UNIVERSITY OF CALIFORNIA PUBLICATIONS IN AGRICULTURAL SCIENCES Vol. 3, No. 8, pp. 135-242, plates 13-24 July 12, 1918 THE CHEMICAL COMPOSITION OF THE PLANT AS FURTHER PROOF OF THE CLOSE RELA- TION BETWEEN ANTAGONISM AND CELL PERMEABILITY BY DEAN DAVID WAYNICK CONTENTS PAGE Introduction , 135 Object of the investigation 137 Review of previous investigations 137 Methods 140 Experimental data 144 External appearances of the plants 154 General review of experimental results 155 Results with salts of the heavy metals 156 Possible effects of variations in the concentrations of the solutions on the plants 160 Consideration of a possible Calcium-Magnesium ratio 160 Permeability and antagonism 162 Summary 164 Introduction A solution of a single salt at certain concentrations is toxic to plants grown in it. The addition of a second salt usually permits of growth superior to that in a solution of a single salt alone even though the added salt is toxic when used by itself. A third salt added may permit of a still further increase over the growth in the two salt solu- tion. Other salts added will increase or decrease growth, depending upon the salt used. Qualitative relationships only have been consid- ered. When we adjust the quantitative relationships of the various salts present, having at the same time due regard for their qualitative 136 University of California Publications in Agricultural Sciences [Vol.3 nature, we get as a result a solution in which the plant grows and functions normally. Such a solution has been termed by Loeb, "phy- siologically balanced. ' ' It is evident that if growth is better in a two salt solution the toxic effects of the solution due to a single salt must be lessened by the presence of the second salt. We may refer to either as the second salt since either may be toxic alone. On the addition of a third salt the increase in growth over that obtained in the two salt solution points to a still further lessening of the toxic properties of the various salts present taken singly. This action of one or more salts in limiting or preventing entirely the toxic effects of one or more other salts, is termed antagonism. Sea water may be taken as an example of a physiologically balanced solution or a solution in which the mutual antagonism between the constituents of the solution is such as to allow of normal growth of numerous organisms. The fact of the existence of antagonism has been proven by a number of investigators working in plant and animal physiology, but the mechanism of antagonistic action is by no means clear. Since salts are very largely ionized in the nutrient solutions usually em- ployed, it is probable that antagonism has to do with ions. Further, antagonism will probably take place between the ions present in, or between, the ionic constituents of the solution, and the living mem- branes in contact with the solution. Loeb 1 first advanced the theory that one ion may prevent the entrance of another ion into living cells and that in this property lies the reason for antagonistic action. On the basis of this hypothesis, penetration precedes the manifestations of toxic effects and where penetration does not occur, due to antagonistic action, there are no toxic effects evident. Used in this way, the term penetration means simply the entrance of ions in greater number than would normally occur were the plant cells in their natural environ- ment. Experimental evidence as to the correctness of this hypothesis has been furnished by Loeb 2 in a very interesting series of experi- ments. Ost^rhout 3 has applied the electrical conductivity method to the measurement of the penetration of ions into plant tissue, while recently Brooks has confirmed Osterh out's results (1) by deter- mining: the diffusion of ions through tissue, 4 (2) by exosmosis, 5 and (3) by the change in the curvature of tissue. i Amor. Jour. Physiol., vol. 6 (1902), p. 411. -Science, n.s., vol. 36, no. 932, p. 637. a Ibid., vol. 35, no. 890, p. 112. ' Proc. Nat. Acad., Sci., vol. 2 (1916), p. 569. ■Anier. .lour. Hot., vol. 3 (1916), p. 483. 1918] Waynick: Antagonism and Cell Permeability 137 It is evident that these methods are limited in their application and give no idea of the quantitative relationships existing between the ions actually entering the cells. They do show, however, that the permeability of the plant tissue may be greatly altered by salt action and that solutions which permit of normal growth, also preserve normal permeability as regards the ions present in the solution. Object of the Investigation In a preliminary paper 7 the results obtained from chemical analy- ses of plants grown in toxic and antagonistic solutions have been reported. These results were of interest and the general method em- ployed seemed to be worthy of a more extended application in the determination of ions absorbed by plants from solutions, of known composition and concentration. From a consideration of the data in the paper referred to above, it was felt that the results obtained in a more extensive investigation would be of importance: (1) from the standpoint of the effect of various salts upon the permeability of the cell tissue of growing plants; (2) from that of the effects of vari- ous salts upon the nutrition of plants as evidenced by growth; (3) from that of a possible correlation of growth with the absorption of ions; and (4) from the standpoint of the quantitative relationships existing between certain ions of the solution and the same ionic rela- tionships in the plant. The various phases of the problem as outlined above will be con- sidered in the discussion of the experimental results following. Review of Previous Investigations It is not intended that the following review of the previous work done in this field of plant physiology be exhaustive. Robertson 8 has reviewed the literature dealing with antagonistic salt action very completely up to a recent date. Brenchley 9 and Lipman and Gericke 10 have referred to all the important work done with regard to the effects of the salts of the heavy metals upon plants. The present review therefore touches only the work bearing directly • upon the a Ibid., p. 562. ' Contribution to the causes of antagonism between ions. (Univ. Calif., Master's thesis, 1915.) sErgeb. Physiol. Jahrb., vol. 10 (1910), p. 216. 9 Inorganic plant poisons and stimulants. New York, Putnam, 1915 (Cam- bridge agricultural monographs). i"Univ. Calif. Publ. Agr. Sci., vol. 1 (1917), p. 495. 138 University of California Publications in Agricultural Sciences [Vol.3 present problem or work so recent as not to be included in the papers cited above. A large share of the contribution to the experimental evidence in regard to antagonism between salts as regards plants we owe to Oster- hout. In a series of papers he has shown that any salt may be toxic to plants when used alone in solution at certain concentrations and further that the addition of a second salt may, in proper concentra- tion, modify or eliminate entirely the toxic effect of the first salt. He has shown further that acids, alkalies, and various organic compounds may likewise be toxic to plants and that their toxic effects may be modified by the presence of a variety of compounds, depending upon the toxic substance employed. By measuring the resistance of cylin- ders of Laminaria in solutions of one salt and in solutions containing two or more salts, he has brought forward much evidence as to the penetration of ions into plant cells. While this method has yielded very valuable results both as to the rate of entrance of ions and also the total number of ions penetrating, it does not yield results which give us a knowledge of the relative amounts of the various ions which penetrate the tissue when the qualitative as well as the quantitative relationships of the nutrient solution are varied. Osterhout has shown, however, that penetration is more rapid, and the degree of permeability is greatly increased, in unbalanced solutions and further that as the permeability of the plant tissue more nearly approaches normal the growth of the plant is also more nearly normal. Szucs 11 has used Cucurbit a pepo as an indicator by immersing the young seedlings in various solutions for varying periods of time and counting those still able to show geotropic movement when placed in a horizontal position in a moist chamber. He found a marked antagon- ism between copper sulphate and aluminum chloride and concludes from his experiments that antagonism consists in the mutual hin- drance of similarly charged ions in entering the cell. He states further that the rate of absorption of equally charged ions is of great importance. His chemical methods are open to question, for in the experiments reported the test for copper used was that of boiling the roots and testing the resulting solution for copper with hydrogen sulphide. By analyzing the solution in which pea seedlings had grown, Pan- ic! li'- has determined ion absorption. The growing period was short. ujahrb. Wies. Bot. (Pringheim) , vol. 52, no. 1 (1912), p. 85. >- Ibid., p. 211. 1918] Waynick: Antagonism and Cell Permeability 139 He found a rapid absorption of zinc, manganese, iron, and aluminum, but the total amounts taken up were small. He gives other evidence of the selective absorption of various other ions from solutions, but these results are of not direct application here. It is of interest to note, however, that he found a direct relation between time and ion absorption. His most important conclusion, which bears directly upon the problem in hand, is that strong narcosis was associated with the penetration of ions in large numbers. Schreiner and Skinner, 13 using a similar method, have determined the amounts of phosphoric acid, nitrates, and potassium remaining in a solution in which plants had been grown. Various ratios of these three ions were employed, the total concentration being 80 parts per million. They found widely varying amounts of these three ions removed from the solution, and further there seemed to be a possible difference of 20 to 30 per cent in the removal of any one without an apparent effect upon the growth of the plants. Under the condi- tions reported by them increased growth was correlated with increased absorption. By means of conductivity measurements of solutions in which pea seedlings were growing, True and Bartlett 14, 15 ' 16 have determined the rate of absorption and of excretion of electrolytes. Their work was done with one, two and three salt solutions. In general they found a greater absorption when a mixture of salts was present than when single salts were used. Further, the absorption relationships of salts with a common kation seem to be similar. For example, from solutions of low concentrations, potassium chloride, potassium sul- phate, and potassium nitrate are not removed, but on the other hand there is an excretion of electrolytes by the plant. In direct contrast, calcium nitrate and calcium sulphate are removed from their solu- tions in every concentration employed and no excretion of electro- lytes from the plants could be detected. It seems probable that the low concentration employed by them acted as a limiting factor in some cases. In a recent paper Breazeale 17 has shown that the presence of sodium carbonate, and sodium sulphate, when used in concentrations of 1000 parts per million in nutrient solutions, decreased the absorp- isBot. Gaz., vol. 50 (1910), p. 1. HAmer. Jour. Bot, vol. 2 (1915), p. 255. is Ibid., p. 311. ^ Ibid., vol. 3 (1915), p. 47. 17 Jour. Agr. Kesearch, vol. 7 (1916), p. 407. 