the j a r s , the p l a n t roots were washed o f f and a f r e s h s o l u t i o n was given them. This procedure was c a r r i e d on weekly. The sample of s o l u t i o n i n which the pla n t had grown and the sample of unused s o l u t i o n were taken to the l a b o r a t o r y and p a r a l l e l analyses run on them f o r the i o n s , - Ca, Mg, S0 A, - f • - — iiO^, E, PO^. Standard s o l u t i o n s c o n t a i n i n g known parts per m i l l i o n of these ions were a l s o analyzed and compared w i t h the check and used s o l u t i o n s . The parts per m i l l i o n of the v a r i o u s ions absorbed by the pla n t could thus be c a l c u l a t e d . The ions were determined c o l o r i m e t r i c a l l y w i t h a K l e t t Top Colorimeter:- The procedure f o r the, n i t r a t e i o n -was ••the phenol d i s u l p h o n i c method (1 ) , f o r the potassium ion the method o u t l i n e d by Cameron and P a i l y e r (4)--, phosphates were determined by the ammonium molybdenum blue method of Parkes and Pudge (37) > magnesium as o u t l i n e d by Hubbard (17), and the calcium and sulphate ions according to the procedures of Richard and Wells (41). Unfortu n a t e l y t h i s f i r s t p l a n t i n g could not be . continued®ny le n g t h of time. A c o l d n i g h t caught the p l a n t s from which set-back they never g r e a t l y recovered. In a d d i t i o n i t was not r e a l i z e d at t h i s p o i n t that a e r a t i o n was re q u i r e d i n the s o l u t i o n s to provide s u f f i c i e n t oxygen f o r the r o o t s . By January 7 so much d i s i n t e g r a t i o n of the roots was going on w i t h the consequent r e l e a s e of the ions that i t - 22 - was decided to abandon these p l a n t s . Table I , page 24, gi v e s the parts per m i l l i o n of the ions absorbed during t h i s p e r i o d . By January 7 a l l ions were being d i f f u s e d . Second p l a n t i n g s were set up i n the same manner as the f i r s t ones on January 22nd. The p l a n t s chosen were 8 cm. h i g h . Hoagland's s o l u t i o n was used f o r the f i r s t two weeks. S h i v e 1 s s o l u t i o n f o r the f o l l o w i n g three weeks and Hoagland's f o r the f i n a l week. At the time Shive's s o l u t i o n was s u b s t i  t u t e d the question of a e r a t i o n was becoming urgent and i t was f e l t t h a t a change of medium might a l l e v i a t e the n e c e s s i t y of arranging f o r the a e r a t i o n . The p l a n t s were given a f r e s h s o l u t i o n weekly as w i t h the previous p l a n t i n g , and analyses were made. Again, u n l u c k i l y , the p l a n t s were c h i l l e d and l i t t l e growth was being made. On February 20, i t was decided to use a e r a t i o n on h a l f the p l a n t s . A system was set up as f o l l o w s i - A pump was i n s t a l l e d and connected w i t h the j a r s by a system of g l a s s and rubber tubings,, The amount of a i r enter ing each j a r was c o n t r o l l e d and kept uniform by clamping the rubber t u b i n g . "A d i s t i n c t improvement i n the .appearance- of the p l a n t s was apparent but i t was found to be too l a t e to s t i m u l a t e them to normal growth, w i t h the hope of reaching m a t u r i t y i n the few weeks s t i l l l e f t . Consequently two weeks l a t e r , t h i s p l a n t i n g was a l s o discarded. whereas considerable d i s i n t e g r a t i o n had been going on before a e r a t i o n , as was i n  d i c a t e d by the a n a l y s i s , the t e s t s run f o l l o w i n g i t s a p p l i c a  t i o n showed a lower c o n c e n t r a t i o n of ions being d i f f u s e d out by the p l a n t . The f i n a l week the s o l u t i o n s from the non -» 2 3 - aerated and the aerated p l a n t s were both compared w i t h the o r i g i n a l * A n a l y s i s showed those p l a n t s which had been aera ted to be, upon the whole, h e a l t h i e r than the non-aerated. A f u r t h e r d i f f i c u l t y w i t h a l l p l a n t i n g s was root i n j u r y , due to the weekly disturbance and the washing of the roots w i t h c o l d water. Table I I , page 24, gives the a n a l y s i s r e s u l t s f o r t h i s p l a n t i n g . I t i s noted that i n the pe r i o d March 4 - 11, the f i n a l week, aerated and non-aerated r e s u l t s are given . A t h i r d p l a n t i n g was made on March 1 8 . S i x larg e p l a n t s about to blossom were set up i n g l a s s j a r s i n the manner p r e v i o u s l y employed. Hoagland's s o l u t i o n was used and a e r a t i o n was i n s t a l l e d . Again d i f f i c u l t i e s were experienced i n maintaining, the continuous f u n c t i o n i n g of. t h e pump, and i n p reventing i n j u r y and breaking of the roots when changing the s o l u t i o n s i n ha n d l i n g p l a n t s of t h i s s i z e . These p l a n t s were grown f o r f i v e weeks, and as before weekly analyses were made. Very l i t t l e growth took p l a c e , but some f r u i t was obtained. Table I I I , page 24, gives the absorption during t h i s p e r i o d . - 24 - TABLE I, N u t r i e n t s absorbed i n p.p.m. by the f i r s t p l a n t i n g . P e r i o d Nov,13-20 Eov.20 -27 Nov .27-Dec.4 Dec.4 -11 Dec.11-18 0,0 25. 62. 6 .0 30.0 K 16. 21„ 0.00 11.0 8.0 PO 4 0.0 30. 1.0 73-0 . 1.0 0.0 * 49 .0 9 .0 7.0 Hg •••: 0.0 0.0 i7»o 9.5 0.0 s o 4 + ... 0.0 0.0 0*0 - 2 5 . 8 - TABLE I I . N u t r i e n t s absorbed i n p.p.m. by the second p l a n t i n g . P e r i o d Jan . 2 2-Eeb . 5 Eeb . 5 - 1 9 Peb .19-26 Eeb.26-Mar.4 March 4 -11 Aer. Non-Aer TSQ'i 67 2*2 30.05 9.3 100 228™* K 25.8 84 . 5 - 4 5 . 27* - 1 2 . 2 0 PO 4 0 - 1 3 . 9 38. - 3 3 » - 5 3 . 6 - 7 6 . 6 Oa 3 - 1 0 . 8 - 1 5 . 7 - 7 .5 - 9 » 2 - 3 2 . 4 Mg 7.2 • - .6 - . 1 3 . 10* - 6.6 s o 4 26.3 - 1 1 . 5 - 5 0 . - - 1 0 . 8 - 2 7 . 2 TABLE I I I . . •" N u t r i e n t s absorbed i n p.p.m. by the t h i r d planting P e r i o d 1st two weeks 3rd week 4th week 5th week Average weekly NO. 97.5 0.0 0.0 100. K $ 46.3 - 1 6 . 8 42 . 5 - 2 3 . PO. • 0. 11.3 33-3 9 .1 •C«T 8 l e - 3 3 . 2 - 3 3 . 0.0 Mg 8 .5 - 3.2 0.0 9.0 S 0 4 2 .75 14 .3 0.0 11 . 5 - 25 - PJSSULTS. The r e s u l t s of the f i r s t two experiments were l a r g e l y negative, showing the a c t i o n of an unhealthy p l a n t r a t h e r than a normal h e a l t h y one. Ca, Mg, and SO were c o n s i s t e n t l y d i f f u s e d out of 4- the plant under the adverse c o n d i t i o n s . lfO^ continued to he absorbed i n some degree at a l l times, except on one occasion when the p l a n t s were d i s i n t e g r a t i n g r a p i d l y . K and PO4 d i d not act w i t h any consistency.but there was l e s s d i f f u s i n g out of the K i o n than of the PO4 from the unhealthy p l a n t s . With a e r a t i o n a n a l y s i s showed a d i s t i n c t l e s s e n i n g of d i s i n t e g r a t i o n . The t h i r d p l a n t i n g , showing the p e r i o d between blossoming and f r u i t i n g , gave the f o l l o w i n g general absorption r e s u l t s ? ITO^ was absorbed at f i r s t , but then ceased. I t was noted that the p l a n t s made l i t t l e v e g e t a t i v e growth at t h i s time, which would c o r r e l a t e the l a c k Of 10^ a b s o r p t i o n . The f i n a l week, abs o r p t i o n again occurred. The E i o n was absorbed i n l a r g e q u a n t i t i e s a l t e r n a t e l y w i t h none at a l l or even a d i f  f u s i o n out. This d i f f u s i o n was explained on the grounds that the roots were i n a d v e r t e n t l y broken i n changing the s o l u t i o n s from week to week. The PO^ i o n was absorbed i n same quan t i t y the t h i r d and f o u r week but very l i t t l e the f i f t h week. There was Ca absorption the f i r s t two weeks and t h e r e a f t e r a d i f f u s i o n out which was again accounted f o r by the breaking - 26 - of the r o o t s , and consequent d i f f u s i n g out of the i o n s . There was d i f f u s i o n out of Mg when l a r g e amounts of Ca were d i f f u s e d out, but t h i s d i f f u s i o n was fol l o w e d by renewed abs o r p t i o n . S0 A showed a f a i r l y c o n s i s t e n t absorption® - 2? - MAIN EXPERIMENT« The P r e l i m i n a r y Experiments were fo l l o w e d "by f u r t h e r a b s o r p t i o n and n u t r i t i o n i n v e s t i g a t i o n s i n 1932-1933* The Absorption Experiment concerned the ions d e a l t w i t h i n the p r e l i m i n a r y work which has been presented, that i s , Ca, Mg, SO4, NO^, K, PO4. The m a t e r i a l s and methods used and the r e s u l t s obtained are presented i n S e c t i o n I . The N u t r i t i o n Experiment concerned the e f f e c t of a d e f i c i e n c y of n i t r a t e , potassium and phosphate, a change i n con c e n t r a t i o n , and complete s t a r v a t i o n . The m a t e r i a l s and methods used are given i n S e c t i o n I I . The r e s u l t s upon p l a n t growth and f r u c t i f i c a t i o n , and a chemical a n a l y s i s are pre sented. Since i n t e r e s t i n g c o r r e l a t i o n s were found to occur, d i s c u s s i o n of the r e s u l t s of the two experiments appears under one heading. VARIETY USED. R i v e r s i d e E a v o r i t e tomato seeds were planted i n well-washed non-nutrient sand, and vrettered w i t h d i s t i l l e d water on l y . These se e d l i n g s were "used i n both the Absorp t i o n and N u t r i t i o n Experiments. - 28 - SECTION I . ABSORPTION EXPERIMENT. M a t e r i a l s and Methods«> A new method was devised f o r the abs o r p t i o n e x p e r i  ment. A pump was no longer a v a i l a b l e , and since i t had been shown i n the p r e l i m i n a r y experiments th a t aeration.;, was essen t i a l , water c u l t u r e s could not be used. A c c o r d i n g l y a s e r i e s of. sand c u l t u r e s was set up. On December 20, four of the s e e d l i n g tomato p l a n t s were planted i n f i v e i n c h c l a y pots of w e l l washed quartz sand. Means were taken to prevent the sand from l e a c h i n g out of the drainage h o l e , by covering i t w i t h an aluminum thimble an inc h i n diameter. The p l a n t s were watered w i t h Hoagland's n u t r i e n t s o l u t i o n of the r e g u l a r con c e n t r a t i o n . A check pot of sand w i t h no p l a n t i n g was al s o set up. The pots were placed upon p i n t s e a l e r s , that any d r i p p i n g s might be caught. Brown paper was wrapped around the s e a l e r s to prevent algae growth i n the d r i p p i n g s and dark paper was f i t t e d over the tops of the pots around the p l a n t s to prevent algae growth on the n u t r i e n t s a l t s i n the sand* The amount o f s o l u t i o n fed to each plant was ta b u l a t e d and the check pot watered w i t h an equal amount. P l a t e I , F i g . (a) (Appendix) shows the arrangement of the pots of the mature p l a n t s upon the s e a l e r s . A f t e r seventeen days the s o l u t i o n s were withdrawn f o r a n a l y s i s . Procedure was as f o l l o w s : D i s t i l l e d water was f l u s h e d through each pot to wash out the s a l t s which were present there. The washings were c o l l e c t e d i n crocks and - 29 - measurement was made of the s o l u t i o n which passed through each pot; s u f f i c i e n t water was used to b r i n g t h i s measurement up to twice the volume of the n u t r i e n t s o l u t i o n fed to the p l a n t . The washings of the f o u r pots were w e l l mixed and a sample was taken. The check pot was s i m i l a r l y t r e a t e d and a sample taken of the washings from i t . I t was f e l t that the use of the check pot would provide an adequate comparison as the unused s o l u t i o n , s i n c e i t was r e a l i z e d that the pots might absorb n u t r i e n t s which must not be a t t r i b u t e d to the p l a n t . The scheme f o r washing out the s a l t s i s shown i n P l a t e I , P i g . (b) (Appendix). A f t e r washing the plants through i n t h i s f a s h i o n i t was found necessary to allow them to dry out f o r a day or more before feeding the new s o l u t i o n : otherwise i t would d r i p through the pots at a l l parts and c o n t r o l of the q u a n t i t i e s of the s o l u t i o n would be l o s t • The f r e s h s o l u  t i o n was sup p l i e d i n such a qua n t i t y as the pot would take without leakage. During the f i r s t p e r i o d , which was seventeen days, each plant received ^00 c.c. The amount fed was gra d u a l l y increased u n t i l 20^0 c.c. was being given i n June, the f i n a l month. The pots were washed through and a f r e s h s o l u  t i o n given, as has been described. A f t e r two three-week pe r i o d s , and t h e r e a f t e r every two weeks, analyses of the used and unused s o l u t i o n s and comparison w i t h a known standard were made i n the manner of the p r e l i m i n a r y experiments. Measure ment of growth was taken at each change of the s o l u t i o n . A l l suckers were removed and the pl a n t s kept to one stem and staked. At the f o u r t h changing of the s o l u t i o n on March 7th, - 30 - the p l a n t s were t r a n s p l a n t e d to 6 i n c h pots, the average height at t h i s time being 49,6 em. They remained i n these u n t i l the c l o s e of the experiment on June 29th, when i t was considered v i t a l a c t i v i t y was ceasing. R e s u l t s . Tables are presented to show the absorption of the var i o u s i o n s : Table IV.