140 University of California Publications in Agricultural Sciences [Vol.3 tion of potassium and phosphoric acid as much as 70 per cent below that of the control cultures. The work of Gile 18 is of interest in this connection. From ash analyses obtained in investigating the cause of chlorosis in pineapples, he found a direct relationship between the absorption of lime and that of iron; that is, when the absorption of lime was high but little iron was taken up. In soil studies Gile and Ageton 19 found no direct relation between the lime content of plants and varying amounts of lime and magnesia in the soil. A few investigations have been made on the absorption of specific elements from solution, but these need only be mentioned in the present connection. Maquenne 20 found that mercuric chloride causes marked increase in permeability of the protoplasm, although it is not necessarily absorbed itself in any considerable quantities. Marsh 21 correlates the amount of barium chloride present in the soil with that found in the plant. Colin and De Rufz 22 always found absorbed barium localized in the roots. A large number of analyses of plants grown under various condi- tions have been reported, but the environmental factors have varied so greatly as to render the results obtained of little value in the present study. From this review it is evident that no quantitative study of the elements actually absorbed 'from the nutrient solutions, balanced and unbalanced, has been made with the idea in mind of a correlation between the absorption of the various ions with their antagonistic or toxic effects in solution cultures. Methods Barley was used as the plant indicator. The seeds were obtained from the University Farm at Davis and were of a pure strain of the Beldi variety. The method of sprouting the seeds, while simple, has not been noted elsewhere and has given such excellent results, both to the writer and to others, that it seems worthy of mention here in detail. A piece of oilcloth about 12x18 inches was covered with sev- eral thicknesses of paper toweling and the whole thoroughly wetted. is I'orto Rico Exp. Sta. Bull., 11 (1911). ifl Ibid., Bull. 10 (11)14). 20C.-R. Acad. Sci. (Paris), vol. 123 (1896), p. 898. 2i Bot. Gaz., vol. 54 (1912), p. 2~>0. 22C. B. Acad. Sci. (Paris), vol. 150 (1910), p. 1074. 1918] Waynick: Antagonism and Cell Permeability 141 Selected seeds were distributed over the toweling so that about two hundred were placed on an area of the size indicated above. Another layer, made up of several sheets of toweling-, was then laid on the seeds and the whole thoroughly soaked with water. The water was allowed to evaporate gradually until the paper was but slightly moist to the touch and the water relation then maintained constant until the seedlings were transferred to the solutions. If the paper is kept too moist the growth of molds is often very abundant, but with a low moisture content no trouble was experienced from this source. By the time the roots were a quarter of an inch long, the upper layer of paper was supported two or three inches above the seedlings. This procedure permits of a straight growth of the shoots, which is of con- siderable importance in placing the seedlings in the corks. The seed- lings were transferred when the shoots were about an inch and a half in length. The paper in which the roots are grown, tears apart readily Avithout injuring them in any way, the oilcloth not permitting their downward penetration. There is no contact with metal containers at any time, the apparatus required is practically nothing, the time period is short — about six days under greenhouse conditions — and strong seedlings are obtained which can be transferred to any contain- ers without injury. The containers used were quart jars of the Mason type, each holding approximately 950 c.c. of solution. The inside of each jar, as well as that of the bottles for the stock solutions, was coated with a layer of paraffin so that the solutions were never in contact with the glass. The outside of the jar was covered with black paper to exclude light, the black surface facing the glass. Flat corks, having a diam- eter of three and a half inches, were used to support the seedlings. Each cork had seven holes, one in the center through which distilled water was added to maintain the volume of the solution as nearly constant as possible, and six equally spaced, one and a quarter inches from the center, for holding the seedlings. After the holes were made the corks were soaked in boiling paraffin. To introduce the seedlings the corks were turned upside down, supported by the rim of the jar, and the shoots stuck through the holes prepared for them and held in place by a small piece of cotton. On turning the corks over the seedlings were in their proper position without being in the least injured, for there was no necessity for touching the roots at any stage since the plant was always picked up by the seed coat. The method suggested by Tottingham 23 was tried, 142 University of California Publications in Agricultural Sciences [Vol.3 but the one outlined above proved very satisfactory and much simpler. The basic nutrient solution used throughout was Shive's three salt nutrient 24 containing the following salts in the given partial molecular concentrations : K H 2 P0 4 0180 M. Ca (N0 3 ) 2 0052 M. MgS0 4 0150 M. The stock solution was made up to twice the strength indicated above and diluted as necessary by the addition of added salt solution, or distilled water, or both. In the case of the chlorides used, viz., calcium, magnesium and potassium, normal or twice normal solutions were prepared and standardized by titrating against a standard silver nitrate solution. Normal solutions of magnesium and potassium sulphate were stand- ardized by weighing the barium sulphate precipitate. Solutions of copper, zinc, iron, and mercury salts were prepared in concentrations of 1000 parts per million by weighing out the carefully dried salts. The final volume of solution required for the duplicate jars was approximately two thousand cubic centimenters. Starting with a thousand of the nutrient solution, various volumes of the standard solutions were added so that when the total volume was made up to two liters with distilled water, the concentrations of the various salts would be those reported in the accompanying tables. The growing period was six weeks. The duplicate cultures were grown in specially constructed mouse-proof cages each holding ninety jars. The tops of the cages were open and the sides made of coarse wire screening. The different parts of the cages were equally well lighted, as shown by the nearly equal growth of the controls in dif- ferent parts of the cages. When necessary the plants were supported by cords strung across from side to side of the cages. The solutions were not changed during the growing period, but the volumes were kept as nearly constant as possible by adding dis- tilled water. There are objections to this method, as there are objec- tions to the method of using water cultures at all. The growth was found to be very satisfactory and compares favorably with the growth 28 Physiol. Researches, no. 4 (1915), p. 174. 24 Arner. Jour. Bot., vol. 2 (1915), p. 157. 1918] Waynick: Antagonism and Cell Permeability 143 obtained by other investigators in comparable periods of time. A further discussion of this point will be taken np below. At the expiration of the six weeks growing period the plants were removed from the corks, the roots rinsed thoroughly with dis- tilled water, placed between layers of paper toweling, dried in the oven at 100°-105?C, roots and tops separated, weighed, and placed in envelopes ready for analysis. For analysis the roots from dupli- cate cultures were combined unless the dry weight was sufficient to allow of separate analysis. Total ash was determined after direct ignition of the dry material in a muffle at a low red heat until no trace of carbon remained. The ash was then taken up in dilute hydrochloric acid and evaporated to dryness to remove possible contamination with silica. Iron was N precipitated as the hydroxide with ammonia and titrated with — — potassium permanganate after reduction with zinc and sulphuric acid. This determination was made because of the relation Gile has shown to exist between calcium and iron absorption by plants. Calcium N was precipitated as oxalate and titrated with — — potassium perman- ganate. The double precipitation of the oxalate assured freedom from magnesium contamination. Magnesium was precipitated by ammonium phosphate and weighed as the pyrophosphate. Potassium, where determined, was precipitated and weighed as the chloroplati- nate. Copper was determined colorometrically by using the ferro- cyanide method. The amount of material available precluded the possibility of a more complete analysis than was made if any degree of accuracy was desired. For example, in Series vn, the weight of the ash varied from 12 to 233 milligrams in the case of the roots and from 32 to 183 milligrams in the case of the tops. While these varia- tions are not extreme, they are fairly representative. The values of these elements actually determined cannot be taken as absolute in every case because of the limited amounts of material available, but the significant differences are so great as to make a small variation in this regard of minor importance. The strength of all solutions is uniformly expressed in terms of molecular concentrations since this mode of expression has been quite generally used in experimental work reported by different investigators. Under experimental results twenty-six series are reported. A series, as used in the present work, may be defined as a number of 144 University of California Publications in Agricultural Sciences [Vol.3 duplicate cultures containing one salt in varying concentrations in each, or one salt constant and varying concentrations of a second salt. In some instances both salts varied but only in concentration, the same ratios being maintained. These are few. The number of con- centrations reported vary from three to fourteen in a series, depend- ing upon the salt used. Before two salts were taken together, the effects of each separately upon the plants were determined. Usually this meant only the establishment of the toxic limits of the salts em- ployed when used in the nutrient solution. Several series of this kind are not reported here, as no analytical work was done upon them. Calcium and magnesium salts were used to a large extent because of the fact that their kations can be determined with less experi- mental error than most other nutrient salts where the small amounts of material dealt with here are considered; also it was of interest to determine whether or not there is a lime-magnesia ratio for plants grown under carefully controlled conditions. Copper, zinc, iron, and mercury salts were used because of the fact that their toxic and antag- onistic effects have not been previously determined as regards absorp- tion. Potassium chloride was the only monovalent salt used. A longer growing period than has usually been employed was con- sidered important. McGowan, 25 in conducting experiments in pure solutions of sodium, potassium and calcium chlorides, found growth better in the first two at the end of six days, but far superior in a solution of calcium chloride in twenty-five days. In a qualitative way the same relationships were observed in the present investigation. It seems reasonable to assume that the results obtained in six weeks with plants are more nearly representative of the true effect of various solutions than those obtained in two or three day periods or even in three week periods. But it is not assumed that the results herein reported are the same as those which might be obtained were the plants grown to maturity. It is hoped that more data may be pre- sented shortly on this point. In the following section, in which the experimental results are given, the time factor and the basic nutrient solutions are constants. Experimental Data All analyses are reported as percentages of the dry weights of the plants. To make the results obtained as clear as possible, graphs and photographs have been used throughout as well as the tables giving the actual percentage composition of the plants. 