- Page 3 3 : The absorption of the n u t r i e n t s i n p a r t s per m i l l i o n (p.p.m.) and i n m i l l i - e q u i v a l e n t s (m.e.) f o r the periods between a n a l y s i s . Table V. - Page 34: The abso r p t i o n of the n u t r i e n t s i n p.p.m. and m.e. du r i n g v e g e t a t i v e growth, blossoming and f r u i t i n g . Chart I . - Page 2%8The absorption of the ions i n m i l l i - e q u i v a  l e n t s charted against growth i n centimeters. I t i s notable t h a t a l l the ions are absorbed through out the l i f e of the p l a n t , Mg only, d e f i n i t e l y ceasing to enter two weeks before the others stopped. At blossoming time there was a s l i g h t f a l l i n g o f f i n n i t r a t e a b s o r p t i o n , but during an unusually f i n e p e r i o d i t went up again to almost as h i g h a point as i t had reached during the p e r i o d of purely v e g e t a t i v e growth. Potassium showed a continuous f l u c t u a t i o n i n absorption u n t i l the f i n a l month or more when i t dropped very low. Phosphate absorption increased g r a d u a l l y u n t i l blossoming time, when i t suddenly ceased f o r one p e r i o d , was low through blossoming time and increased again during f r u i t i n g . Absorption of calcium i n  creased g r a d u a l l y as e a r l y growth was being made, a f t e r which - 3 l - i t f l u c t u a t e d . A reference to Chart I , Page-1% however, shows that both calcium and n i t r a t e absorption f a i r l y w e l l f o l l o w the growth curve. Ho calcium was absorbed e a r l y i n June and then l i k e the phosphate there was suddenly a large amount absorbed the l a t t e r part of the monthj the f r u i t had been, and s t i l l was, r i p e n i n g i n l a r g e q u a n t i t i e s . Magnesium abso r p t i o n a l s o f l u c t u a t e d though there was l i t t l e c o r r e l a  t i o n w i t h calcium a b s o r p t i o n . Sulphate was absorbed i n the l a r g e s t amount e a r l y i n the v e g e t a t i v e growth but was used i n a f a i r l y uniform degree u n t i l h a r v e s t i n g of the p l a n t s . The r e l a t i v e amounts of the s i x ions absorbed i n m i l l i e q u i v a l e n t s i s a l s o i n d i c a t e d i n Chart I , Page- 29a. I t i s shown th a t n i t r a t e s are absorbed i n the l a r g e s t amounts, w i t h calcium next i n q u a n t i t y . The phosphates ions enter i n the s m a l l e s t amounts. Table V I , Page y% presents data on the r e l a t i o n of the absorption of n i t r a t e and potassium to the r a t i o of sun shine to p o s s i b l e sunshine and to the s o l a r r a d i a t i o n as measured by a black bulb thermometer. Data was obtained from the Dominion M e t e o r o l o g i c a l Service and i s therefore r e l i a b l e , but i t i s pointed out that complete s t a t i s t i c s were not a v a i l a b l e f o r the e a r l y months of the year. The averages, however, are f e l t to be r e p r e s e n t a t i v e . There seems to be very l i t t l e c o r r e l a t i o n between the elementary absorption and s o l a r r a d i a t i o n , but w i t h the r a t i o of sunshine to pos s i b l e sunshine there i s a notable c o r r e l a t i o n . During per iods of greater sunshine there i s g e n e r a l l y greater n i t r a t e - 3 2 - a b s o r p t i o n , and i n periods of l e s s e r sunshine there i s l e s s e r potassium absorption,, - '33. '- TABLE IV The absorption of the nutrients .in iparts .per million and mmi , V for the periods between analyses.- - «uiion and .milli- equivalents Hons p.p.m. m.e. p.p.m. in.e. p.p.m. m.e. /•pv-pim."''.'mVei' ;p.p.m.''m.e. 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H m o • « (4 M ft ft to ro • CM e» © S H H •rt o3 ft s S <S o ra tN-O • r a CM o © • r-i u u 3 a H r>- CM 10) eg .Sift S - P o ,Q rO CD <D 0) l> •rt +3 «3 CO ta r>-H CM H Q)-}h> M O O -rOrrt * » o a a) a5 • d o • r t M 0) Irv o O O S 4 g O O O O N O O o O C O CM © <& s o- ro CM O N rrt rrt CM O O O CM H 0 N ® e * ® O co m rO O CM rH O O O O O » e * © o- o GTN o- vo o CM H H O O O \ D O C O « ® O N ro r-i l f \ O rrt O O O O ITS o- C O ro O N CM O N CM CM no O ' O o tr\ O N O ro © * ® O co CM CM O r-i O CM O O O O o © to C O ITS O ro O CM r-i O N O on O N O © s © 1r\ O CM C O CM CM o O O O O rO co O O « & e C O ro CM o- C O G N O O O CM CM N O e ® © « CM O N CM ro t>- CM rt • r t a •rt ra to ra S3 3 ra a) « H - r t o ra ra o o •rt ft 03 O ro O o o •rt . p ft PH ?H • O ft ra S M O o • •rt 0 ft ft Pi » o ft ra rt O •rt - P CS •rt •d a CI3 r-i O - 36,- : TABES vji RESULTS OS1 N U T R I T l 6 : i r E X P E R I M E H T ; ——showing; size atiH, vrei^ht. Series Av.Ht.of Effect on Plant at the plant Change of f i r s t Treatment visible Mar. 4. First Average Average' ' Fruit height at length ." harvesting of root's in feet. in inches Average • Average - Total " g r? 6?x Hreer.: -green •weight weight : weight of tops, of roots, in grs I. l u l l Nutrient 34.375 32.75• II. -Nitrate Deficiency May 14 III. Potassium Deficiency IV. Phosphate Deficiency V. 2 x Concen tration VI. Complete Starvation Mar. 23 Yellowing '27.1 33-1 33-75 32.00 Bronzing May 16 May 19th Purple Mottling May 2 • Mar. 23 May 9 May 4 • 7*20 • 4.80 • 6.75 • 7 ..10 • 7*25 • 4 . 5 5 1 6 . 7 5 ' . 1 2 . 6 0 20.50 •21.00 22.25 13.00 Average -Average Percentage • Average,' 'Average ' ^Average I l o t ^ t - - - t " of tops of roots per plant per -fruit in grs. ; plant .:' 485.20 88.80 • 149.-00 - 33.00 333.75 54.50 477.38 69.25 429.25 92.5 137.25 35.25 574.0 182,5 388.2 546.6 522.0 I 7 2 . 5 110.00 , 22.00 24.50 4.10 77.7 .00 83.20 16.25 102.50 .14.25 26.75 4.50 77.33 83.56 76.66 . 8 2 . 5 8 76.12 80.55 2354.8 58.0 460.8 . .20.75 1550.4^ 1402.3 2269.8 407.8 27.0 34.75 36.3 15.0 X - The treatment was changed in these plants at an earlier date. 40.6 23.9 57.4 40.2 54.2 . 27.2 - ^ ^ ^ »lle those cf the-Potassi™ deficiency show a larger average sized fr u i t , there was consid.ra,!, . -TABLE VIII. - Series !. Pull Hutrlent 1 1 • titrate Deficiency III. Potassium Deficiency IV. Phosphorus Deficiency V. 2 x Concen tration VI. Complete Starvation LEAVES free reduc- Total" •-Starch i T o t a l C a ? ~ ^ S M a E § Sugars' ' V.r.o^ H-,.-.^  STEMS • "Y;' '}.'•'•'- 2.70. 2.70 £.6o 3.05 -3-50 2.50 6.4.6 5.10 2.70 5.00 6.85 6.80 3.05 6 .42 4.60 6.75 2.50 4:. 25. 11.50 ~7.7Q: 13.65 0.47 10 . 7.5 6.75 •-^ree^edu-o- '-Total .i,Stardg?Total^Ca7 1.60 3.04 1.35 1.81 1.45 2.22 10.24 9 .36; . 19.80 8.99 . I6 .72 25.71 8.56 - '8.22- 16.78 9.42 12.00 .31.82 12.72 10.95 83.67 11.20 14.72 25.92 'percentage proteins . in oven-dry weiphta ROOTS ;A. >ree reduc-,Total - . .Starch'7Total Oar" mg sugars- Sugars ./ - hohvrtrat.PS| 1.48 1.33 O.90 0.73 2.14 1.40 0 1.49 5.75 7.24 2.20 14.00 15.2Q 17.70 10.66 1.12 10.00 11.12 2.36 4.14 6,60 1.47 14.40 15.87 Leaves Stems- Roots 12.66 7.35 .12.125 .8.200 2.895 4.375 14.00 9.25 15.25 16.25 .8.62 14.75 11.62 8.25 19.81 .§•375 3.45- 19.45 - 39 T- Series I. F u l i Hutrien't I I . t i t r a t e Deficiency I I I . Potassium Deficiency IV. -^osphorus Deficiency ' V. 2 x Concen tration VI. Complete Starvation ;Total p.H. Aoidity in • Specific •'" Terms of Gravity Citric Acid. 4 . 8 6 4 . 7 ? 4 . 8 5 4 . 8 5 4 . 9 6 5-3 ..512 1.64 : ,512 •5376- .512 " .704 1.0747 1.033 1.0378 1.0271 1.059 1.057 CHEMICAL ANALYSIS Of THE:. T^tTTT. Free Reducing/Sugars' Percentage Extracted.... Green - ,';-:JDry.; . Juice .'. Weight. 'Weight 2.00 i . 8 p : 1.6,3 1.7 2.20 1 .?6 l , 8 i 16.36 1.58 18.12 1.44 16.21 1-5 17.52 1 .85 18.60 1.50 15.18 .Total Sugars in Percentage Extracted;- Sreen . .Dry. ' Juice Weight. Weight 2.19 I..92 I .83 2.08 2 . 3 I l.'ftn 1,63 1.82 1.97 T FT:) 17.08 19.2.6 1B..35 .21.26 19.60 T n" A Q Total Proteins in Percentage  Extracted Ureen ...Dry Juice,; Weight "Weight! Percent Percent Dry' Wt. 'Moisture of in Fruit. F r u i t " • 3725 .454 .818- .716 .5691 .323 2.92 11.06 . 88.94. •399 4*70 8.72 91.28 .722 8.02 8.88 91.12 .628 7.33 8.56 91.44 .486 4 . 8 3 10.05 89.95. - 40 - SECTION I I . NUTRITION EXPERIMENT. • M a t e r i a l s and Methods. N u t r i t i o n experiments were commenced on November 8. Twenty-four young tomato p l a n t s of a: h e i g h t of 4.5 cm. -were planted i n f i v e i n c h pots of well-washed quartz sand. They were to be d i v i d e d i n t o s i x s e r i e s which were to be t r e a t e d ag f o l l o w s : I . P u l l n u t r i e n t throughout growth and f r u i t i n g . I I . E u l l n u t r i e n t u n t i l blossoming time and then a n i t r o g e n omission. I I I . Potassium d e f i c i e n c y treatment. IV. P u l l n u t r i e n t u n t i l blossoming time and then a phosphate omission. V. E u l l n u t r i e n t throughout, but f e d twice the concen t r a t i o n once a week i n s t e a d of the r e g u l a r c o n c e n t r a t i o n twice a week. VI . P u l l n u t r i e n t u n t i l blossoming time and then an omission of a l l n u t r i e n t s . A l l p l a n t s were watered w i t h Hoagland's s o l u t i o n , f o u r times the r e g u l a r concentration being used f o r the f i r s t two waterings. Por the potassium d e f i c i e n c y s e r i e s ( S e r i e s I I I ) sodium was s u b s t i t u t e d f o r some of the potassium. Series I , I I , I I I , IV and VI were fed the s o l u t i o n s twice a week and Series V once a week w i t h the double concentration. Further waterings were given w i t h d i s t i l l e d water as seemed necessary. Por s e v e r a l weeks the normal c o n c e n t r a t i o n of the s a l t s was given u n t i l , on January 3, i t was doubled and was kept so f o r the remainder of the experiment. The amount of s o l u t i o n given each p l a n t was g r a d u a l l y increased from 5° c. c. to 150 c . c - 41 - The p l a n t s were kept to a s i n g l e stem and a l l suckers were removed. On January 20, the pots w i t h the potassium d e f i c i e n c y treatment ( S e r i e s I I I ) were w e l l washed through w i t h d i s t i l l e d water and a l l potassium was omitted from t h a t time on. . Sub s t i t u t i o n was made w i t h sodium according to Hoagland's d e f i  ciency s o l u t i o n s . At t h i s time the pl a n t s averaged 16 cm. i n h e i g h t . By January 31 , the f i v e s e r i e s of n u t r i e n t plants had reached an average height of 23.3 cm. and the potassium d e f i c i e n c y p l a n t s a height of 17 cm. They were thereupon t r a n s p l a n t e d to 10 i n c h pots, 14 of the pots being glazed and 10 c l a y . The p l a n t s were staked and 250 c c . of the n u t r i e n t s o l u t i o n s g i v e n . This q u a n t i t y of s o l u t i o n was fed f o r s e v e r a l waterings and was then g r a d u a l l y brought up to 400 c c . at which i t was maintained f o r the duration of the experiment. Waterings w i t h d i s t i l l e d water were given as r e q u i r e d , the c l a y pots evaporating considerably more than the glazed. The f i r s t week i n March i t was f e l t blossoming time had a r r i v e d , at which point the treatment was to be changed. March 7 the pots were w e l l washed through w i t h hose water and f i n a l l y w i t h d i s t i l l e d water, so that a l l t r aces of s a l t s might be removed. Measurement of the plants was taken. On March 10 waterings were given according to the s e r i e s out l i n e d above. Hoagland's d e f i c i e n c y s o l u t i o n s being used f o r the omission treatments. 