25 Hot. Gaz., vol. 45 (1908), p. 45. 1918] Wayhick: Antagonism and Cell Permeability 145 The relationships of calcium to magnesium salts are reported in the first seven tables. For a review of the more important literature bearing directly upon the relationships of the salts to these two ele- ments reference is made to McCool, 26 who has considered these in some detail, and to a recent critical survey of the lime-magnesia ratio hypothesis b}^ Lipman. 27 As is evident from table 1, calcium chloride does not become toxic until present in concentration of over .24 M. Up to and including this concentration the growth seems to be but little affected by the increasing concentrations of the salt added. The percentage of cal- cium in the plants shows no direct increase with increasing concen- tration of calcium chloride in the solution. The lowest percentage of calcium given occurs in a concentration of .20 M. calcium chloride. In table 2 there is a close parallelism between the growth of roots and tops. Two low points on the dry weight graph are evident, the first occurring at cultures 4 and 5 and the second from 7 to 11. At these low points we have a high percentage of magnesium in both roots and tops, but of calcium only in the second low point. Calcium is low where growth is good in cultures 2 and 3. But the most inter- esting feature is the decreased absorption of both elements at cul- ture 6, where there is a distinct increase in dry weight. Iron was not present in sufficient concentration to allow of titration until cul- ture 11 is reached. It may be stated here that the iron determined is limited to that in the seed as a maximum, for it was purposely ex- cluded from the solutions except where its toxic or antagonistic action was under observation. In many instances the titration of this residual iron is of interest. Table 3 is a record of one of the most interesting and significant series reported. The root growth was so limited in nearly every cul- ture that no attempt was made to segregate roots from tops for sep- arate determinations except where the total dry weight was so greatly increased as in cultures 6 and 11. In the first place we have double maxima of growth, the first in culture 6 and the second in 11. The total dry weight at culture 11 is twice that at 6, but the dry weight in culture 6 amounts to a 35 per cent increase over that in culture 7. A direct inverse relationship is shown between total growth and ab- sorption at these two high points ; the maximum growth in culture 11 is accompanied by the lowest absorption of calcium and magnesium. The percentage of magnesium is low in culture 6, but that of calcium 26 Cornell Univ. Agr. Exp. Sta. Mem. 2 (1913), p. 127, 27 Plant world, vol. 19 (1916), p. 83. 146 University of California Publications in Agricultural Sciences [Vol.3 is higher than in the cultures of slightly higher or lower concentra- tions. No explanation of the narrow ratio between these two ele- ments at this point can be offered. It is of interest to note the very great increase in the amounts of calcium and magnesium found in the plants grown in concentrations of .20 M. calcium chloride alone. While magnesium chloride is constant throughout the series, the amount of magnesium does not increase proportionately to that of calcium. A still higher concentration of magnesium chloride was used in the series reported in table 4. The percentage of magnesium found in the roots is very high and would indicate that it was not entirely removed from the roots by washing. In general the percentages of calcium and magnesium found are high, the calcium content increas- ing as the concentration of calcium chloride present in the culture, but not proportionately. Magnesium is lower at the greater dry weights for the tops, the decrease amounting to 50 per cent in the case of culture 6. Magnesium sulphate was used alone in the series reported in table 5. The decrease in growth is nearly proportional to the increase in concentration of the added salt. In this series w T e have a very marked decrease in the percentages of calcium and magnesium present in the roots without any evident effect upon the growth of the plants, especially that of the tops. Here again, however, we have increased absorption of calcium as the percentage of magnesium increases, even though the concentration of the former in the nutrient solution is constant. It is of interest to note that the percentages of both ele- ments in the tops throughout this series are low and vary but little, regardless of the increasing concentration of the nutrient solution. Very marked antagonism between calcium chloride and magnesium sulphate is shown in table 6. The dry weight of the plants grown in a solution of magnesium sulphate .18 M. concentration was .29 gram, but when .04 M. concentration of calcium chloride was added the aver- age dry weight was 1.20 grams and in a concentration of .18 M. magnesium sulphate and .24 M. calcium chloride the average dry weight was .98 gram. Between these two concentrations of calcium chloride the dry weights recorded are uniformly high. Correlated with the rapid decrease in growth, in concentrations of .24 M. of cal- cium chloride, is the marked increase in the percentage of both calcium and magnesium found in the plants. The graphs representing the amounts of these elements found crosses the growth graph coincident 191S] Waynick: Antagonism and Cell Permeability 147 with its sharp decline. The low percentage of magnesium is of interest since the concentration of the culture solution was uniformly high with respect to this ion. It is striking that there is a marked decrease in the growth of roots at the concentration which gave the best growth of tops, and further that the percentage of calcium in the tops and magnesium in the roots parallel this decrease in the growth of the roots. A comparison of the results obtained with magnesium sulphate as against those with magnesium chloride is reserved for later discussion. In table 7 we have an opportunity to compare indirectly anion effects, or possibly the effects of combinations of the same kation with different anions. From preliminary results it seemed advisable to use .15 M. magnesium sulphate in this series instead of .18 M. as used in the preceding series, so that the concentration of magnesium ion is not equivalent in the two series. A solution containing magnesium sulphate .15 M. plus calcium nitrate .08 M. proved highly toxic, while a solution containing calcium chloride of the same concentration as the nitrate in the above solution supported normal growth. It is possible that the difference is due to the toxic action of the nitrate ion on the plant directly. Tottingham has shown that the total ionization of a nutrient solution was decreased 10 per cent below the theoretical by the addition of calcium nitrate in low concentrations. It is pos- sible that the ionization of some other salt is repressed so that there is an actual lack of some ion necessary for growth. The percentage of calcium found was not high enough in any case to account for the toxic effects shown. Magnesium Avas found in extremely large amounts, 9.20 per cent in the case of culture 6, the largest percentage recorded in any culture studied. Unfortunately the series in which the toxic effects of calcium nitrate alone were studied was lost, so it cannot be reported here. Potassium chloride was the only monovalent salt studied, and the results are given in tables 8 and 9. The growth shown in the various concentrations of potassium chloride used was approximately the same as that found when magnesium sulphate was used alone. The increase in the percentage of ash, as far as the tops are concerned in table 8 is very striking. The percentage of calcium found in the tops and of magnesium found in the roots remains practically constant throughout. The amount of potassium absorbed increases as the con- centration of potassium chloride in the solution increases and in- versely as the growth of the plants. The toxic effects due to the 148 University of California Publications in Agricultural Sciences [Vol.3 addition of potassium chloride to the solution are much more evident in the tops than in the roots with respect to the increasing concen- trations of potassium chloride. Using a constant concentration of potassium chloride of .18 M., which is an increase of .02 M. over the highest concentration of that salt reported in table 8, against varying concentrations of magnesium sulphate, the results reported in table 9 were obtained. There is a marked increase in total ash as the concentration of the nutrient solu- tion with respect to magnesium sulphate increases. Parallel with this increase is the higher percentage of potassium. The growth decreases inversely. Antagonism between the two salts is evident where the lower concentrations of magnesium sulphate were used. In cultures 2 and 4 of this series, we have a marked increase in growth over that of culture 3. Absorption is markedly lower at the two high points than at the intermediate concentration, where the solution is evidently more toxic. The least growth obtained in the series was recorded in culture 7, which shows the highest absorption of all the elements determined. In the two higher concentrations of magnesium sulphate used the growth was increased somewhat while the percentage of cal- cium, magnesium, and potassium in the plants decreased markedly. It seems worthy of note that the amount of iron in the ash was not sufficient to allow of titration at any concentration employed in the series. This series very well illustrates the point which has been brought out a number of times before of the relationship between absorption and growth. Here we have five cultures in the one series of which this relationship is evident. The relations are not absolute in every instance, but there can be no doubt whatever of the tendency toward decreased absorption as growth increases, or that antagonism between ions results in decreased absorption of at least some of the ions present in the nutrient solution. We turn now to a consideration of the effects of a few of the salts of the heavy metals upon growth and absorption. In table 10 the effects of adding various concentrations of aluminum chloride are shown. Growth is decreased in every concentration of the salt used. The high percentage of magnesium is marked in both roots and tops. On the other hand, the percentage of calcium is increased relatively little. The percentage of iron found was practically constant and in total quantity is in marked contrast to the last series considered in which the amount was so small that it could not be determined. In a solution of .20 M. calcium chloride, the results with the vary- 191S] Waynick: Antagonism and Cell Permeability 149 ing concentrations of aluminum chloride are shown in table 11. In general the toxic effects of the two salts seem to be additive, that is. the growth in this series in which two salts are present together is less than in the preceding series