400 c c . of s o l u t i o n , of twice the normal concentration was given twice a week as before the change of treatment? s e r i e s V received four times the con- c e n t r a t i o n once a week. P o l l i n a t i o n was secured, p a r t l y by hand and p a r t l y by tapping the stakes; a l l p l a n t s , however, were given iden t i c a l treatment. Observations were made as to the subsequent be h a v i o r of the s i x s e r i e s as regards amount of growth, d i s  c o l o r a t i o n of f o l i a g e , time of f r u i t r i p e n i n g . The f r u i t s were p i c k e d as they ripened and records of s i z e and weight were kept• P l a t e s I I , P i g . (a) and ( b ) , (Appendix), show the stand of p l a n t s i n the greenhouse at maturity. On June 20 the p l a n t s were harvested. Top and root measurements and weights were taken and the plants were spread out t o dry on the greenhouse benches. Throughout the summer they were c o n t i n u a l l y turned i n order to become tho roughly a i r - d r i e d . 'Shen q u i t e dry they were weighed and s t o r e d . T o t a l weights of the f r u i t produced were recorded and r e p r e s e n t a t i v e samples were preserved, some i n a l c o h o l and others by s t e r i l i z a t i o n i n quart j a r s . Chemical ...Analysis.. Chemical analyses were made i n the l a b o r a t o r y upon oven-dried samples of leaves, stems and roots f o r reducing sugars, t o t a l sugars, starches and p r o t e i n s . The p.H. and t o t a l a c i d i t y of the f r u i t s were taken and they also were analyzed f o r reducing sugars, t o t a l sugars, starches and p r o t e i n s . - 43 - The e x t r a c t i o n of the sugars from leaves, stems, and roots was accomplished i n the f o l l o w i n g manner? Samples which ranged from 10 to 25 grams were placed i n Erlenmeyer f l a s k s , and 100 c.c, of a l c o h o l plus 1.5 gram of Ca 00^ were added. These were b o i l e d f o r 15 minutes when the c l e a r l i q u i d was decanted o f f . This was repeated twice by adding f u r t h e r 5° c.c. po r t i o n s of a l c o h o l and again decanting the l i q u i d . A f t e r the t h i r d e x t r a c t i o n the residue was t r a n s  f e r r e d to a S a x h l e l t e x t r a c t e r ; 200 c.c. of a l c o h o l with 1 gram of Ca C0^ was used f o r the e x t r a c t i o n which continued f o r s e v e r a l hours u n t i l the l i q u i d siphoned over c o l o r l e s s . This Was added to the l i q u i d p r e v i o u s l y decanted and d i s  t i l l e d down to about 75 c.c. 200 c.c. of d i s t i l l e d water were added, the mixtures were placed on a, hot p l a t e and again brought down to about 100 c . c , care being taken both at t h i s point and during d i s t i l l a t i o n not to l e t the tem perature go beyond 9 0 ° C. The al c o h o l s now having been d r i v e n o f f the mixture of sugars and water was s t r a i n e d through cheese-cloth, and the f i l t r a t e was cleared with n e u t r a l lead acetate and deleaded w i t h sodium oxalate. A c t u a l determination of sugars was made by the Lane and Eynon Method (28) by the use of Pehling's S o l u t i o n and methylene blue as an i n d i c a t o r . This was found to give ex c e l l e n t r e s u l t s . T o t a l sugars were found by i n v e r t i n g 50 c.c. of the sugar s o l u t i o n w i t h 5 g r s . of p i c r i c a c i d . The mixture was b o i l e d e x a c t l y 10 minutes, was cooled, and n e u t r a l i z e d - 44 - w i t h 207* Ma OH, and then the sugars determined. Starches were determined by b o i l i n g the pulp remain ing from the sugar e x t r a c t i o n s i n 20 c c . of c o n e H Gl d i l u  ted.to 2 50 c c . f o r 2|- hours. The mixture was allowed to stand, was n e a r l y n e u t r a l i z e d w i t h Ha OH and f i l t e r e d . Deter mination of the r e s u l t i n g sugars was again made by the Lane and Eynon Method. Determination of the sugars of the s t e r i l i z e d f r u i t was made by s t r a i n i n g out the pulp through a cotton sack and squeezing i t very dry, then c l e a r i n g the j u i c e w i t h lead te and deleading w i t h sodium o x a l a t e . Determination was made as' w i t h the v e g e t a t i v e p a r t s . The K j e l d a h l Method of p r o t e i n a n a l y s i s was used f o r l e a v e s , stems, ro o t s and f r u i t . The p.H. of the f r u i t was taken by the quinhydrone method; and t o t a l a c i d i t y was determined by t i t r a t i o n against Ma OH using p h e n o l t h a l e i n as an i n d i c a t o r . R e s u l t s . Complete records of the morphological changes i n the p l a n t s as a r e s u l t of the s i x d i f f e r e n t treatments were kept. A report at the end of three and a h a l f weeks i s quoted: "The P u l l N u t r i e n t p l a n t s show good healthy growth w i t h an abundance of f r u i t forming. The E i t r a t e omission p l a n t s are now showing the e f f e c t s . Yellowing of the lower leaves commenced w i t h i n ten days of the change of the t r e a t  ment . The v e g e t a t i v e part of the plant i s l o s i n g i t s suc culence, though there i s growth from the upper pa r t . - 4-5 - Blossoms are s t i l l appearing and f r u i t i s being s e t , but h a r d l y more than i s the case w i t h p l a n t s deprived of a l l nut r i e n t s . Potassium d e f i c i e n t p l a n t s have made almost as good growth as the f u l l n u t r i e n t , but they look s l i g h t l y l e s s v i r i l e and s l i g h t l y l e s s f r u i t i s being borne. Blossoms appeared q u i t e as e a r l y as w i t h the other s e r i e s . So f a r the omission of phosphorus i s not n o t i c e a b l e i n any way. The p l a n t s are h e a l t h y , growing and f r u i t i n g w e l l . Those plants w i t h which the concent r a t i o n of the s o l u t i o n was v a r i e d ( S e r i e s V) from the beginning showed s l i g h t l y the best growth. There are many f r u i t s which are somewhat f u r t h e r advanced than those of the P u l l N u t r i e n t S e r i e s i n which the same amount was fed, but i n two periods instead of one© Within a week the Complete S t a r v a t i o n p l a n t s were showing some yellow i n g * There has been l i t t l e i f any f u r t h e r growth. Blossom i n g and f r u i t i n g i s continuing though the f r u i t s are not maturing as r a p i d l y as i n the P u l l N u t r i e n t S e r i e s . The whole v e g e t a t i v e part of the plant i s l i f e l e s s i n appearance." Table V I I , P a g e g i v e s the f i n a l morphological e f f e c t ' o f the v a r i o u s treatments. P l a t e I I I , f i g . ( a ) , ( b ) , ( c ) , (d), ( e ) , and ( f ) (Appendix), portray the t y p i c a l mature plant of each s e r i e s contrasted with a t y p i c a l plant of the P u l l N u t r i e n t S e r i e s . In a l l photographs the P u l l N u t r i e n t p l a n t s are designated "C" - Complete. . A comparison of the leaves of a l l s e r i e s i s given i n P l a t e IV (Appendix). Unfortunately they are not i n color - 46 - but, i n that they were photographed from the same spot, the comparison i n s i z e i s a true one. The f r a g i l e nature of the a i t r a t e d e f i c i e n c y and the Complete S t a r v a t i o n (-ALL) plan t s i s v i s i b l e . A l l leaves were cut from the same p o s i t i o n on the p l a n t and were r e p r e s e n t a t i v e ones. P l a t e V (Appendix) shows t y p i c a l roots of the plants of the s i x s e r i e s . Table V I I I , Page 37, presents the data obtained by chemical a n a l y s i s of l e a f , stem,and root^and Table IX, Page 39, presents the data obtained by chemical a n a l y s i s of the f r u i t s . The leaves of the Nitrogen d e f i c i e n c y p l a n t s con t i n u e d to y e l l o w throughout the treatment and f i n a l l y whitened u n t i l by June 2 0 ,the date of h a r v e s t i n g , they had become very f r a g i l e , w i t h the veins prominent and a p u r p l i s h c a st. Ref erence to Table V I I , Page 3&, and P l a t e I I I , f i g . ( a ) , (Appen d i x ) , shows growth, green and dry weights, the amount and s i z e of the f r u i t s to have been much l e s s than that of the F u l l N u t r i e n t p l a n t s . The leaves are smaller and there was poor root growth. None of the f r u i t s were l a r g e r than 6 cm. i n diameter but they were u s u a l l y w e l l shaped. The percentage of moisture i n the tops was r e l a t i v e l y h igh. Chemical analy s i s showed the v e g e t a t i v e parts to be hig h i n carbohydrates; e s p e c i a l l y so were the stems and roots where large q u a n t i t i e s of s t a r c h were present; reducing sugars were high i n the sterns. In the f r u i t s , the p.H. was 4 .75 more a c i d than any of the other s e r i e s . T o t a l a c i d i t y was 1.64 ( i n terms of C i t r i c Acid) which was three times as great as that of any of the other 47 - s e r i e s except the S t a r v a t i o n p l a n t s s i n which case i t was twice as gr e a t . S p e c i f i c G r a v i t y was lower than that of- the P u l l N u t r i e n t f r u i t s i n d i c a t i n g a l e s s e r amount of mineral and v o l a t i l e matter,, The dry weight i s a l s o lower. T o t a l sugars were s l i g h t l y h i g h e r than i n the check p l a n t s . Proteins were r a t h e r higher i n the f r u i t s hut low i n leaves, stems and r o o t s . In the Potassium d e f i c i e n c y s e r i e s a bronzing grad u a l l y made i t s appearance on the leaves. By June 6 a yellow i n g had appeared around t h i s bronzing and by harvest time many of the leaves so a f f e c t e d had s h r i v e l l e d . The e f f e c t of the potassium omission was t h e r e f o r e gradual. The plan t s were . s c a r c e l y l e s s t a l l than those of the P u l l N u t r i e n t Series but the green and dry weight was l e s s and there was l e s s f r u i t produced. P l a t e IV, (Appendix), shows the leaves to be some what l e s s vigorous than those of Series I . The roots show an i n t e r e s t i n g c o n d i t i o n i n being long and s p i n d l i n g , and l e a t h  ery w i t h few root h a i r s ; they were dark brown i n c o l o r . The average s i z e and weight of the f r u i t s was considerably greater than those of any other s e r i e s , there being more f r u i t s meas uri n g 6 . 5 - 7 cm. i n diameter; but, as has been noted the t o t a l f r u i t production was lower. Chemical a n a l y s i s showed a h i g h reducing sugar content i n the leaves. The roots have a low p r o p o r t i o n of both reducing and t o t a l sugars, but starches are p a r t i c u l a r l y h i g h , g i v i n g a h i g h carbohydrate f i g u r e . The p.H. and t o t a l a c i d i t y of the f r u i t j u i c e s was much as i n the c o n t r o l p l a n t s , the p.H. being 4 . 8 5 and the a c i d i t y . 