where aluminum chloride was used alone. The decrease is not great from the standpoint of total weight, but proportionately is very considerable, amounting to from 33 per cent to 100 per cent in the various concentrations employed. The percentage of magnesium in the two series is about the same. The amount of calcium absorbed, on the other hand, is increased over 300 per cent and remains constant throughout. The total absorption with respect to calcium and magnesium, at least, is uniformly high. This fact is reflected in the increase in the percentage of ash over that of the control. In the next series all factors are the same except that magnesium chloride was used instead of calcium chloride, there being no difference whatever in partial or total concentration. The antag- onism shown between magnesium chloride and aluminum chloride in culture 4 is very marked, and correlated with the increased growth is the marked decrease in the percentage of both magnesium and cal- cium found in tops and roots. The percentage of magnesium found in the plants is not proportional to the concentration in the solution as was true with calcium chloride. An interesting case of the in- creased absorption of one element with a decrease in the other is well illustrated in the case of culture 6 of this series. Such a relationship has been noted previously, but is apparently of no direct importance from the standpoint of growth. Ferric chloride, a second trivalent salt, was used in the nutrient solution in the concentration shown in table 13. In the concentration employed, growth is nearly normal and absorption is very nearly the same as Avith plants in the control cultures, except in the case of cal- cium. The decrease in some instances in the percentage of calcium found, as iron increases in the nutrient solution, is notable, and will be referred to later in connection with the action of ferric and zinc sulphates. The effects of adding .20 M. calcium chloride, together with vari- ous concentrations of ferric chloride, are given in table 14. The growth of roots and top parallel each other closely. Marked toxic effects are evident in certain combinations as in cultures 3 and 7. The percentage of calcium found in both roots and tops is high in plants grown in the same cultures. The magnesium present in the tops shows the same relationships as the calcium, although the 150 University of California Publications in Agricultural Sciences [Vol.3 amount absorbed varies but little from that of the control. In the roots magnesium is present in large amount when growth is low in culture 2, but in succeeding cultures the percentage found falls off sharply and remains abnormally low without any relation to growth or concentration of the solution. The percentage of iron is high in cultures 6 and 7, in which the weight of the plants was small. Substituting magnesium chloride in equivalent concentration for the calcium chloride used in the preceding series, the results are of a very different order from those in table 15. The absolute growth of the tops is greater than in series 14. Root growth does not parallel the growth of the tops. The toxicity of the solution is scarcely evident at some concentrations while markedly increased at others. Absorp- tion, with the exception of the magnesium in the roots, is usually low, amounting to about that of the control, but the percentages of calcium and magnesium found bear no apparent relation to the differences in growth. Iron, however, shows the inverse relation already noted in many other series with calcium and magnesium, that is, high percent- age present when growth is low, and vice versa. The toxic and antag- onistic effects as well may be due in this instance to the ferric ion, but this statement is by no means indisputable. In several tables following, the effects of copper salts are given. Previously copper salts have been shown to be highly toxic to plants as well as to a wide variety of vegetative forms. That they may also be stimulating has been shown recently by Forbes 28 using solution cultures, and by Lipman and Gericke 29 in soil cultures. The reader is referred to the latter paper for an extensive review of the subject. The results with copper chloride are reported in table 16. Growth, especially that of the roots, was limited in every concentration re- ported. In fact, the growth of the roots was so limited that their weights are not given. There is a suggestion of antagonistic action between the nutrient solution and copper chloride in cultures 3 and 5. The percentage of magnesium found is high where growth is low. The same is not true of calcium, the percentage of which is low and decreases as growth decreases to a certain extent. A trace of copper was found in every case and appreciable amounts had penetrated the plant tissue at the two higher concentrations. When ferric chloride is added together with copper chloride marked antagonism is shown. Table 17 will make this effect evident. In this series, as in zs Univ. Calif. Publ. Agr. Sci., vol. 1 (11)17), p. 395. 20 Ibid., p. 495. 191S] Waynick: Antagonism and Cell Permeability 151 several following, the concentrations of both salts added increase, that is, both increasing but bearing the same ratio between the two. There is an increase of approximately 100 per cent in the dry weight of culture 2 over cultures 1 and 3. The low absorption of culture 2 as related to 1 and 3 is evident. There is a marked decrease in the percentages of calcium and magnesium found in the plants grown in culture 5, in which the dry weight of the plants was also low. At this second point, however, iron and copper were found in larger amounts than at any other concentration used. As in the previous series the percentage of calcium in the tops does not seem to parallel that in the roots or of magnesium in either roots or tops. A similar relationship was brought out in the previous series in which copper chloride alone was used. No apparent precipitation took place upon the addition of iron in the concentrations given, but a precipitate composed of ferric phosphate was present at the time of harvesting. It is possible that double salts of copper or iron with calcium or magnesium and, for instance, the phosphate ion were formed at the higher concentrations. Their complexes may not be taken up by the plants and hence actual starvation as far as these elements are concerned, may be responsible for the low amounts found in the plants. Such a condition contrasts directly with one in which there is low permeability due to antagonistic effects between the ions in the solution. In table 18 mercuric chloride was used with copper chloride, since it was desired to determine the effects produced by the addition of two highly toxic salts to the nutrient solution. The results with mer- curic chloride alone are given in table 26. They are somewhat irregu- lar, but there can be no doubt of the correlation between the quanti- tative presence of calcium and magnesium in the tops, of magnesium in the roots, and growth. There is evidence of a distinct antagonistic action between copper and mercuric chlorides both from the stand- point of growth and that of absorption. The root growth was very limited. The percentage of calcium and magnesium in the roots was very high ; high enough to account for the decreased growth by itself if we use the results of other series in interpreting this one. Not enough iron was present in any culture to permit of its determination. Considering the most common salt of copper used in solution cul- tures and soil work, the results as given in table 19 are especially noteworthy. The concentrations of the sulphate used are low. Dis- tinct evidence of the toxic effects of the salt, together with only slight decrease in growth in culture 4 of the series is shown. High percent- 152 University of California Publications in Agricultural Sciences [Vol.3 ages of calcium and magnesium accompany low growth; low percent- ages of calcium and magnesium go with much increased growth. No iron could be quantitatively determined in cultures 8 and 9. The copper content shows no variations which may be regarded as impor- tant, in fact the amount taken up by the plants is somewhat lower where decreased growth is shown. Zinc sulphate was used with copper sulphate as shown in table 20. There is little evidence of antagonism between the two salts. At the same time there is evidently no direct relationship between concen- tration and toxic effect, since growth does not decrease regularly with increasing concentration. While the percentages of calcium and mag- nesium found are somewhat irregular, they increase rapidly as growth becomes less. The percentage of magnesium found in the tops in culture 8 was 1.10 per cent, and in the roots 1.91 per cent. This occurred with the same concentration of the magnesium ion in the nutrient as in culture 1. The percentage of copper found in the dry matter is distinctly larger than that found in the preceding series, in which copper sulphate alone was used. Copper sulphate used with ferric sulphate shows no evidence of antagonism between the two if the growth of the tops alone is con- sidered, but with the roots there is a marked increase in growth in cultures 3 and 4 of the series. The percentages of magnesium found in the roots is low and constant, which contrasts markedly with the amounts determined in the previous series. The calcium likewise varies but little in the tops and its percentage remains low. On the other hand, the percentages of calcium in the tops and magnesium in the roots show marked increases as growth decreases. The amount of iron remains very uniform until the last culture of the series is reached, when a marked increase is recorded. It will be noted that the percentage of calcium decreases to nearly one-third of the original in the same culture. This relation has been noted previously in other series. The stimulation resulting from the addition of ferric sulphate to the nutrient solution in the concentrations given in table 22 is remark- able, a total dry weight of 3.9016 grams for the tops of six plants being recorded. The growth of the roots does not parallel that of the tops. In the highest concentration of ferric sulphate employed, the root growth decreased while the growth of the tops was increased. Attention has already been called to cases of this kind in which there may be an increase in the growth of tops with a decrease in 191S] Waynick: Antagonism and Cell Permeability 153 the root growth, or vice versa. As will be noted, the percentages of calcium and magnesium found are low, in fact below the control in every case. Whether or not ferric sulphate would be stimulating in still higher concentrations is not known, but it is probable that the limit of stimulation was reached, since the roots show a marked de- crease in growth in the highest concentration used. The percentage of iron found is comparatively high. The reason for this increased growth is evidently bound up with the presence of the ferric salt, but no idea of the nature of its action can be given. It is very evi- dent from the present data, however, that the amounts of the elements present in the plants were low. In table 23 the results with zinc sulphate alone are reported. There is no stimulation or no antagonism between zinc sulphate and the other constituents of the solution evident in any concentration. As growth decreases magnesium was found present in larger amounts than in the cultures in which growth was more nearly normal. The percentage of calcium remains very much the same in the tops and decreases rapidly in the roots with decreasing growth. Here we have a suggestion of a relationship between zinc and calcium as has already been referred to in the case of iron. It can only be stated, however, that the results as regards calcium penetration are exceptional in the light of the results in other series previously referred to. Turning to table 24, in which the results with zinc sulphate and ferric sulphate are given, there is a marked contrast on the one hand with series 20 in which zinc sulphate and copper sulphate were used, and on the other hand with the preceding series in which zinc sulphate alone was used. In this series there is marked antagonism shown be- tween the salts employed. This is true for both tops and roots, but the most marked increase in both does not occur in the same culture. The marked increase in growth of the tops evident in culture 4 is accompanied by a decrease in the percentages of calcium and mag- nesium present in the tops but not in the roots. The percentage of magnesium in the roots increases with decreased growth throughout the series. The calcium in the tops is low and abnormally so in the roots. Growth is good throughout the series and in culture 4 is in- creased about 50 per cent above the control. This result would hardly be expected from the decreases recorded where zinc sulphate was used alone in the preceding series. The percentage of iron varies some- what, but does not increase or decrease with any regularity in any one direction. Attention is again called to the low calcium content, especially of the roots. 