5 1 2 . - 48 The S p e c i f i c G r a v i t y again was somewhat low. The percentage of dry weight of the f r u i t was l e s s than that of the P u l l N u t r i e n t f r u i t s , so that while t o t a l sugars were somewhat higher i n the Potassium d e f i c i e n c y s e r i e s when c a l c u l a t e d to dry weight, they were s l i g h t l y lower i n the extracted j u i c e . P r o t e i n content was found to be h i g h i n l e a f , stem, root and f r u i t * No e f f e c t of Phosphorus s t a r v a t i o n was apparent i n the S e r i e s IV p l a n t s u n t i l May 9 when a purple m o t t l i n g or b l o t c h i n g appeared on the leaves of the lower h a l f of one of the p l a n t s . A week l a t e r i t had made i t s appearance on two of the other s . Only j u s t before h a r v e s t i n g d i d i t appear on the f o u r t h . This m o t t l i n g crept upwards to some degree and then h a l t e d . A stro n g c o r r e l a t i o n was obtained here with the Absorption Experiment. At no time was there a se v e r e ' e f f e c t ; the p l a n t s were otherwise p e r f e c t l y healthy and f r u i t i n g went on i n an apparently normal manner. A study of Table and P l a t e s , however, shows the average dry weight of the tops to be low and the moisture content to be hig h . P r u i t production i s c o n s i d e r a b l y lower, the average s i z e i s the same but the number of f r u i t s produced per plant i s much l e s s . L i k e the Potassium d e f i c i e n c y p l a n t s there was much i r r e g u l a r i t y i n s i z e which ranged from 4 to 7 cm. This same i r r e g u l a r i t y , , however, was found i n plant s given the P u l l N u t r i e n t t r e a t  ment. P l a t e IV, f i g . ( d ) , (Appendix), shows the l e a f ; i t i s d i f f i c u l t to dis c o v e r any b l o t c h i n g and i t i s apparently stronger than the "K" l e a f . The roots were u s u a l l y l e s s _ 4 9 - extensive than those of the complete feeding, as P l a t e V, f i g . (d ) , (Appendix) i n d i c a t e s , and there were r e l a t i v e l y fewer root h a i r s . These r o o t s , too, were brown i n c o l o r . A n a l y s i s showed no s i g n i f i c a n t d i f f e r e n c e between these phosphorus d e f i c i e n t p l a n t s and the check p l a n t s i n t o t a l and reducing sugars Reducing sugars were s l i g h t l y higher i n the leaves and lower i n the roots of p l a n t s w i t h the phosphorus l a c k . The stems showed the same r e l a t i v e increase of t o t a l over reducing sug ars i n both s e r i e s . Starches were hig h i n both stems and root The f r u i t j u i c e s showed no v a r i a t i o n i n p.H. but the t o t a l a c i d i t y was s l i g h t l y higher. The S p e c i f i c G r a v i t y of the j u i c e s was lower than f o r any of the other s e r i e s and the per centage of the dry weight of the f r u i t was also lower. This w i l l be d i s c u s s e d l a t e r . Both reducing and t o t a l sugars were found to be a l i t t l e higher than the check pla n t s when c a l c u  l a t e d to dry weight but lower on the green weight b a s i s . Pro t e i n s were notably high i n r o o t , l e a f and f r u i t , but were not found so i n the stem. The p l a n t s of Series V, double the concentration fed h a l f as o f t e n , have been reported as making s l i g h t l y b e t t e r growth than the S e r i e s I check p l a n t s . Both continued strong h e a l t h y p l a n t s . At h a r v e s t i n g some of the leaves had become a p a l e r green or the normal dark green had become somewhat blotched and r e c e n t l y formed blossoms on both were s h r i v e l l i n g . The t a b l e s and p l a t e s show l i t t l e d i f f e r e n c e i n any respect except i n f r u i t production. The amount produced was almost the same, but the average number of f r u i t s on the Series I - 50 - p l a n t s was greater than on the S e r i e s V p l a n t s , but those of the l a t t e r s e r i e s were l a r g e r than those of the former. Both reducing and t o t a l sugars taken f o r the plant as a whole, that i s l e a f plus stem plus r o o t , were i n approximately the same percentage i n both s e r i e s . The F u l l N u t r i e n t plants showed a s l i g h t l y h i g h e r t o t a l sugar content i n the l e a f , and the double co n c e n t r a t i o n p l a n t s i n stem and r o o t . Both showed high t o t a l sugars i n the stem. The p.H. of S e r i e s I was 4 .86 , w h i l e of S e r i e s Y i t was 4 . 9 6 . T o t a l a c i d i t y , nevertheless was the same. P r o t e i n content was higher i n the roots and f r u i t s of the double-concentration plants5 i t was s l i g h t l y h igher i n the stems of t h i s s e r i e s but higher i n the leaves of S e r i e s I . The S p e c i f i c G r a v i t y of the j u i c e s and the dry •weight of the f r u i t s of the two s e r i e s approximated each other. The Complete S t a r v a t i o n p l a n t s e x h i b i t e d a contin uous y e l l o w i n g , w i t h c e s s a t i o n of growth. Some of the f r u i t s developed brown l e s i o n s and spots. Later the f o l i a g e became bronzed and purpled and f i n a l l y c o l o r l e s s , with a blue cast about the v e i n s , much as the f o l i a g e of the N i t r a t e d e f i c i e n c y p l a n t s . A comparison of these two s e r i e s i s shown i n P l a t e I I I , f i g . ( b ) , (Appendix). The Tables and P l a t e s show that p l a n t s which have been starved of a l l n u t r i e n t s r e f l e c t the treatment i n a much r e s t r i c t e d growth of root, stem and l e a f , and a small f r u i t production. On the other hand, the chemical a n a l y s i s evidenced reducing and t o t a l sugar content to be equal to that of the f u l l y fed p l a n t s i n root, stem and l e a f , and almost so i n the f r u i t , w i t h the exception only of a low - 51 - i n v e r t sugar content i n the l e a f of the starved p l a n t . Since the leaves of the s t a r v a t i o n plants had hegun to wit h e r , i t i s p o s s i b l e the i n v e r t sugars had passed from the leaves to the stem. The percentage of moisture i n the tops of these p l a n t s , n e v e r t h e l e s s , was greater than that i n the P u l l Nut r i e n t ones. P r o t e i n s were higher i n roots and f r u i t s of the starved p l a n t s , and higher i n the stem and leaves of the w e l l - nourished ones. The p.H. of the j u i c e s was higher f o r the s t a r v a t i o n p l a n t s , but t o t a l a c i d i t y was greater. The Speci f i c G r a v i t y of the j u i c e s and the percentage of dry weight of the f r u i t were a l i t t l e l e s s f o r the s t a r v a t i o n s e r i e s , i n  d i c a t i n g a lower mineral and v o l a t i l e matter content. - 52 - SUBSIDIARY EXPERIMENTS. (a) Complete Potassium Omission - Seeds of p l a n t s which had been grown i n 1932 with no potassium whatever were planted. Plant (a) of P l a t e VI, (Appendix), r e c e i v e d no potassium throughout i t s l i f e t i m e , whereas p l a n t (b) was f e d potassium i n the usual amounts from blossoming time. The d i f f e r e n c e i n height i s notable, though n e i t h e r a t t a i n e d any great s i z e . Both, however, bore f r u i t as i s evidenced i n the photograph. The green weight of the top of p l a n t (a) was 14.2 grams, and that of plant (b) 31.5 grams• (b) Seed Experiment - Seeds from a l l s e r i e s of the N u t r i t i o n Experiment, from the Absorption Experiment, from the 1932 Potassium omis s i o n seeds, and seeds from plant (a) of the Subsidiary E x p e r i  ment (a) were planted i n 1933* Germination t e s t s were run by p l a n t i n g the seeds i n f l a t s of w e l l washed quartz sand and watering w i t h d i s t i l l e d water only. A d e s c r i p t i o n of the seeds and the percentage of germination obtained i s given i n Table X, page 53• A second s e r i e s of f l a t s was planted s i m i l a r l y , but was fed a complete n u t r i e n t s o l u t i o n . These plants were cared f o r i n the manner of the N u t r i t i o n Experiment and are now i n ten i n c h glazed and clay pots. P l a n t s whose parents and - grand-parents had no potassium are s l i g h t e r stemmed and - 53 ~ T A B L E X s D e s c r i p t i o n and Percentage Germination of geeds i n S u b s i d i a r y E x p e r j m e ^ t J ^ D e s c r i p t i o n and Si z e Percentage Germination P u l l N u t r i e n t taken as normal 58 2 x Concen t r a t i o n as F u l l N u t r i e n t but more wringled 81 Potassium d e f i c i e n c y as P u l l N u t r i e n t but depressed 54 Potassium omiss ion 193 2 seeds s l i g h t l y smaller but uniform i n s i z e 68 Potassium omission 1933 seeds smaller than 1932 parents and more v a r i a b l e 65 Phosphate d e f i c i e n c y s l i g h t l y smaller and l e s s uniform 49 N i t r a t e d e f i c i e n c y s m a l l e r and l e s s uniform i n s i z e and shape 66 Complete s t a r v a t i o n s m a l l e r and l e s s uniform i n s i z e and shape 77 Absorption P l a n t s smaller than F u l l N u t r i e n t but uniform 74 - 54 - r a t h e r l e s s vigorous than those whose parents only were depriv ed, and these l a t t e r i n t u r n , are only s l i g h t l y l e s s so than the p l a n t s from the seeds of the Potassium d e f i c i e n c y p l a n t s of the N u t r i t i o n Experiment. The p l a n t s from the seeds of the other give s e r i e s of the N u t r i t i o n Experiment show a more or l e s s uniform growth. Seedlings from the Absorption Experiment had a much b e t t e r r o o t system than any others, and i t i s n o t i c e a b l e that the p l a n t s are making b e t t e r growth. The e f f e c t of s t a r  v a t i o n i n one generation has l i t t l e permanent e f f e c t . (c) Phosphate and Potassium^Omission - Seed l i n g tomatoes from seeds of p l a n t s completely starved of phosphates and from seeds of potassium d e f i c i e n t p l a n t s are being grown. They are being e n t i r e l y starved of the n u t r i e n t i n ques t i o n . H i s t o l o g i c a l examinations of these and the potassium range i n Experiment (b) are being c a r r i e d on. (d) Ir o n D e t e c t i o n i n Potassium.; Starved P l a n t s - A t e s t f o r i r o n deposits i n the t i s s u e s of p l a n t s t r e a t e d w i t h a potassium omission according to the method des c r i b e d by H o f f e r (20) were performed. P l a n t s from seeds of the Potassium d e f i c i e n c y s e r i e s of the N u t r i t i o n Experiment which were fed no potassium were used. In the absence of potassium, i r o n i s b e l i e v e d to accumulate i n that i t becomes i n s o l u b l e . A very d i s t i n c t browning of the t i s s u e s r e s u l t e d i n d i c a t i n g t h i s accumulation. when seeds of the various potassium d e f i c i e n t treatments being grown i n S u b s i d i a r y Experiment (b), now being fed a complete n u t r i e n t were t e s t e d , no brown d i s c o l o r a t i o n appeared. - 55 - DISCUSSION. The f i n d i n g s of the i n v e s t i g a t i o n would seem to i n  d i c a t e that P e t r i e (38) i s c o r r e c t i n h i s theory that the rate of r e s p i r a t i o n of the root c e l l s of the p l a n t w i l l determine the r a t e of absorption of the i o n s . Evidence points c l e a r l y to the f a c t that the p l a n t absorbs the n u t r i e n t s which i t s metabolism r e q u i r e s . I t i s impossible to agree w i t h Loew ( 2 9 ) that i n most cases they take up not only an excess, but also substances which are "perhaps u s e f u l , but not a b s o l u t e l y neces-. sary to the p l a n t " . Absorption of the ions s t u d i e d was found to take place throughout the l i f e of the p l a n t , and a strong c o r r e l a  t i o n was obtained between the Absorption and N u t r i t i o n E x p e r i  ments. A d i s c u s s i o n of the i n d i v i d u a l ions i s presented, when these c o r r e l a t i o n s w i l l be pointed out. Calcium, magnesium and sulphate were worked w i t h only i n the absorption e x p e r i  ment so that statements cannot be made regarding t h e i r essen t i a l n ature. There were no p l a n t s i n the N u t r i t i o n Experiment which-were w i t h h