154 University of California Publications in Agricultural Sciences [Vol.3 Little can be said of the mercuric chloride ferric sulphate series given in table 25. Growth is uniformly low throughout, with con- siderable variation between duplicate cultures. The percentage of magnesium is very high in the roots and while less in the tops, is much above that of the control. The percentage of calcium is uni- formly low in both tops and roots. Attention is called to the fact that no iron could be determined quantitatively, except in the highest concentration of salts used. This condition is striking when the rather large amounts of ferric sulphate in the solution are considered. A short series is reported in table 26 in which the toxic effects of mercuric chloride when used alone, are evident. There is a de- crease in growth with increasing concentration of the added salt and also an increasing percentage of both calcium and magnesium found. The very low ash content given by the plants in this series is of interest and will be discussed below. External Appearances of the Plants It seems worth while to note here a few of the more striking appearances of the plants. Since iron salts were purposely excluded from all solutions except those in which it was planned to study their effects, the control plants were of a more or less yellowish green color. Aside from this no differences were noted between control plants grown with or without the addition of a little ferric phosphate to the nutrient. In every series in which growth was limited by the presence of magnesium salts the roots were short and much thickened. With a high concentration of magnesium in a balanced solution, this effect was not noted however. High concentrations of magnesium were also apparent from the decided yellowing of the older leaves. Excessive amounts of calcium were characterized by the appearance of brown spots or streaks on the leaves. 30 When any considerable growth was permitted the plants grown in solutions of copper salts were dark green in color. 31 Where growth was good the roots were apparently normal. In several of the higher concentrations used, copper hydroxide was deposited upon the roots, especially about the tips. A suggestion is made that possibly copper may replace iron as a catalyzer in connection with the building or activation of chlorophyll. 30 Jost, Plant physiology (Oxford, Clarendon Press, 1907), p. 85. f" CJniv. Calif. Agr. Sci., vol. 1 (1917), pp. 495-588. 1918] Waynick: Antagonism and Cell Permeability 155 Several cultures in which mercuric chloride was used and in which growth was good, displayed the same dark green color as noted for copper salts and the same suggestion as made for the functioning of copper in this color relationship may hold for mercuric salts as well in very dilute solutions. The color was light green when iron salts were present; with the other salts used no marked external effects were noted. General Review of Experimental Results It seems advisable to consider the results reported in the previous tables together, so that the data presented in one table may be more closely correlated with those given in another. It is proposed to do this in the present section and further to discuss briefly the more important relationships shown. It will be noted in the accompanying tables that there is consid- erable variation between the controls grown at different seasons of the year. This was to be expected, since conditions in the green- house varied between the different growing periods. For this reason it is not possible to compare one series of cultures with another so far as absolute weights of the dry matter are concerned. Within any one series or between series grown at the same time the absolute weights are comparable. This point must be borne in mind in con- sidering the results as a whole. In some cultures, however, growth was stimulated to such an extent as to far surpass any variation between series due to differing external conditions. Such a case is that of series 22, in which ferric sulphate was added to the nutrient solution in varying amounts. In culture 5 of this series, the dry weight was over twice that of any control plants grown during the entire time. The experimental work with the salts of calcium plus magnesium was rather extensive. McCooP 2 has reviewed the previous work with calcium and magnesium salts as related to plants, so a discussion of that phase of the relationships between the two need not be entered into here. In his own work McCool found that calcium chloride was effective in antagonizing the poisonous effects of magnesium chloride and magnesium sulphate. He found a slight increase in the growth of pea seedlings over the controls based upon the green weight of the plants. This was the case in distilled w T ater and in nutrient solution. It seems probable that the nutrient solution used by McCool was not 32 Cornell Univ. Agr. Exp. Sta. Mem. 2 (1913), p. 129. 156 University of California Publications in Agricultural Sciences [Vol.3 a balanced solution, since the addition of either magnesium or cal- cium chloride resulted in an increased growth of the pea seedlings. In the present investigation there are only two cases in which the growth of the plants was greater with both calcium and magnesium chlorides present than when calcium chloride was used alone in vari- ous concentrations, one in culture 6, series 2, the other in culture 11, series 3. In the latter culture the dry weight of the plants was twice that in the same concentration of calcium chloride alone. There are marked differences in growth recorded between different combinations and concentrations of the two salts, and as can be easily seen from the graphs, the percentages of the two ions found in the plants show an inverse relation to growth in nearly every instance. Proceeding from series to series, the amount of magnesium found in the plants increases with the concentration of the magnesium chloride in the nutrient solution. Magnesium sulphate is not as toxic as magnesium chloride in equivalent concentrations of the kation. Growth in solutions of mag- nesium sulphate plus calcium chloride was superior in every case to that found when the salts were used separately. There is a marked contrast between calcium chloride and calcium nitrate in antagoniz- ing the toxic effects of magnesium sulphate, the nitrate proving less effective than the chloride in concentrations of .12 M. and over. This is of especial interest, since the qualitative ionic relations of the nutri- ent are not altered. It is possible that we are dealing with the effects of undissociated molecules in the higher concentrations, which may be very different from ionic effects. Results with Salts of the Heavy Metals Since salts of aluminum, copper, zinc, iron and mercury were used, it will be necessary for the sake of clearness to treat each more or less separately. Miyake 33 has shown aluminum chloride to be highly toxic, in con- N centrations above — -, to rice seedlings grown in water cultures. Similar results have been reported by House 34 and Gies, Micheels and De Heen, 35 Duggar, 36 and Ruprecht, 37 working with several aluminum 33 Jour. Biol. Chem., vol. 25 (1916), p. 23. 84Amer. Jour. Physiol., vol. 15 (1905), p. 19. 35 Bull. Acad. Roy. Belg. (1905), p. 520. 36 Plant Physiology, New York, Macmillan, 1911, 37 Mass. Exp. Sta. Bull. 161 (1915), p. 125. 1918] Waynick: Antagonism and Cell Permeability 157 salts. Probably the work of Abbott, Conner and Smalley 38 is of more direct interest here. These investigators found aluminum nitrate to be toxic to corn seedlings in the presence of nutrient solutions. E. Kratzmann 39 has reported stimulation due to the presence of small amounts of aluminum salts. Miyake 40 concludes further that the effects observed with aluminum chloride cannot be attributed to the hydrogen ion resulting from the dissociation of the salt. Aluminum chloride was found to be toxic in every concentration used in the present work. The effect of the presence of calcium chloride in a concentration of .20 M. was to decrease growth still fur- ther, indicating that its toxic effect, as reflected in growth, was but additive to that of aluminum chloride. With magnesium chloride present in equivalent concentration as the calcium chloride, there is a marked antagonism at a concentration of .000066 M. of aluminum chloride with .20 M. magnesium chloride. The increase in dry weight was 100 per cent greater than in an equivalent concentration of aluminum chloride alone and 300 per cent greater than, with mag- nesium chloride in the concentration given. This culture has been referred to especially since it furnishes a striking example of antag- onism between bivalent and trivalent salts, both of which are highly toxic when used alone. The chloride ion was a constant as far as this and the preceding series are concerned, the only difference between the two cases being the use of calcium chloride in one and magnesium chloride in the other. It seems logical to conclude that the action is specific as regards the magnesium and aluminum ions. Whatever the nature of this action may be, it is certainly not shown between calcium and aluminum ions. The same general relationships are brought out between ferric chloride and calcium and aluminum chlorides. Ferric chloride did not prove toxic in the concentrations used, growth differing but little from that of the control. When calcium chloride was present in a concentration of .20 M. throughout the series, growth was half or less than half that recorded when ferric chloride alone was present. Magnesium chloride in equivalent concentrations, as the calcium chloride above, affected growth but little. In other words, magnesium chloride did not prove toxic in the presence of certain concentrations of ferric chloride. The relations between the four salts may be briefly summarized as follows : There is no antagonism shown between alumi- 38 Incl. Exp Sta. Bull. 170 (1913), p. 329. 39Chem. Ztg., vol. 38 (1914), p. 1040. 40 Jour. Biol. Chem., vol. 25 (1916), p. 23. 