e l d these ions. But such a strong c o r r e l a t i o n was obtained between the two experiments i n regard to the other ions that i t would seem p e r m i s s i b l e to suppose that these too were probably being absorbed i n only the required amounts. - pb - Calcium. In the P r e l i m i n a r y Experiments 1 and 2, the d i f f u s i o n of calcium i n t o the s o l u t i o n i s accounted f o r by the d i s i n t e g  r a t i o n of the p l a n t s , when the c e l l s i n separating would r e l  ease the calcium of the middle l a m e l l a . In the t h i r d P r e l i m i  nary Experiment, the e a r l y absorption during growth would be expected since calcium i s re q u i r e d i n the b u i l d i n g up process and i s of importance i n the absorption of the other e s s e n t i a l elements (Co l b y ) , when these p l a n t s began to d i s i n t e g r a t e there was again a d i f f u s i o n out. In the Ma.in Absorption Experiment there was a gradual increase of absorption during e a r l i e r growth. The calcium was re q u i r e d f o r the d i v i s i o n of the c e l l i n the formation of the middle l a m e l l a , and f o r normal root and l e a f growth. The e f f e c t of calcium on n i t r a t e absorption i s an i n t e r e s t i n g p o i n t . I t i s noted that Dorothy Day (8) t h i n k s there to be too much c a l  cium i n some n u t r i e n t s o l u t i o n s . However the importance of calcium i n c e l l p e r m e a b i l i t y , through i t s e f f e c t on other ions, and as a c o r r e c t o r of poor s o i l c o n d i t i o n s , cannot be over estimated. No calcium was absorbed e a r l y i n June when growth had reached i t s maximum, but the renewed absorption i n large q u a n t i t i e s when f r u i t i n g was at the maximum opens up an i n t e r  e s t i n g f i e l d . One wonders i f calcium i s present i n large amounts i n the f r u i t s , and i t i s suggested that a mineral a n a l y s i s of the f r u i t f o r calcium would be of value i n e x p l a i n  in g t h i s renewed a c t i v i t y i n the absorption of calcium. Such i s l a t e r shown to be the case i n regard to phosphates a l s o . - 57 - Magnesium* During the adverse c o n d i t i o n s of the f i r s t two P r e l i m i n a r y Experiments there was a d i f f u s i n g out which would he accountable to breaking down of the c e l l s and d i s i n t e g  r a t i o n of chlorophyll,, When absorption d i d occur, as i t d i d i n s e v e r a l p e r i o d s , magnesium entered i n f a i r l y l a r g e amounts to be u t i l i z e d i n the b u i l d i n g up process. In the t h i r d P r e l i m i n a r y Experiment, magnesium absorption f l u c t u a t e d : i t was d i f f u s e d out when calcium was d i f f u s e d out i n l a r g e q u a n t i t i e s . Whether there i s a l i n k a g e here i s u n c e r t a i n . In the Main Absorption Experiment, there was again a f l u c t u a t i o n i n magnesium absorption. One might hope to e x p l a i n i t on the grounds of c h l o r o p h y l l development i n the photosynthetic r e a c t i o n s , but there appears to be no cor r e l a t i o n with s u n l i g h t , as one would expect, i f such were the cause. The c o r r e l a t i o n of growth and magnesium absorp t i o n , however, i s f a i r l y w e l l defined, as a reference to Chart I (Pag-£...29a), w i l l show. In the e a r l y stages of growth more p r o p o r t i o n a t e l y was required than at any other time. R e c a l l i n g the migratory q u a l i t y of magnesium t h i s would seem reasonable. - 58 - Sulphate. L i k e calcium and magnesium, the sulphate i o n , on the whole, was c o n s i s t e n t l y d i f f u s e d out of the plant under the co n d i t i o n s of P r e l i m i n a r y Experiments 1 and 2. Indeed i t was even more continuously d i f f u s e d out. In the t h i r d P r e l i m i n a r y Experiment i t was, i n a l l hut one p e r i o d , c o n s i s t e n t l y absor bed. In the Main Absorption Experiment sulphate was absorbed f a i r l y u n iformly throughout the whole period of growth and f r u c t i f i c a t i o n . The l a r g e s t amount of absorption occurred d u r i n g v e g e t a t i v e growth, however, and again during the heavy r i p e n i n g of f r u i t s . Since i t enters i n t o the com- •\"f p o s i t i o n of p r o t e i n s , i t would be c o n s t a n t l y required as long as growth was going on. The l i t e r a t u r e has shown i t to be present i n the seed, and i t would seem that the f r u i t must co n t a i n a considerable q u a n t i t y . Again a chemical a n a l y s i s of the f r u i t would be v a l u a b l e ; the p e r s i s t e n t absorption of sulphate through a l l phases of development and i n f r u i t i n g , would suggest t h a t i t must be present i n some qua n t i t y i n the f r u i t . - 59 - g i t r a t e s , N i t r a t e s , i t would seem from the data of both the N u t r i t i o n and Absorption Experiments, are being constantly u t i l i z e d by the p l a n t as long as growth and f r u i t i n g i s prog r e s s i n g . I f the l i f e processes of the tomato are s i m i l a r to those of the small g r a i n s , and the r a t e of transformation of n i t r a t e n i t r o g e n i n the p l a n t i s very r a p i d as shown by McCool and Cook ( 3 0 ) , i t would l o g i c a l l y f o l l o w that a constant sup pl y of n i t r a t e must be maintained that protoplasm might be b u i l t up. The consequences of the w i t h h o l d i n g of n i t r a t e from the S e r i e s I I p l a n t s , t h e r e f o r e , were the expected ones i n l a c k of growth and d i s c o l o r a t i o n of the f o l i a g e . The leaves have l o s t the power of c h l o r o p h y l l formation. In contrast to the f r a g i l e pale lavendar of the l e a f of the tomato i n the f i n a l stages, MacMurtrey ( 3 2 ) w i t h the tobacco plant obtained a y e l l o w i n g f o l l o w e d by a " f i r i n g " of the lower leaves to a b r i g h t brown c o l o r . N i g h t i n g a l e ( 3 4 ) i n contrast to McCool and Cook and to Murneek, t h i n k s n i t r a t e s may be stored w i t h i n the p l a n t u n t i l the proper co n d i t i o n s a r i s e f o r synthesis to other forms of n i t r o g e n . I f t h i s i s so the c e s s a t i o n of n i t  r a t e absorption i n the t h i r d p l a n t i n g of the P r e l i m i n a r y Ex periment could be accounted f o r . There was l i t t l e growth during two p e r i o d s , p o s s i b l y <doae to adverse c o n d i t i o n s , and l i t t l e n i t r a t e was needed. As f r u i t i n g commenced more n i t r a t e s would be r e q u i r e d , according to Murneek ( 3 3 ) , and there was a f u r t h e r absorption of them. - 60 - The l i t e r a t u r e c i t e d i n the i n t r o d u c t i o n by T o t t i n g - ham and Stephens (4-5) and Tottingham and Lowsna (44) on the i n f l u e n c e of sho r t e r l i g h t rays i s of intense i n t e r e s t i n reference to the increased absorption of n i t r a t e s on b r i g h t days, Table VI, page 35» I t has been mentioned that u t i l i z a  t i o n of n i t r o g e n i s very r a p i d ; we know photosynthesis to be c o n t i n u a l l y going on; and have often observed, the r a p i d i t y and suddenness of growth almost and i n t r u t h overnight, f o l l o w i n g sunny weather, when there would, be much carbohydrate manufac tured and much n i t r a t e enter the p l a n t . I t would seem, there f o r e , that there i s an immediate response i n metabolism to an adequate n i t r a t e and carbohydrate supply. The potassium and n i t r a t e r e l a t i o n s h i p i s i n t e r e s t i n g i n t h i s connection, i n t hat l e s s potassium i s absorbed on b r i g h t days• This w i l l be more f u l l y discussed under potassium, but i t i s suggested here that potassium possesses some k i n e t i c or e l e c t r i c a l energy a k i n to l i g h t rays, which make i t s absorption i n large amounts unnecessary during b r i g h t weather. Brazeale's (3) c l a i m that the supply of n i t r o g e n i s the l i m i t i n g f a c t o r i n the absorption of potassium would s t i l l be a v a l i d one under t h i s hypothesis, though l e s s e r amounts would be required during sunshine. That n i t r a t e s should be more r e a d i l y a v a i l  able from a KNO^ s o l u t i o n than from other n i t r a t e s o l u t i o n s i s an i n t e r e s t i n g p o i n t . Both K and NO-^  however are r a p i d l y moving ions and would have some e f f e c t one upon the other. We obtained l i t t l e c o r r e l a t i o n between temperature and n i t r a t e and potassium absorption as evidenced by So l a r R a d i a t i o n , - 61 - Table V I , page 35, while Tottingham d i d . As i s c o n s i s t e n t w i t h Kraus and K r a y b i l l ' s work, f r u i t p roduction was very low, but the percentage of moisture was r e l  a t i v e l y h i g h . The h i g h s t a r c h content of stems and roots would also-be i n accord w i t h ltraus and K r a y b i l l ' s theory. The accu mulation of these carbohydrates would be due to the n o n - u t i l i z a  t i o n of the manufactured sugars of the leaves, now stored as s t a r c h i n stem and r o o t , P r o t e i n s n a t u r a l l y were low. Here there i s a low p r o t e i n content, which would i n d i c a t e a low- amino n i t r a t e content, balanced against a h i g h carbohydrate supply, which has r e s u l t e d i n a low f r u i t production. Of i n t  e r e st at t h i s point i s Harrison's work (13) already r e f e r r e d to where, by f r e q u e n t l y c u t t i n g o f f the carbohydrate supply and s t i l l m a i n t a i n i n g a h i g h n i t r a t e supply, Kentucky Blue Grass p l a n t s were a c t u a l l y k i l l e d . By lowering the n i t r a t e sup pl y and thereby keeping a b e t t e r balance the plants continued to send up rhizomes which t i l l e r e d . Murneek's c l a i m that the f r u i t of the tomato draws on the n i t r a t e and carbohydrate supply i s probably a v a l i d one« Though f r u i t production was low, they continued to develop a f t e r n i t r a t e s were w i t h h e l d , and were q u i t e as high i n pro t e i n s and sugars as were the P u l l N u t r i e n t p l a n t s . The h i g h a c i d i t y i n the f r u i t s might be considered to be due to a p o s s i b l e calcium-nitrate r e l a t i o n s h i p . I t has been shown by N i g h t i n g a l e and h i s co-workers (3^) that n i t r a t e s are not absorbed i n the absence of " f r e e " calcium. P o s s i b l y the opposite c o n d i t i o n may a l s o be t r u e . The "free calcium" might have a n e u t r a l i z i n g e f f e c t on the organic a c i d s . An - 6 2 - a n a l y s i s of the f r u i t s f o r calcium would he of i n t e r e s t , i n order to l e a r n i f a n i t r o g e n d e f i c i e n t plant also contains l e s s calcium. — 6 3 " Potassium. The r e l a t i o n s h i p of n i t r a t e and potassium absorption has already been discussed and reference made to Table V I , page 3 ! ? > showing the e f f e c t of s u n l i g h t on t h e i r absorption; i t has been pointed out that l e s s e r amounts of potassium are absorbed on b r i g h t days. This corroborates James and Penston's f i n d i n g s , though r e f u t i n g those of Jansen and Bartholomew (24). I t i s suggested that the reason f o r t h i s i r r e g u l a r absorption may be found i n the f a c t that potassium i s r a d i o - a c t i v e , and that an important part of i t s f u n c t i o n i n the plant i s such, when the sun s u p p l i e s t h i s energy, l e s s e r amounts of potassium are r e q u i r e d . The p l a n t has adjusted i t s e l f to t h i s s i t u a t i o n . In f a c t , i t would seem that p l a n t s are not merely the creatures of casual circumstance, absorbing m a t e r i a l s whether they are u s e f u l or not as i s sometimes thought, but that they have devel oped during t h e i r long