158 University of California Publications in Agricultural Sciences [Vol.3 num chloride and calcium chloride. There is very little, if any, be- tween ferric chloride and calcium chloride. Magnesium chloride and ferric chloride show marked antagonism in all concentrations used as do magnesium chloride and aluminum chloride in certain concen- trations of the two salts. Magnesium chloride and ferric chloride show marked antagonism in all concentrations as do magnesium chloride and aluminum chloride in one concentration of the latter salt. Reference has already been made to Miss Brenchley's monograph 41 and to the paper by Lipman and Gericke, 42 in which the literature relating to the effects of copper, zinc, and iron salts on plants is reviewed. Suffice it to say that the results reported by different investigators are very conflicting, due largely to the widely different methods used and the varying conditions under which the various data were obtained. In the present work, copper chloride was toxic in every concen- tration used. There was marked antagonism between copper and ferric chlorides both from the standpoint of growth and of absorption. Copper sulphate did not prove to be uniformly toxic. Growth was nearly normal in one concentration used while very much diminished in a lower concentration. The term stimulation might be applied here, but in the present discussion it is applied only when growth due to the presence of an added salt or salts is undoubtedly greater than that in the control. Toxic effects are correlated with increased absorption and antag- onistic effects with decreased absorption as in other series reported. Growth was always less with zinc sulphate present in the nutrient solution than in the latter alone. Copper and zinc sulphate together were no more toxic than a solution of zinc sulphate alone. The case with ferric sulphate is clearly one of stimulation. The dry weight was over twice that of the controls in one concentration of the salt used and far superior in several concentrations to that of the plants grown in the controls. Wolff 43 has reported similar results when iron was used in the form of the citrate, an increase in growth comparable to that noted above having been obtained. He found further that nickel or chromium could not be used to replace iron. The toxic effects of copper sulphate were markedly reduced by the presence of ferric sulphate when we consider the results as a 11 Inorganic plant poisons and stimulants. 1915. '- Univ. Cal. Pub. Agr. Sci., vol. 1 (1917), p. 395. 43C.-E. Acad. Sci. (Paris), vol. 157 (1913), p. 1022. 1918] Waynicl*: Antagonism and Cell Permeability 159 whole, although in one instance growth was greater with copper sul- phate alone than when both salts were added together. The second case of stimulation was noted with zinc sulphate and ferric sulphate in certain concentrations. In series 26 four cultures gave growth superior to that obtained in the control for the series, and throughout growth was good when the two salts referred to above were present together, over the range of concentrations employed. Low absorption was noted. In summarizing the relations of ferric, cupric and zinc sulphates, it is evident, from the discussion above, that zinc sulphate was toxic in every concentration used. Copper sul- phate was toxic, but marked variation in degree was shown between various concentrations. Ferric sulphate was stimulating. Copper sulphate and zinc sulphate were no more toxic together than when each was used alone. Ferric sulphate modified somewhat the toxic effects of copper sulphate. Zinc sulphate and ferric sulphate together proved stimulating to the growth of plants. As contrasted with the chlorides, the sulphates of copper and iron were less toxic to barley over the range of concentrations used in this investigation. Taking the results as a whole, twelve instances of a marked in- crease in growth at certain definite concentrations of one or more added salts have been noted. With every such increase there is a very notable decrease in the amount of calcium and magnesium absorbed. The increase in growth is attributed to antagonistic salt action; de- creased absorption is undoubtedly due to the same action, which tends to preserve the normal permeability of the plasma membrane. In addition to the twelve instances referred to above, we find in series after series, the toxic effects of the solution in which the plants were growing, noticeable not alone by decreased growth but also by increased absorption. The roots and tops may not show the same relations as regards the amounts of calcium and magnesium taken up. For example, in series 25, in which ferric sulphate and mercuric chloride were used together, the toxicity of the solutions was evident by the very limited growth, yet the composition of the tops was about normal. In the roots, however, the percentage of mag- nesium was found to be tremendously increased. It is of interest to refer again to the very low ash content and relatively low absorption, considering the very limited growth, in the few cultures in which mercuric chloride was used alone. It is pos- sible that relatively large amounts of mercuric salts were taken up by the plants which were volatilized on ashing the residue ; thus the low percentage of ash may be less surprising. 160 University of California Publications in Agricultural Sciences [Vol.3 Possible Effects of Variations in the Concentrations of the Solutions on the Plants No attempt was made to maintain the total concentration of the nutrient solution constant. This would be exceedingly difficult to do in work of this character, since it would be necessary to vary the concentration of the nutrient solution to maintain the balance of the solution as regards total concentration. The conclusion seems justi- fied that within the range employed the concentration of the nutrient solution is of minor importance as far as growth is concerned. For instance, in table 1, the variation in the concentration of the solution was .279 M. in terms of calcium chloride, yet the total growth varied but little from .001 M. to .28 M. Again in table 2 the growth is very nearly the same at a concentration of .25 M.. with calcium and mag- nesium chlorides, and a total concentration of .54 M. of the same salts. In table 3 the greatest growth occurred in a concentration of .46 M. in terms of the salts above mentioned, while at the lower concentra- tions of .304 M., growth was but a third that obtained in the higher concentrations. These examples make clear the point above men- tioned, namely, that the concentration over the range used was of but minor importance. It is obvious that the above discussion does not apply to the series in which salts of the heavy metals were used, since the variations in concentration in those series were but slight. Consideration of a Possible Calcium-Magnesium Ratio Since Loew 44 first advanced the hypothesis of the lime-magnesia ratio, much experimental evidence has been collected by various inves- tigators both for and against the existence of an optimum ratio be- tween these two elements as regards the growth of plants. The literature bearing upon the subject has been very fully reviewed by Lipman, 45 so that detailed references are not necessary here. Since the ratios of calcium to magnesium in the solution used by the writer were known and also because of the fact that the analytical data allowed of the calculation of such a ratio for the plants, it seemed of interest to present some of these data here. The following two tables give the results obtained from two series in which widely varying proportions of calcium and magnesium were used. 44 Flora, vol. 75 (1892), p. 368. >■> Plant world, vol. 19 (1916), p. 83. 1918] Waynick: Antagonism and Cell Permeability 161 Table , 27 Rati Mgto in soli, 41 Ca tion 1 Dry weight tops .3536 Ratio Mg to Ca in tops 2.3 : 1 Dry weight, roots .1218 Ratio Mg to Ca in roots 1.2 : 1 16 1 .5484 4.6 : 1 .1519 1 2 8 1 .5885 6.0 : 1 .1266 1 4.1 1 .4774 2.6 : 1 .1509 3.3 2.7 1 .3433 2.6 : 1 .1497 1.2 2.0 1 .6775 2.7 : 1 .2119 1.3 1.6 1 .4136 1.2 : 1 .1500 1.3 1.3 1 .4431 1 : 1.2 .1421 1.2 1 1 .3745 1.3 : 1 .1138 1 1.3 1 1.2 .3268 1.5 : 1 .1254 1 1.5 1 1.4 .2815 1.8 : 1 .1053 1 1.2 1 1.8 .5030 1 : 1.2 .1044 1.2 Table 27 was computed from the results given in table 2. Mag- nesium chloride was present in uniform concentration of .24 M. with varying concentrations of calcium chloride. It will be noted that the dry weights with a ratio of magnesium to calcium of 16 : 1, 8 : 1, and. 1 : 18 are nearly the same. The ratios of these two elements found in the plants grown in these solutions were 2 : 1, 1 : 1. 1 : 1.2 for the roots, and 1 : 4.6, 1 : 1.6, 1 : 1.2 for the tops. Further, the dr}^ weight of plants grown in a solution in which the ratio was 41 : 1 and with a ratio of 1 : 1 are nearly the same. It is evident that the same ratio for the roots may not hold for the tops. Table 28 Ratio Mg to Ca in solution Dry weight, tops Ratio Mg to Ca in tops Dry weight, roots Ratio Mg to Ca in roots 20.2 : 1 .3033 5.5 : 1 .0822 4.4 : 1 10.5 : 1 .4716 5.4 : 1 .1572 4.7 : 1 6.8 : 1 .2799 4.8 : 1 .0780 6.3 : 1 5.1 : 1 .1999 5.8 : 1 .0342 6.0 : 1 4.0 : 1 .1999 4.8 : 1 .0636 7.4 : 1 3.4 : 1 .4363 1.3 : 1 .1122 4.1 : 1 2.5 : 1 .4013 1.8 : 1 .1143 2.8 : 1 2.0 : 1 .2734 2.3 : 1 .0677 4.7 : 1 1.7 : 1 .2603 2.5 : 1 .0867 6.8 : 1 Table 28 gives the ratios in the solutions used in series 4, in which the ratios of magnesium to calcium varied from 20.2 : 1 to 1.7 : 1. Growth is nearly the same in solutions in which the ratio was 10.5 : 1 as in those in which the ratio is 3.4: 1 or 2.5: 1. The plants grown in these cultures gave the following values for the tops : 5.4 : 1, 3.4 : 1, 2.5 : 1, and for the roots, 4.7 : 1, 4.1 : 1, and 2.8 : 1. There is a tendency for the ratio of calcium to magnesium in the plants to become narrower 162 University of California Publications in Agricultural Sciences [Vol. 3 as the ratio of these two ions in the solution becomes narrower. Where a wide ratio exists in the solution, there is always a much nar- rower ratio in the plants. From the brief discussion above it is evident that the barley plants grew equally well in solutions having widely different ratios of cal- cium and magnesium ions. There is no "optimum lime-magnesia ratio," as Gile 46 and Wyatt 47 as well as others have shown, and their results are confirmed in the present investigation. The balance between all the ions present in the solution appears to be of far greater importance than any single ratio. A considera- tion of the ratios existing between the various ions of the nutrient solution, aside from calcium and magnesium used, is reserved for further study. Permeability and Antagonism It is not proposed to enter into a discussion of the structure and composition of the plasma membrane. Davidson 48 has recently sum- marized our present knowledge concerning it with special reference to selective permeability. A discussion of the various theories which have been advanced to explain antagonistic salt action need not be taken up in detail here. The reader is referred to papers by Clark, 49 Loeb, 50 Osterhout, 51 Loew, 52 Koenig and Paul, 53 True and Gies 54 , True and Bartlett, 55 Kearney and Cameron, 65 and Ostwald 57 , for a discus- sion of the various factors which may be of importance in this con- nection. The recent work of Clowes 58 and Fenn 59 is important and some very striking similarities between the action of toxic and antagonistic solutions on oil emulsions and on gelatine on the one hand, and plant cells on the other, have been reported by these investigators. 46 Porto Eico Exp. Sta., Bull. 12 (1912). 