existence a working r e l a t i o n s h i p toward outward c o n d i t i o n s . This, and other phases of the Absorption Experiment which w i l l be pointed out l a t e r most c l e a r l y point to the plant absorbing only the n u t r i e n t s which i t r e q u i r e s . I t was - shown i n the Absorption Experiment that growth and n i t  r a t e absorption were s t r o n g l y c o r r e l a t e d and the N u t r i t i o n Experiment bore t h i s out. Now a c o n s i d e r a t i o n of potassium and i t s r a d i o - a c t i v e q u a l i t y evidences that the t h r i f t y plant ab sorbs l e s s potassium i n f i n e weather. Temperature, as recor ded by the black bulb thermometer, was without e f f e c t . The l i t e r a t u r e shows, however, that James and Penston (22) and Jansen and Bartholomew (24) t h i n k that plants may c o l l e c t and - 64 - store potassium. They both q u a l i f y t h e i r statements, however. Undoubtedly the absorbed potassium must be r e u t i l i z e d as w i l l be shown l a t e r . This p e c u l i a r r a d i o - a c t i v e q u a l i t y of potassium i s als o outstanding i n c o n s i d e r a t i o n of the water-holding q u a l i t  i e s of p l a n t s . Many workers have noted that potassium d e f i c i e n t p l a n t s r e q u i r e much more water than do those with a normal sup p l y . The water i s q u i c k l y t r a n s p i r e d and the leaves soon w i l t . A d i g r e s s i o n at t h i s p o i n t to a c o n s i d e r a t i o n of a recent paper by Shu11 (42) i s i l l u m i n a t i n g : He points out that the r e l a  t i o n between root and s o i l and s o i l water i s a dynamic one, and that i t i s i n the water i t s e l f that t h i s dynamic f o r c e p r i n c i p a l l y r e s i d e s . A water d e f i c i t which reduces vapor pres sure of the c e l l c o l l o i d s by 7% below that of pure water i s b e l i e v e d to develop f o r c e s close to 100 atmospheres. "The c e l l may not, f o r some reason develop s u f f i c i e n t l y high forces to a t t r a c t the water, which passes on up the t r a n s p i r a t i o n stream." A water d e f i c i t may e x i s t i n the root when the t e n s i o n a l p u l l of the water column i n the tracheae draws the water more r a p i d  l y from the l i v i n g c e l l s i n the xylem than i t can be supplied by t r a n s f e r of the water from the epidermis across the i n t e r  vening c e l l s to the p e r i c y c l e . This p u l l may exceed the force of osmotic d i f f u s i o n and the hindrances of the c e l l w a l l and protoplasm. The forces which are important i n determining the osmotic pressure of the c e l l are p h y s i c a l , chemical, c o l  l o i d a l , or e l e c t r i c a l . I m b i b i t i o n also plays an important r o l e i n supplying the plant w i t h water and i n the t r a n s f e r of - 65 •- water i n the p l a n t . " I t i s suggested that i n the absence of potassium, the e l e c t r i c a l p r o p e r t i e s of the c e l l are lessened, and that the osmotic forces are the r e f o r e reduced, r e s u l t i n g i n a decreased a b i l i t y on the part of potassium d e f i c i e n t c e l l s to draw water from the stream. A c e r t a i n osmotic fo r c e would s t i l l e x i s t due to c o l l o i d a l matter, sugars and such l i k e , and i m b i b i t i o n would play i t s p a r t . An abundance of water would, t h e r e f o r e , be r e q u i r e d by potassium starved pla n t s James (21) corroborates t h i s theory i n p o i n t i n g out that the potassium present i s i n many cases capable of ex e r t i n g a consid erable p o r t i o n of the osmotic pressure recorded by Dixon, and that a h i g h e r concentration of potassium w i l l r e s u l t i n a greater a b i l i t y on the part of the c e l l to maintain i t s t u r - g i d i t y . E s p e c i a l l y w i l l t h i s be the case, i f as recorded by Eotyschew and E l i a s b e r g [26), a l l of the potassium i n the c e l l e x i s t s i n an i o n i c form. In many pl a n t s potassium s a l t s seem to be the predominant so l u b l e s a l t s , and t h i s may be the case w i t h tomatoes. Warne (50) has shown, wi t h potatoes, there i s a withdrawal of potassium from the a s s i m i l a t i n g areas r e  mote from the main v e i n s , and suggests there may be as a con sequence l e s s photosynthesis i n those areas; the carbohydrate co n c e n t r a t i o n w i l l be reduced i n those spots, w i l t i n g w i l l f o l l o w due to a reduced osmotic pressure and d i s c o l o r a t i o n and scorch w i l l r e s u l t , i t i s suggested that the bronzing of the leaves of potassium d e f i c i e n t tomato p l a n t s may be also i n part due to t h i s . The linka g e of the bronzing with an i r o n accumulation also forms an i n t e r e s t i n g s p e c u l a t i o n . MacMurtrey - 6 6 - (32) has a l s o found a l o c a l i z a t i o n of potassium s t a r v a t i o n e f f e c t . ' A study of Table V I I , page 36, shows the Series I I I p l a n t s to have produced l e s s growth and f r u i t than d i d those of S e r i e s I which were fe d the normal amount of s o l u t i o n . Jan- sen and Bartholomew (23) and James (21) have both shovm that l e s s carbohydrates are manufactured and James (21) has shown fewer p r o t e i n s are synthesized. An absence of growth, there f o r e , would be expected. Chemical a n a l y s i s shows a high per centage of s t a r c h to be s t o r e d i n the r o o t s , and t h a t p r o t e i n s are h i g h i n a l l p a r t s . E v i d e n t l y a h i g h percentage of t h e carbohydrates which were manufactured were not u t i l i z e d and there was i n t u r n a v a i l a b l e nitrogenous matter. The leaves, too, were high i n reducing sugars which have n e i t h e r been a s s i m i l a t e d nor t r a n s l o c a t e d . This i s i n accordance with Jan- sen and Bartholomew's f i n d i n g s that i n the absence of potas sium there w i l l not be an a s s i m i l a t i o n of carbohydrates and of s o l u b l e n i t r a t e s ; and r e c a l l s h i s statement that high pro t e i n s and h i g h sugars found i n the absence of good growth and f r u i t i n g i n potassium d e f i c i e n t p l a n t s are l i k e l y due to a l a c k of p o l y m e r i z a t i o n of these to higher compounds. A t t e n t i o n i s drawn to the brown leathery roots of the potassium d e f i c i e n t p l a n t s , w i t h a suggestion of a storage 0rgan. This i s i n t e r e s t i n g i n view of the large amount of s t a r c h present i n them. I t i s r e c a l l e d t h a t there are few root h a i r s , but these roots are, nevertheless, very e f f i c i e n t as absorbing agents, f o r l a r g e q u a n t i t i e s of water were t r a n s p i r e d . - 6? - . While f r u i t production was low i n q u a n t i t y , there were many larg e f r u i t s . These though were apt to be i r r e g u  l a r i n shape. Both sugar and p r o t e i n content was high i n these f r u i t s . Probably the a v a i l a b l e potassium was u t i l i z e d by the e a r l i e r formed f r u i t s of a c l u s t e r and from them there would be no r e - t r a n s l o c a t i o n . Then the abundance of u n u t i l i z  ed nitrogenous matter and sugars would be r e a d i l y a v a i l a b l e to s w e l l the s i z e of the f r u i t s . I r r e g u l a r i t y i n shape might be due to an improper balance. The l a t e r f r u i t s formed i n the c l u s t e r would n e c e s s a r i l y be s m a l l e r . The average weight of the f r u i t s , n e v e r t h e l e s s , was h i g h . The dry weights of the tops and f r u i t s were l e s s and the S p e c i f i c G r a v i t y of the j u i c e s was l e s s i n these pot assium d e f i c i e n c y p l a n t s than those r e c e i v i n g P u l l N u t r i e n t , thus i n d i c a t i n g t h a t there i s l e s s mineral and v o l a t i l e matter. Sugars and p r o t e i n s being higher, the percentage of other dry matter would be s t i l l l e s s . I t i s r e c a l l e d that James and Penston (22) consider potassium to make up a large part of the dry weight of p l a n t s , and t h e r e f o r e i n the absence of potassium from the plant food there i s a n o t i c e a b l e d i f  ference i n dry weight. The p r e l i m i n a r y experiments i n Section I showed l e s s d i f f u s i o n out of potassium i n the f i r s t two experiments and then the same i r r e g u l a r i t y of absorption i n the t h i r d experi ment which was obtained i n the main experiment. The success i n growing p l a n t s from seeds whose parents had no potassium f o r two generations would lend strong - 68 - support to the theory of the t r a n s l o c a t i o n of potassium i n potassium d e f i c i e n t plants, I t must undoubtedly be stored i n the seed and then be r e u t i l i z e d repeatedly i n the new p l a n t . These seeds, as Table X, page 53 > shows are smaller i n s i z e than normal seeds and become more so each year of continued potassium s t a r v a t i o n . The seeds from the potassium d e f i c i e n c y p l a n t s of 1933 showed l i t t l e d i f f e r e n c e from the F u l l N u t r i e n t seeds; those from the complete potassium omission grown i n 1932 were considerably s m a l l e r ; whi1e the 1933 complete potas sium omission seeds whose forebears had received no potassium f o r two generations were f u r t h e r d e c l i n e d i n s i z e and showed greater v a r i a t i o n . They d i d , however, carry enough potassium to ensure a reasonable development to the new seedling i n the germination t e s t . A l l pla n t s from, seeds of even the second generation of omission when fed a complete n u t r i e n t are making good growth and f r u i t i n g ' though, as was pointed out under " R e s u l t s " ; t h e y are l e s s vigorous. - 69 - Phosphate• That a small t o t a l amount of phosphates is. reauired f o r p l a n t growth (Andre) would he i n accordance with the f i n d - i n t s i n the Absorption Experiment where l e s s phosphate was ab sorbed i n m i l l i - e q u i v a l e n t s than any other i o n . The gradual absorption d u r i n g the growing pe r i o d and then the sudden cessa t i o n about blossoming time, f o l l o w e d 1 by an absorption of small q u a n t i t i e s , and the l a t e r increase during f r u i t i n g i s i n t e n s e l y i n t e r e s t i n g i n view of the c o r r e l a t i o n i n the N u t r i t i o n E x p e r i  ment. I t i s notable that no e f f e c t of phosphate s t a r v a t i o n appeared i n S e c t i o n I I Phosphate d e f i c i e n c y p l a n t s u n t i l n e arly two months a f t e r the omission from the feedings. The plants of the two s e c t i o n s were about two weeks apart i n growth processes. At the time corresponding to the development of the N u t r i t i o n p l a n t at which phosphates were w i t h h e l d , i t ceased to be absor bed by the p l a n t of the Absorption Experiment. Apparently the pl a n t had a l l t h a t i t required f o r i t s present needs or was r e u t i l i z i n g what i t d i d have. There was none being absorbed and the plant of the N u t r i t i o n Experiment showed no e f f e c t of i t s omission.. L a t e r , i n blossoming, small amounts were absorbed and, f i n a l l y , as f r u i t i n g became more p r o l i f i c , l a r g e r amounts were used. I t was only at t h i s p o i n t that the N u t r i t i o n E x p e r i  ment p l a n t developed the p u r p l i n g which i s accepted as an i n d i c a  t i o n of a phosphate l a c k . Brazeale (3), i t has already been men ti o n e d , found a l s o , t h a t phosphate absorption by wheat p l a n t s increased up to a c e r t a i n concentration a f t e r which there was a sharp d e c l i n e - 7 0 - According to K a c G i l l i v a r y ( 3 1 ) , phosphate i s neces sary f o r c e l l d i v i s i o n and therefore for a l l stages of growth; but i f there i s a shortage there w i l l be t r a n s l o c a t i o n and r e u t i l i z a t i o n . This was probably the case -with our p l a n t s * He a l s o points out that l a r g e q u a n t i t i e s of phosphorus are used i n the f r u i t : " h a l f the phosphorus of the plant i s i n the f r u i t " . I t i s at t h i s stage t h a t the Section I plan t s again absorbed l a r g e q u a n t i t i e s of phosphates, and that the Section I I p l a n t s showed i t s l a c k . Table V I I , page 3 6 , and P l a t e I I I , (appendix) show the growth of the phosphorus d e f i c i e n c y plants to be very s i m i l a r to that of the P u l l N u t r i e n t p l a n t s . A f u r t h e r study of the t a b l e , nevertheless, i n d i c a t e s a lower dry weight of tops and a d i s t i n c t l y lower f r u i t production, This would support the theory of M a c G i l l i v a r y again, that phosphates are necessary i n the synthesis of nucleo-proteins, and that phosphorus i s high i n the f r u i t s . The a c t i v i t y main t a i n e d would be due to the t r a n s l o c a t i o n and r e u t i l i z a t i o n * The M c G i l l i v a r y p l a n t s were t r e a t e d with an e n t i r e phosphate omission from the s e e d l i n g stage and where h i s plants showed a very decided e f f e c t i n decreased growth and f r u c t i f i c a t i o n ours showed a r e l a t i v e l y s l i g h t one. I t would seem that the plant can manage w i t h a r e l a t i v e l y small amount of phosphorus. The p u r p l i n g of the leaves would be due to a discon tinuance of c h l o r o p h y l l productions accompanied by other changes as yet unaccounted f o r : p o s s i b l y accumulations of end products due to a d i s o r g a n i z a t i o n of the c e l l , on account of a derangement i n f u n c t i o n i n g , decomposition of c h l o r o p h y l l , - 71 - • or some obscured reason. MacGillibrary suggests that an accu mulation of sugars would stop a f u r t h e r synthesis of sugars, and i t i s u n l i k e l y c h l o r o p h y l l would be formed i f i t were not going to be used. The roots of the M a c G i l l i v a r y p l a n t s were a dark brown. Curs a l s o were darker than those of the P u l l N u t r i e n t f e e d i n g . Whether t h i s i s an i n d i c a t i o n of high s t a r c h storage i s a l o g i c a l question, f o r a s i m i l a r but more pronounced e f f e c t was found i n the roots of the potassium d e f i c i e n t plants where starches were i n an even higher percentage. The phos phorus d e f i c i e n t p l a n t s d i d not l a c k root h a i r s to any great extent as d i d the potassium d e f i c i e n t s e r i e s . The high s t a r c h content of the roots would be due to the f a c t that n u c l e o - p r o t e i n s are not formed i n the absence of phosphorus and the carbohydrates are not u t i l i z e d . I t seemed e a r l y , that there would be f r u i t produc t i o n equal to the P u l l N u t r i t i o n p l a n t s , f o r there were an equal number of f l o r a l buds. M a c G i l l i v a r y too, found t h i s , but suggests that there i s l a t e r a f a l l i n g o f f . The e a r l i e r buds p o l l i n a t e d would draw on the phosphorus and continue to grow at the l o s s of the l a t e r ones. As was pointed out i n the case of the potassium d e f i c i e n c y p l a n t s , t h i s would account f o r the i r r e g u l a r i t y of s i z e . While we d i d not obtain the pronounced d i f f e r e n c e i n amount of pulp and d i f f e r e n c e i n seed s i z e , our r e s u l t s ran f a i r l y p a r a l l e l with h i s . The lower dry weight and the lower S p e c i f i c Gravity of the f r u i t i s accounted f o r by the f a c t that phosphate i s considered to make up such a l a r g e part of the tomato f r u i t . P r o t e i n and - 72 - sugar content of the f r u i t s were hi g h . Since nucleo-proteins and phospholipoids are not formed i n the absence of phosphorus and the carbohydrates are not u t i l i z e d , l a r g e r amounts of nitrogenous matter and sugars would be a v a i l a b l e . With an increased d e p l e t i o n of phosphorus, M a c G i l l i v a r y obtained a higher percentage of carbohydrates and t o t a l n i t r o g e n i n l e a f and stem a l s o . His records throughout are extreme i n the l i g h t of ours. E v i d e n t l y a small amount of phosphorus w i l l have very e f f e c t i v e r e s u l t s . Seeds grown i n the S u b s i d i a r y Seed Experiment (b) are shown by Table X, page 5 3 ? to be only s l i g h t l y smaller than the F u l l N u t r i e n t seeds and give a s i m i l a r germination t e s t . Again, i t i s pointed out that the p l a n t s of t h i s i n  v e s t i g a t i o n were not so g r e a t l y starved f o r phosphorus as were M a c G i l l i v a r y ' s . Seeds of the S u b s i d i a r y Experiment ( c ) , on the other hand, which were from parents which had no phos phate f e d , and are now being fed none/have produced a seedling, at present 12 inches h i g h , which d i f f e r s i n l e a f shape from the parent>, so that here one would indeed "expect a d i f f e r e n c e i n v a r i e t y " . This same e f f e c t of a phosphorus omission has been found by other workers i n the U n i v e r s i t y . A g e n e t i c a l study and chromosome count show no d i f f e r e n c e i n such cases. I t seems to be a c y t o l o g i c a l mutation. - 7 3 - Do uble Goneentrati on. Hoagland (18) has s a i d that y i e l d s seem to he deter mined by a f a v o r a b l e supply and concentration i n the e a r l y stages of growth, r a t h e r than by intense absorption i n the l a t e r stages. Tyson, (46) has s t a t e d that the l i f e processes are more i n f l u e n c e d by the r a t i o and concentration of the n u t r i e n t s than by a supply of any one element* Both are pro bably r e f e r r i n g to a well-balanced s o l u t i o n and to concentra t i o n i n that sense. The two statements are only a p p l i c a b l e to t h i s i n v e s t i g a t i o n i n a general way. Nevertheless, the chang ing of the concentration i n S e r i e s V, g i v i n g double the con c e n t r a t i o n , but only h a l f the feedings, undoubtedly gave i n t e r e s t i n g r e s u l t s . The p l a n t s were normal i n every way, f r u i t i n g w e l l . They d i d , however, seem to make s l i g h t l y more succulent growth than the Series I , P u l l N u t r i e n t p l a n t s | the f r u i t s were somewhat l a r g e r i n average s i z e , but since p r a c t i  c a l l y the same t o t a l weight of f r u i t was produced, they were n e c e s s a r i l y fewer i n numbers. T o t a l sugars were s l i g h t l y h i g her i n the l e a f of the P u l l N u t r i t i o n p l a n t , but i n the double c o n c e n t r a t i o n they were higher i n stem and r o o t . Since the t o t a l sugars were p r a c t i c a l l y the same f o r the two types of p l a n t s taken as an e n t i t y , i . e . , root plus stem plus l e a f , i t i s probable that the Double Concentration p l a n t s may have been a l i t t l e more advanced than the checks and the sugars were passing out of the l e a f to the stem and r o o t . T o t a l starches were i d e n t i c a l i n the two s e r i e s . P r o t e i n s were much higher i n the r o o t s and somewhat so i n the f r u i t s . These _ 74 - d i f f e r e n c e s i n p r o t e i n content are unaccounted f o r , but i t would seem the d i f f e r e n c e i n concentration may have produced a s l i g h t l y more succulent v e g e t a t i v e growth, perhaps wi t h more nitrogenous matter which has r e s u l t e d also i n l a r g e r f r u i t s . In c o n s i d e r i n g the question of concentration, a t t e n  t i o n i s drawn to the S u b s i d i a r y Experiment (b), where seeds from the Absorption Experiment p l a n t s were grown. In the Ger mination Experiment, the roots were c o n s i s t e n t l y l a r g e r and more vigorous. Those plan t s from the Absorption P l a n t seeds which are being fed a complete n u t r i e n t now, are making n o t i c e  ably b e t t e r growth than those from the P u l l N u t r i e n t seeds. I t i s p o s s i b l e the question of a balance of s a l t s may enter i n t h i s connection. In the Absorption Experiment, the plants were washed through w i t h d i s t i l l e d water f o r t n i g h t l y , thus keeping the r o o t s washed cl e a n . The f r e s h n u t r i e n t s absorbed would be unaffected by any deposit of unused m a t e r i a l s . I f the p l a n t was thus able the more r e a d i l y to keep the balance i t d e s i r e d , the advantage might be r e f l e c t e d i n the seed. These p l a n t s are being kept under observation, and they w i l l be f o l l o w e d to m a t u r i t y and compared w i t h the p l a n t s from Series I seeds. I t i s suggested that the more frequent waterings with d i s t i l l e d water which the Double Concentration p l a n t s would r e c e i v e over the other Series might perform the f u n c t i o n of keeping the r o o t s washed.off. I t i s noted that the roots of S e r i e s Y p l a n t s were not superior to those of Series I . In deed i t cannot be s a i d that the p l a n t s as a whole were s u p e r i o r . - 7 5 - But seemingly a change i n concentration does have an e f f e c t . I t i s pointed, out elsewhere that t h i s i n v e s t i g a t i o n seems to show that p l a n t s absorb only those n u t r i e n t s which they require and w i l l use. There may, however, be an a l t e r a t i o n i n these requirements i n response to a change of environment. - 76 - Oo^Elg-t g S t a r v a t i o n Plants« The change of c o l o r i n the leaves i s an i n t r i g u i n g p o i n t i n the Complete S t a r v a t i o n p l a n t s . N i t r a t e omission r e s u l t e d i n a y e l l o w i n g and f i n a l l y a whitening with a purple c a s t ; potassium omission produced a bronzing;and phosphate d e f i c i e n c y gave a p u r p l i n g . I t would seem that the symptoms of a l l three types of s t a r v a t i o n were present here i n some deg ree, though that of the n i t r a t e s t a r v a t i o n was most apparent. In the Complete S t a r v a t i o n plants a y e l l o w i n g f i r s t appeared, and l a t e r a bronzing and p u r p l i n g , u n t i l f i n a l l y they became completely c o l o r l e s s and f r a g i l e . That growth and f r u i t i n g should be l e s s would be expected i n that the p l a n t had only the m a t e r i a l s w i t h i n i t s e l f to draw upon. That f r u i t i n g should occur would be i n confor mity w i t h the law that reproduction w i l l tend to occur when an organism i s deprived of i t s accustomed growth environment. In t h i s case, however, the n u t r i e n t s were not cut o f f u n t i 1 the blossoms had formed. Since growth was so g r e a t l y decreased, the percentage of sugars to weight d i d not vary g r e a t l y from the P u l l N u t r i e n t p l a n t s % though the percentage of dry weight of stems and f r u i t s and the S p e c i f i c G r a v i t y of the f r u i t s was s l i g h t l y l e s s , i t i s probable the mineral content would not vary to any great extent e i t h e r . Starches were high i n the r o o t s . Before the leaves began to y e l l o w , photosynthesis would be c o n t i n u i n g and i f the carbohydrates were not being u t i l i z e d i n growth there would be an accumulation. Apparently these - 77 - were stored i n the r o o t s . The pr o t e i n s i n the plant as an e n t i t y , were equal to those of the check p l a n t s , hut "by f a r the l a r g e s t amount was i n the r o o t . A c i d i t y was s l i g h t l y higher i n the f r u i t s of the f u l l y starved p l a n t s . Organic a c i d content must have been greater? t h i s i s not accounted f o r . Ebte Regarding P o t s i - Two types of l a r g e pots were used i n the Nutrition Experiment, glazed and c l a y . The c l a y pots r e q u i r e d more watering, but no apparent d i f