47 Jour. Agr. Research, vol. 6 (1916), p. 589. 48 Plant World, vol. 19 (1916), p. 331. 40Bot. Gaz., vol. 33 (1902), p. 26. ■ r >oArchiv. ges. Physiol., vol. 88 (1902), p. 68. 5i Science, n.s., vol. 35 (1912), p. 112. 52 Flora, vol. 75 (1892), p. 368. r > 3 Zeitschr. Hygiene u. Infektionskranklieiten, vol. 25 (1897), p. 1 54 Bull. Torr. Bot. Club, vol. 30 (1903), p. 390. ■>■> U. 8. Dept. Agr., Bull. 231, 1912. • r '0 U. S. Dept. Agr., Bull. 71, 1902. 57 Archiv. ges. Physiol., vol. 120 (1907), p. 19. 58 Jour. Phys. Chem., vol. 20 (1916), p. 407. 59Proc. Nat. Acad. Sci., vol. 2 (1916), p. 539. 1918] IV ay nick: Antagonism and Cell Permeability 163 To define normal permeability is very difficult. There seems to be a comparatively wide range of concentration of salts over which the amount of any element taken up may vary without affecting the growth of the plant to any considerable extent. There is likewise a wide range over which the ratio of any one element to any other may change without being detrimental to plant growth. The latter point has been discussed above in connection with a possible optimum calcium-magnesium ratio for plants. The first point referred to has been very well treated by Gile and Ageton, 60 so that further reference need not be given here. For the work in hand the percentage composition of the plants grown in the control cultures seemed to be the most logical criterion of normal permeability available. There are variations between the controls as regards composition, but they are relatively small. On the other hand, the percentages of magnesium, for instance, range from .02 per cent to 9.21 per cent, depending upon the solution used. The percentages of calcium differ over a wide range as well. From the data presented there can be no doubt whatever that the composi- tion of the plant, as regards inorganic constituents at least, may be altered enormously by variations in the surrounding solution. That portion of the root system in any plant which functions as a semipermeable membrane is obviously of greatest importance in a study of the present kind. The actual area of the membrane which is in contact with the solution must be known in every case before it can be said that the permeability of one root system is greater than that of another. The actual area of the plasma membrane cannot be measured directly because, in the first place, we have no means of determining just how much of the root is involved, and secondly, the area concerned may be changing continually. Length of the roots and their number and length together as well as green weight and dry weight have been taken as criteria of the existence of antagonism. In the present paper the dry weight has been taken as proportional to the area of the plasma membrane through which salts may enter the plant. It cannot be stated defi- nitely that the two are proportional. They have only been so con- sidered since the dry weight of the plant was the most logical criterion to employ. The reservation must always be made that the two may not be directly proportional, even though they are treated as being so. That the permeability of the plasma membrane of the plant cells so Porto Eico Agr. Exp. Sta. Bull. 16, 1914. 164 University of California Publications in Agricultural Sciences [Vol. 3 is changed by the nature and balance of the solution surrounding the roots there can be no doubt from the data already given. That a number of ions are capable of acting in a very similar manner to one another as regards permeability is also evident from the present work. Further, the same salt may act differently at different concentrations, preserving nearly normal permeability at some and allowing the pene- tration of large numbers of ions at others. As previously stated, the total balance of the solution is of vital importance in the preservation of normal permeability, which is in turn correlated with normal growth. In connection with the salts of the heavy metals, the amounts of the kation of cupric and ferric salts which had penetrated the plant tissue were determined in a number of instances. The percentages found were low. Further, whenever these salts proved toxic, the amounts of calcium and magnesium found in the plants were high ; high enough in fact to account for the toxic effect alone. In many instances the percentages of those two elements found were as high in toxic solutions of copper, iron, or zinc salts as when toxic concentra- tions of calcium or magnesium chlorides were used. We might, therefore, in the light of our present knowledge, be justified in attrib- uting the decreased growth of the plants to the abnormally high ab- sorption of calcium and magnesium and the consequent reactions taking place within the plant cells. The permeability of the mem- brane must be altered to allow of the presence of these ions in large numbers. The toxic effects due to the presence of large amounts of calcium or magnesium salts might be evident if we could inject solu- tions of these salts into the plant without altering the permeability of the plasma membrane. But from the present data it seems that the alteration in the permeability of the membrane is the essential consideration. It is probable also that the toxicity of any solution is accompanied by the increased permeability of the plant tissue to all inorganic salts which are normally found in plants. There may be exceptions as noted already for iron and calcium, but in general this relation holds from the data now at hand. Ruprecht 61 has localized the effects of aluminum salts in the few layers of cells surrounding the root hairs and attributes the death of the plants grown in solutions of aluminum salts to starvation incident upon the inability of the plant to obtain nutrient salts for normal ii Mass. Exp. Sta. Bull. 161 (1915), p. 125. ]918] Way nick: Antagonism and Cell Permeability 165 metabolism. Forbes 62 has likewise localized the effects of copper salts, when present in toxic concentrations, and concludes that the toxic effect of copper is due to the combination of metal with protein at the growing tips of the roots. From the experimental results given in the present paper, it is evident that the presence of the salts of each element in toxic concen- tration results in an increased permeability of the plant tissues to calcium and magnesium at least. Ruprecht's view that plants starve for lack of nutrient salts when grown in toxic solutions is untenable, in the light of the above discussion. The results of both investigators are significant in indicating the localization of the effect of the two metals studied in the extreme outer portion of the roots, in which the plasma membrane is located. The results obtained by Loeb with Fundulus eggs, by Osterhout with Laminaria, using electrical conductivity methods, and by Brooks employing microscopical methods with various plant tissues, all point to the preservation of normal permeability as the result of antago- nistic salt action. The results reported by these investigators using widely different methods have been confirmed in the present work by the use of a more direct and more nearly quantitative method than any hitherto employed. It must be recognized, however, that a picture of but one stage in the growth of the plant has been given and that only a portion of the inorganic constituents have been determined. The results reported are essentially those of a static system and must be so considered in comparing them with results obtained by the use of other methods referred to above. Summary In the present paper results are given showing the effect of vari- ous salt solutions upon the chemical composition of plants, with spe- cial reference to a correlation between toxic and antagonistic effects and composition. A uniform nutrient solution was used throughout. The cultures were arranged in series in which the concentration of one salt was kept constant while the concentration of a second salt varied over a wide range. In several series the concentration of both varied, but the ratio between the two remained constant. The ana- lytical data cover the percentages of calcium and magnesium found in the plants grown in every culture, together with determinations 62 Univ. Cal. Publ. Agr. Sci., vol. 1 (1917), p. 395. 166 University of California Publications in Agricultural Sciences [Vol.3 of potassium, iron and copper in certain series. With these facts in mind the results of the investigation may be briefly stated as follows : The composition of the plants grown in different solutions varied widely. Normal growth, i.e., approximately that of the controls, was always accompanied by approximately equal percentages of calcium and magnesium in the plants. In nearly all cases in which the growth of the plants was decreased to a marked extent, the amounts of the two elements referred to above were increased greatly. The degree of absorption of any salt seems to be independent of the concentration present in the solution over a wide range. Certain relationships are pointed out between calcium and mag- nesium absorption and the presence of iron and zinc salts in the solution. Antagonism as evidenced by growth is correlated with absorption of the ions, which were determined, in every instance. Stimulation of growth was recorded when ferric sulphate was present in the nutrient solution in certain concentrations and with ferric sulphate and zinc sulphate together. The amounts of the two ions uniformly determined were not neces- sarily found in the same proportions in roots and tops. The possible effects of changes in concentrations of the various solutions are considered, and the conclusion reached that the changes in concentration were of secondary importance over the range of con- centrations of the various salts used. Data are presented showing that growth is the same with widely varying ratios of calcium to magnesium in the nutrient solution. The results in general confirm those of Loeb, Osterhout, and Brooks in finding that antagonistic salt action tends toward the preservation of normal permeability of the plasma membrane in living tissue. This problem was suggested by Dr. C. B. Lipman. The writer wishes to express his thanks for this and for many other valuable suggestions offered while the work was in progress. The writer is also indebted to Prof. L. T. Sharp for helpful advice. 