f e r e n c e i n growth was obtained. The pots were mixed throughout the s e r i e s . - 7 8 - MogmssmATt CMS % I t i s recommended t h a t i n growing tomatoes, s o i l s must he kept s u p p l i e d with m i n e r a l n u t r i e n t s a t a l l stages of growth. Some q u a l i f i c a t i o n s as to the supply may he made, how ever. "While n i t r a t e s are needed throughout growth, a l a r g e r amount may he s u p p l i e d i n the f i e l d where l i g h t c o n d i t i o n s are at the maximum, than i n the greenhouse under poorer l i g h t c o n d i t i o n s . The v a l u e of c a l c i u m i s too w e l l known to r e q u i r e s t r e s s i n g h e r e . Magnesium and sulphates seem to he r e q u i r e d throughout the l i f e o f the p l a n t , but small amounts are a c t u a l  l y used. Potassium and phosphate f e r t i l i z e r s need not be a p p l i e d i n l a r g e q u a n t i t i e s to an annual crop s i n c e t h e r e i s a r e u t i l i z a t i o n of both these n u t r i e n t s . F i e l d t e s t s upon the p l a n t s themselves are an e x c e l  l e n t i n d i c a t i o n o f the a v a i l a b l e n u t r i e n t s i n the s o i l . I t i s suggested that an i n v e s t i g a t i o n i n regard to the m i n e r a l content of the p l a n t , and p a r t i c u l a r l y of the f r u i t , under omission treatments such as were c a r r i e d out i n t h i s i n v e s t i g a t i o n , w i t h a d d i t i o n a l treatments of a calcium, and a sulphate omission or d e f i c i e n c y , would be of considerable i n t e r e s t and v a l u e . - 7 9 - • SUMMARY. Ic I n v e s t i g a t i o n s were c a r r i e d out w i t h the tomato p l a n t , Lycopersicum esculentum M i l l . , as to; - (a) the period of development at which the n u t r i e n t ions Ca, Mg, S0 4, NO^, K, and PQ^ were absorbed! and (b) the e f f e c t the p a r t i a l with h o l d i n g of n i t r a t e , potassium, and phosphorus, a change i n con c e n t r a t i o n of the i o n s , and a complete s t a r v a t i o n might have on the growth, f r u c t i f i c a t i o n , and sugar and p r o t e i n content of the v e g e t a t i v e parts of the plant and the f r u i t . 2. The methods employed and the r e s u l t s obtained f o r e ach of the two p a r t s of the i n v e s t i g a t i o n are given separately as S e c t i o n I and S e c t i o n I I , but since i n t e r e s t i n g c o r r e l a  t i o n s developed between the two experiments, d i s c u s s i o n i s reserved u n t i l the f i n d i n g s of both are presented, and i s t h e r e f o r e g e n e r a l . 3. In S e c t i o n I , which i s nominated, Absorption Experiment, the p l a n t s were grown i n n u t r i e n t c u l t u r e s and a n a l y s i s of the c u l t u r e was made f o r t n i g h t l y to f i n d the absorption f o r the designated ions during that period. 4. In S e c t i o n I I , designated as N u t r i t i o n Experiment, the p l a n t s were grown i n n u t r i e n t sand c u l t u r e s and s i x s e r i e s of treatments were given as f o l l o w s : - I E u l l N u t r i e n t , I I N i t r a t e omission from blossoming time. I I I Potassium d e f i c i e n c y throughout, Iv Phosphorus d e f i c i e n c y from b l o s  soming time, V Double the concentration of Series I, but - 8 0 - f e d once a week i n s t e a d of tvri.ce, VI Complete S t a r v a t i o n from blossoming time. 5- Tables, p l a t e s and d e s c r i p t i o n s are given to show that the elements, calcium, magnesium, sulphur, n i t r o g e n , potassium and phosphorus are e s s e n t i a l to normal p l a n t growth, and that t h e i r omission causes a t y p i c a l s t a r v a t i o n e f f e c t . 6. Calcium, magnesium and sulphates are absorbed at a l l stages of growth and f r u i t i n g , but the e f f e c t of t h e i r omis s i o n was not s t u d i e d . 7® N i t r a t e s are necessary at a l l stages of growth, blossoming and f r u i t i n g ; more are u t i l i z e d during b r i g h t weather. 8. Potassium i s absorbed throughout the l i f e of the p l a n t . I t seems to have a :. r a d i o - a c t i v e or e l e c t r i c a l q u a l i t y and i s not r e q u i r e d i n as large q u a n t i t i e s i n s u n l i g h t as i n d u l l weather. In i t s absence there wi11 be a r e u t i l i z a t i o n of the potassium already present. The a b i l i t y of plants completely starved of potassium to continue i n t o the t h i r d generation, without being s u p p l i e d any, gives strong sup port to the theory of r e u t i l i z a t i o n . 9„ Phosphorus i s needed f o r both growth and f r u i t i n g . But, as evidenced by a c e s s a t i o n of absorption at blossoming time, c o r r e l a t e d w i t h an i n d i f f e r e n c e to i t s omission at that time, i t would seem not to be e s s e n t i a l , at l e a s t i n added q u a n t i t i e s at th a t p o i n t . Since i t i s more abundant i n the growing parts of the p l a n t , i t i s p o s s i b l e there i s - 81 - a r e u t i l i z a t i o n at that t i m e . S t o r i n g would he a p o s s i b i  l i t y , but the evidence of t h i s i n v e s t i g a t i o n does not point to a storage of mineral n u t r i e n t s by p l a n t s . 10. A change i n concentration gave s l i g h t l y more succulent growth and a l a r g e r average s i z e of f r u i t . There were fewer f r u i t s , but the t o t a l weight was almost equal to that produced by the P u l l N u t r i e n t check p l a n t s . The t o t a l amount of ions f e d i n the two s e r i e s was i d e n t i c a l . 11. A complete s t a r v a t i o n of a l l n u t r i e n t s and of n i t r a t e s only produced s i m i l a r r e s u l t s . 12. The amount of f r u i t produced i n a l l omission s e r i e s was c o n s i d e r a b l y l e s s than i n the check p l a n t s : i n the Potas- • sium d e f i c i e n c y i t was 60% of the check, i n the Phosphate omission from blossoming i t was 65%> i n the N i t r a t e omis si o n from blossoming i t was 20/», and i n the Complete Star v a t i o n 18/b. 13. Analyses were made of the l e a f , stem, root and f r u i t of a l l s e r i e s i n S e c t i o n I I , f o r carbohydrates and proteins and t a b l e s are given presenting the f i n d i n g s . 14. A h i g h storage of s t a r c h was found i n the roots of the p l a n t s of a l l omission treatments, and of p r o t e i n s i n the roots of the Potassium d e f i c i e n c y , Phosphate omission and Complete S t a r v a t i o n p l a n t s . 15. The v a r i a t i o n s i n sugar, s t a r c h and p r o t e i n content are discussed i n d e t a i l . - 82 - l 6 e Sugars were found not to vary to any great extent i n the f r u i t s ; p r o t e i n s v a r i e d r a t h e r more. 17. A c i d i t y was found not to vary to any extent except i n the N i t r a t e omission s e r i e s . 18. Seedlings grown from seeds of the Absorption Experiment pl a n t s had a more extensive root system than had any of the others, and t h i s i s r e f l e c t e d i n a somewhat b e t t e r growth they are making over the E u l l N u t r i e n t s e e d l i n g p l a n t s . I t i s suggested that a b e t t e r balance of s a l t s was maintained i n washing the roots of the Absorption p l a n t s f o r t n i g h t l y . 19. The r e s u l t s of the i n v e s t i g a t i o n s t r o n g l y suggest that p l a n t s absorb only those n u t r i e n t s which they r e q u i r e and w i l l u t i l i z e . 20. I t i s recommended that a mineral a n a l y s i s of the f r u i t s of s i m i l a r l y grown p l a n t s , w i t h a Calcium d e f i c i e n t and a Sulphate d e f i c i e n t s e r i e s added, would be of value. 21. I t i s f u r t h e r recommended that a l l mineral n u t r i e n t s must be kept c o n t i n u a l l y s u p p l i e d to the growing p l a n t , but that other than calcium and n i t r a t e they need not be s u p p l i e d i n l a r g e amounts. - 83 - ACKNOY/L1SDGMENT S S i n c e r e thanks are r e n d e r e d to Bean F. M. Clement for- a v e r y k i n d c o n s i d e r a t i o n and i n t e r e s t * To Dr. A. 3 7 . B a r s s f o r h i s k i n d l y and sy m p a t h e t i c a t t i t u d e towards the work, the w r i t e r i s i n d e b t e d . G r a t e f u l acknowledgment and a p p r e c i a t i o n i s ex p r e s s e d t o Dr . G. H. H a r r i s , who suggested t h e i n v e s t i g a t i o n and whose a d v i c e and a s s i s t a n c e were a t a l l times most g r a  c i o u s l y and g e n e r o u s l y g i v e n . Thanks a r e a l s o extended to Mr. Frank G a r n i s h f o r many c o u r t e s i e s a t the greenhouse. 84 - LITERATURE CITED (1) American A s s o c i a t i o n of A g r i c u l t u r a l Chemists (A.O.A.C.) Washington, D.C. (2) " Andre G. • I918 R e p a r t i t i o n des elements minereaux et 1 'azote chez l e Tegetat etoile.Comp. Rend. Acad. S c i . (Paus) 167.1004 - 1006. (3) Brazeale, J . I . 1928. E f f e c t s of one element of plant food upon the absorption by pl a n t s of another element. Univ. A r i z . Agr. Exp. L t a . Tech, B u i . 1 9 (4) Cameron and P a i l y e r . Jour. Arner. Chem. Soc. 1068. (5) Colby, Harold L. 1933* Seasonal absorption of n u t r i e n t s a l t s by the French prune grown i n s o l u t i o n c u l t u r e s . Plant P h y s i o l . 8 ( l ) s I -34 ' (6) . Davidson, J e h i e l . 1933• The p o s s i b l e e f f e c t of H ion concentration on the absorption of potassium and phosphorus under f i e l d c o n d i t i o n s . Jour. A g r i c . Res. 46 (5) : 449-506. (7) Davis, M.B. I93O. Some e f f e c t s of the d e f i c i e n c i e s of nit r o g e n , potassium, calcium and magnesium w i t h s p e c i a l reference to the behavior of cer t a i n v a r i e t i e s of apple t r e e s . Jour. Pomo1. and Hort. S c i . - V o l . 8 No . 4 , (8) Day, Dorothy. 1929. Some e f f e c t s of calcium d e f i c i e n c y on Pisum sativum. P l a n t P h y s i o l . 4 ( 4 ) : 493-506. (9) Davy, S i r Humphrey. l 8 l 4 . Elements of A g r i c u l t u r a l Chemistry. - 85 - (10) Ernmert, E.K. ' 1931 8 The e f f e c t of s o i l r e a c t i o n on the growth of tomatoes, and l e t t u c e , and on the n i t r o g e n , phosphorus and manganese content of the s o i l and p l a n t . Univ. Kent., Agric.Exp. Sta. Res. B u i . 314'. 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The i n f l u e n c e of s o i l c o n d i t i o n s , f e r t i l i z e r t r e a t m e n t s , and l i g h t i n t e n s i t y on growth, c h e m i c a l c o m p o s i t i o n and enzymic a c t i v i t y - 89 - of sugar beets. Mich.. State C o l l . Agr. Exp. Sta. Tech. B u i . 108. (47) Wallace, T. 1925» Experiments on the manuring of f r u i t t r e e s . I I . Jour. Pomol. and Hort. S c i . V. Ho. 1. (48) Wallace, T. and Mann, C.E.T. 1926. I n v e s t i g a t i o n s on the c h l o r o s i s of f r u i t t r e e s . Jour. Pomol. and Hort. S c i . V. Ho .2 (49) Wallace, T. 1928. Experiments on the manuring of f r u i t t r e e s . Jour. Pomol. and Hort. S c i . V I I . Ho.l (5*0) Warne, L.G.G. 1934. The d i s t r i b u t i o n of potassium i n normal and scorched f o l i a g e . Ann. Bot. XLVIII:5 7 - 67. ( 5 1 ) Watts, W.M* I93I. Some f a c t o r s which i n f l u e n c e growth and f r u i t i n g of the tomato. Univ. Ark. Agr. Exp. Sta. B u i . 267. ( 5 2 ) Weevers, T. I9II. Untersucheengenuber die L o c a l i s a t i o n und Eunktion des Kaliurn i n der P f l a n z e . Rec. Trav. Bot. Heerland. V I I I . ( 5 3 ) WooIf, E. I87O. Ashen A n a l y s i s . - b -- f) „ PLATE IV. Showing V a r i a t i o n i n leaves. - f "PLATE V. Showing v a r i a t i o n i n r o o t s . PLATE V I . The Potassium. O m i s s i o n P l a n t s o f S u b s i d i a r y Experiment ( a ) , showing on the l e f t t he p l a n t which has had no p o t a s s i u m f o r two g e n e r a t i o n s . The one on the r i g h t was from the same seeds but was f e d p o t a s s i u m a t b l o s s o m i n g . 

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