1918] Waynick: Antagonism and Cell Permeability 167 NOTE The following key applies to all the graphs. The numbers on the abscissas represent both the actual weight of tops and roots and percentages of calcium and magnesium, or of iron, when the latter were plotted. The numbers on the ordinates correspond to the number of cultures as given in the table on the opposite page. The heavy lines always refer to the roots, the light lines to the tops. The following type lines are used: (Solid line) Weight of tops. (Short dashes) Weight of roots. (Long dashes) Percentage of calcium. (One long and two short dashes) Percentage of magnesium. (One long and one short dash) Percentage of iron. The numbers given in the "Explanation of Plates" always refer to the plants arranged in order from left to right, the control being on the extreme right in every case. 168 University of California Publications in Agricultural Sciences [Vol. 3 Table 1 Calcium Chloride No. 1 Solution CaCl 2 .002 .004 10 11 12 Ful .01 .02 .04 .06 .08 .10 .12 .16 .20 .24 Dry weight Tops .7633 .7104 Roots .3608 Tops .6486 .4976 Roots .6555 Tops .6419 .6138 Roots .5628 Tops .4814 .6600 Roots .5750 Tops .5950 .5442 Roots .5959 Tops .5692 .4998 Roots .5048 Tops .6706 .5015 Roots .5250 Tops .6114 .4182 Roots .3827 Tops .4832 .4778 Roots .3918 Tops .5668 .5218 Roots .6123 Tops .5687 .7637 Roots .6305 Tops .5266 .5793 Roots .6067 Nutrient Tops .7937 .7418 Roots .6900 .7368 .1804 .5731 .3277 .6279 .2814 .5707 .2875 .5696 .2979 .5845 .2524 .5860 .2625 .5148 .1913 .4805 .1959 .5443 .3066 .6662 .3154 .5529 .3033 .7677 .3450 I « ft* o 15.38 15.34 26.24 15.34 17.34 29.98 17.29 16.81 28.10 16.80 17.69 31.56 17.52 16.59 33.54 17.67 17.00 29.45 13.92 11.82 29.46 18.46 19.94 30.18 16.80 17.15 29.12 1 7.45 18.47 29.43 17.03 17.13 31.82 17.62 17.34 27.08 18.80 19.10 20.03 Pi 15.36 16.34 17.05 17.24 17.05 17.33 12.87 19.20 16.97 17.96 17.08 17.48 18.95 p4 .477 .231 .400 .470 .261 .514 .517 .219 .504 .450 .121 .640 .565 .113 .718 .713 .098 .407 .684 .363 .428 .440 .290 .443 .425 .273 .387 .486 .392 .348 .388 .204 .354 .392 .373 .393 .390 .227 pi cS g« 3 be a 1 * go 3 a) c fee Ph a 0) S ® ? ° S3 No. MgClo CaCl 2 4) 3 1 .24 .004 Tops .3407 12.90 .483 1.21 .3666 .3536 13.00 12.95 .500 .491 1.06 1.16 Eoots .2456 .1218 20.05 .557 .670 .022 2 .24 .01 Tops .5106 16.84 .198 .883 .5862 .5484 16.88 16.86 .196 .197 .953 .918 Eoots .3038 .1519 16.75 .437 .216 .025 3 .24 .02 Tops .2758 15.22 .490 1.45 .5885 .4321 15.94 15.58 .189 .339 1.09 1.13 Boots .2533 .1266 21.02 .407 .433 .010 4 .24 .04 Tops .4450 16.30 .361 .967 .943 .4828 .4619 .17.55 16.92 .342 .351 .920 .943 Eoots .3018 .1509 16.32 .550 1.82 .009 5 .24 .06 Tops .3150 18.50 .445 1.07 .3716 .3433 17.87 18.18 .403 .422 1.13 1.10 Eoots .2994 .1497 19.54 .309 .388 .006 6 .24 .08 Tops .6871 15.97 .154 .440 .6679 .6775 15.97 .172 .163 .441 .440 Eoots .4238 .2119 14.25 .206 .280 .007 7 .24 .10 Tops .4797 16.75 .458 .645 .4476 .4636 13.23 14.99 .457 .457 .507 .576 Eoots .3000 .1500 19.95 .430 .577 .002 8 .24 .12 Tops .4583 16.20 .817 .665 .4279 .4431 17.43 16.81 .732 .774 .635 .650 Eoots .2843 .1421 23.09 .376 .454 .006 9 .24 .16 Tops .3243 15.14 1.160 1.46 .3543 .3393 15.45 15.29 1.210 1.18 1.59 1.52 Eoots .2276 .1138 24.60 .970 .714 .010 10 .24 .20 Tops .3404 16.62 1.140 1.69 .3132 .3268 17.50 17.06 1.330 1.23 1.98 1.83 Eoots .2508 .1254 24.60 1.200 .785 .012 11 .24 .24 Tops .2707 18.53 1.640 .807 .015 .2924 .2815 17.83 18.18 1.290 1.46 .826 .816 .009 .012 Eoots .2107 .1053 24.62 1.200 .995 .010 12 .24 .30 Tops .4834 18.48 .563 .675 .010 .5226 .5030 17.64 18.06 .585 .574 .707 .691 .012 .012 Eoots .3888 .1944 25.16 .437 .538 .010 13 .24 Tops .4281 16.83 .387 .907 .018 .5106 .4693 17.20 17.02 .383 .385 .890 .898 .013 .015 Eoots .2463 .1231 23.07 .208 .800 .011 Grown January 2-February 13, 1916. 1918] Waynick: Antagonism and Cell Permeability 171 .1 _ li 13 Fig. 2 Magnesium Chloride + Calcium Chloride (See Table 2) 172 University of California Publications in Agricultural Sciences [Vol.3 Table 3 Magnesium Sulphate + Calcium Chloride Solution No. 1 9 10 1 1 ll 1 MgCl s .30 .30 .30 .30 .30 .30 .30 .30 .30 .30 .30 .30 13 .30 14 .30 Full Nutrient CaCl 2 .001 .002 .004 .01 .02 .04 .06 .08 .10 .12 .16 .20 .24 .30 Dry Weight Tops .1986 Eoots .2650 Tops .2700 Roots .2766 Tops .2566 Roots .2236 Tops .3194 Roots .1744 Tops .2074 Roots .2249 Tops .5249 .4166 Roots .1309 Tops .3036 .1836 Roots .0850 Tops .2184 .2403 Roots .0648 Tops .3978 Roots Tops .3842 .2759 Roots .1059 Tops .8819 .9600 Roots .6900 Tops .2100 Roots .2700 Tops .2576 Roots .1930 Tops .2600 Roots .2000 Tops .7937 .7418 Roots .6900 a cS .2318 .2733 .2401 .2469 .2163 .4707 .0654 .2436 .0425 .2293 .0324 .3300 .0529 .9209 .3450 24.00 .2253 .2300 .7677 .3450 16.60 15.60 16.44 16.39 18.43 17.39 16.49 14.28 14.56 15.73 15.19 15.57 18.10 16.83 17.42 14.70 17.90 15.99 13.05 30.08 17.92 19.57 27.29 17.41 18.11 33.71 18.80 19.10 20.03 16.10 16.41 17.91 15.38 15.14 15.38 17.12 16.94 30.08 18.74 17.76 14.08 15.40 14.30 18.95 .300 .209 .198 .143 .216 .274 .229 .189 .526 .423 .396 .443 .254 .145 .263 .235 .213 .104 .525 .329 .388 .623 .136 .184 .435 .393 .390 2'27 .259 .170 .255 .209 .526 .409 .254 .224 .525 .358 .160 1.390 1.490 2.200 .391 M C3 ££ .440 .413 .571 .514 .595 .533 .515 .088 .749 .565 .405 .437 .649 .790 .700 .663 .230 1.15 .399 .563 .124 .047 .037 .047 .235 .202 .259 .426 .542 .564 .515 .607 .418' .719 .681 1.15 .481 .042 3.660 4.210 3.790 .218 Grown October 24 to December 5, 1915. 1918] WaynicTc: Antagonism and Cell Permeability 173 • chloride + Calcium Chloride Magnesium Chloride -r 8 (See Table 3) 174 University of California Publications in Agricultural Sciences [Vol.3 Table 4 Magnesium Chloride + Calcium Chloride Solution A Dry Weight pi /M ■/ v--r 4 Fig. 8 Potassium Chloride (See Table 8) 184 University of California Publications in Agricultural Sciences [Vol. 3 Table 9 Magnesium Sulphate + Potassium Chloride Solution A Drv Weight a 03 3 bfl s -a a So c3 i * CD P-l o3 OJ 3 03 C3 p-i 03 C 03 No. MgS0 4 KC1 Oj 3 1 .04 .18 Tops .6579 19.65 .141 .490 2.13 .6379 .6479 21.20 20.42 .162 .151 .500 .495 1.72 1.92 Eoots .2800 .1400 20.00 2.79 .67 2 .08 .18 Tops .6800 22.07 .138 .430 3.21 .7162 .6981 25.10 23.58 .171 .154 .480 .455 3.00 3.10 Boots .2682 .1341 23.50 .590 1.350 .72 3 .12 .18 Tops .4957 22.60 .292 3.78 .5202 .5079 26.50 24.55 .310 .301 .923 .923 3.82 3.80 Roots .2200 .1100 25.25 .745 .104 .86 4 .16 .18 Tops .6229 23.50 .152 .382 4.12 .6464 .6346 24.70 24.10 .169 .161 .381 .381 d 4.21 4.16 Eoots .3123 .1561 22.70 .291 .561 f-{ 1.40 5 .20 .18 Tops .3643 .4819 .4231 26.20 26.20 .440 .440 .977 .977 03 O 5.01 5.01 Roots .2063 .1031 24.00 .678 1.060 4J 2.20 6 .24 .18 Tops .3259 33.20 .407 1.040 7.22 .2141 .2700 37.60 35.40 .423 .415 2.920 1.970 O o 8.13 7.66 Roots .1297 .0648 22.10 .750 1.490 +3 2.80 7 .28 .18 Tops .1522 31.10 .299 1.030 o 9.20 .2050 .1786 35.10 33.10 .265 .282 .990 1.010 a 11.30 10.25 Roots .0913 .0456 24.10 • .735 2.900 < 3.10 8 .32 .18 Tops .2704 36.10 .263 .890 11.20 .2997 .2850 36.20 36.15 .211 .237 .810 .850 7.20 9.20 Roots .1467 .0733 22.57 .229 .818 2.17 9 .36 .18 Tops .2800 23.90 .261 .801 3.21 .2846 .2823 25.00 24.45 .278 .269 .895 .848 4.17 3.64 Roots .1417 .0708 21.60 .699 .248 2.18 Fu] 11 Nutrient Tops 1.0992 20.17 .310 .268 1.0750 1.0872 19.12 18.69 .297 .303 .228 .224 Roots .8120 20.00 .271 .233 Grown January 31-March 13, 1916. 191S Waynick: Antagonism and Cell Permeability 185 1 .4 1.3- 1 . 2 1 . 1 1 . . 9 0.8- . 7 . 6 . 5 0.4- 0.3- . 2 .1 Fig. 9 Magnesium Sulphate + Potassium Chloride (See Table 9) 186 University of California Publications in Agricultural Sciences [Vol. 3 Table 10 Aluminum Chloride £S No. Solution A1C1 3 Dry Weight P4 c8 3 3 Ph 03 3 03 1 .0000033 Tops .4438 22.42 .098 .450 .075 .4688 .4563 20.40 21.40 .092 .095 .590 .520 .077 .076 Eoots .4315 .2157 20.20 .055 .590 .089 2 .0000165 Tops .2934 23.20 .196 .793 .151 .2745 .2839 22.70 22.95 .180 .188 .793 .793 .140 .145 Roots .1976 .0988 18.70 .250 .990 .061 3 .000033 Tops .2872 21.09 .316 .953 .173 .3595 .3233 22.60 21.84 .321 .318 1.000 .970 .154 .163 Eoots .2964 .1482 19.55 .153 .504 .130 4 .000066 Tops .3028 21.09 .342 .605 .128 .3200 .3114 22.80 21.94 .334 .338 .615 .610 .155 .141 Roots .2422 .1211 19.55 .334 .840 .171 5 .000132 Tops .2720 22.60 .423 .835 .101 .2479 .2599 19.00 20.80 .450 .436 .837 .836 .134 .117 Roots .2645 .1322 26.60 .382 1.110 .104 6 .000331 Tops .3178 20.20 .246 1.130 .137 .4016 .3597 21.90 21.05 .247 .246 .690 .91 .124 .130 Roots .2550 .1275 27.90 .108 .108 7 .00331 Tops .3863 22.30 .258 .535 .114 .2369 .3116 21.82 22.60 .268 .263 1.230 .88 .187 .150 Roots .2153 .1076 29.40 .583 .330 .231 Grown August 26-October 7, 1916. 1918] Wayniclc: Antagonism and Cell Permeability 187 1 .2 1 . 1 1 .0 0.9 - .7 0.6- .5 . 4 .3 .2 .1 I I Fig. 10 Aluminum Chloride (See Table 10) 188 University of California Publications in Agricultural Sciences [Vol.3 Table 11 No. 1 Cal cium C !hlorid( i 4- Aluminum Chloride Solution A Dry Weight 03 u es I" ® 3 Percentage of Ca Mean 03 C fcJD 03 3) 3 03 M 03 a A1C1 3 CaCl 2 .0000033 .20 Tops .2962 24.20 1.630 .745 .2087 .2524 23.10 23.65 1.820 1.72 .830 .787 Eoots .1897 .0948 20.40 .730 .872 .0000165 .20 Tops .2542 24.23 1.610 .890 .123 .2678 .2610 22.18 23.2'0 1.720 1.66 9.30 .910 .313 .21 Eoots .2745 .1372 26.17 .731 .0000331 .20 Tops .3324 23.00 1.250 .787 .503 .3706 .3515 25.20 24.10 1.460 1.35 .903 .845 .540 .52 Roots .2942 .1471 26.10 .790 .817 .075 .0000662 .20 Tops .2139 23.20 1.210 .821 .237 .2337 .2238 21.70 22.45 1.030 1.12 .733 .772 .114 .17 Eoots .1839 .0919 24.20 .621 1.02 .000132 .20 Tops .2148 20.58 1.400 .932 .248 .18 .3085 .2611 24.10 22.34 1.060 1.23 .897 .913 .119 .18 Eoots .2239 .1119 20.20 .513 .947 .000331 .20 Tops .2337 24.10 1.420 .632 .105 .2137 .2237 21.32 22.71 1.270 1.34 .711 .621 .210 .15 Eoots .1682 .0841 20.16 .672 1.310 00331 .20 Tops .2496 24.10 1.330 .490 .155 .1946 .2221 22.90 23.50 1.550 1.44 .516 .503 .285 .22 Eoots .1296 .0648 26.40 .780 .253 .021 Nutrient Tops .7103 18.17 .322 .221 .7327 .7215 19.31 18.74 .377 .349 .270 .245 Eoots .6600 .3300 19.99 .300 .233 Crown August 26-October 7, 1916 1918] IV ay nick: Antagonism and Cell Permeability 189 1 .4 1 .3 1 . 2 1.1 - 1 .0 0.9 - 0.8 - 0.7- 0.6 - 0.5 - .4 0.3 - 0.1 - Fig. 11 Calcium Chloride + Aluminum Chloride (See Table 11) 190 University of California Publications in Agricultural Sciences [Vol. 3 Table 12 Aluminum Chloride + Magnesium Chloride Solution A Dry Weight 03 Ph 03 a 3 fe«H Pi 03 3 03 No. A1C1 3 MgCl 2 3 1 .0000033 .20 Tops .2819 19.02 .426 2.13 .121 .5457 .4138 18.71 18.86 .371 .398 1.91 2.02 .131 .126 Eoots .2936 .1468 21.31 .321 2.12 .122 2 .0000165 .20 Tops .3539 17.31 .327 1.71 .141 .3000 .3269 19.21 18.16 .421 .374 1.83 1.77 .132 .136 Eoots .1750 .0875 22.17 .400 1.71 .101 3 .0000331 .20 Tops .2243 21.05 .352 2.51 .148 .2259 .2251 17.70 19.39 .448 .400 2.80 2.65 .147 .147 Eoots .1239 .0614 24.10 .366 1.96 .135 4 .0000662 .20 Tops .6327 19.00 .147 1.46 .050 .7644 .6985 18.10 18.55 .130 .138 1.36 1.42 .047 .049 Eoots .5526 .2763 19.95 .111 .333 .040 5 .000132 .20 Tops .2406 19.75 .280 2.78 .069 .3156 .2781 19.55 19.65 .282 .281 3.11 2.94 .064 .066 Eoots .1250 .0625 24.00 .186 5.88 .181 6 .000331 .20 Tops .2279 16.50 .407 .940 .091 .2048 .2163 19.90 18.20 .467 .437 .557 .74 .081 .086 Eoots .1700 .0850 22.76 .302 2.51 .098 7 .00331 .20 Tops .2400 17.40 .131 2.14 .161 .1450 .1925 18.80 18.10 .161 .145 1.71 1.92 .192 .176 Eoots .0990 .0495 32.60 .200 4.53 .439 Grown August 26-October 7, 1916. 1918] JVaynicJc: Antagonism and Cell Permeability 191 1.4- 1.3- 1 .2 1.1- 1.0 - 0.9 0.8 0.7 - .6 0.5 - 0.4- .3 .2 0.1 - \^ - I I I I I 12 3 4 5 Fig. 12 Aluminum Chloride + Magnesium Chloride (See Table 12) 192 University of California Publications in Agricultural Sciences [Vol.3 Table 13 Ferric Chloride No. Solution FeCl 3 Dry Weight 3 Ph 3 So Ph Ph c 5! O) £^ Sh«w Ph 3 1 .000089 Tops .7712 14.50 .199 .258 .214 .8376 .8044 19.20 16.85 .228 .213 .237 .247 .199 .206 Roots .7143 .3571 2 .000168 Tops .7884 17.90 .138 .244 .200 1.2762 1.0323 19.30 16.60 .106 .122 .299 .271 .197 .198 Roots .6462 .3231 17.60 .178 .250 .220 3 .00168 Tops 1.3519 15.70 .045 .259 .247 1.1269 1.2394 18.10 16.90 .046 .045 .225 .242 .232 .239 Roots .9034 .4517 17.10 .054 .213 4 .0168 Tops 1.0000 16.78 .033 .254 .109 1.2750 1.1370 16.51 16.64 .028 .030 .222 .238 .138 .123 Roots .7219 .3609 20.98 .068 .203 .231 Full Nutrient Tops 1.0992 20.17 .310 .268 1.0750 1.0872 19.12 18.69 .297 .303 .228 .224 Roots .8120 20.00 .271 .233 Grown January 24-March 6, 1916. 1918] Waynick: Antagonism and Cell Permeability ]93 1 . 4 1 .3 1.1- 1 .0 .9 .7 0.6 .5 .4 .3 .2 .1 v^—-"~- i i i i Fig. 13 Ferric Chloride (See Table 13) 194 University of California Publications in Agricultural Sciences [Vol. 3 Table 14 Ferric chloride 4- Calcium Chloride Solution A, Dry Weight 'a 3 0-3 Ph 3 go No. FeCl 3 CaClo PM 1 .000089 .20 Tops .4826 18.27 1.02 .4116 .4471 18.50 18.38 1.83 Eoots .4143 .2071 28.35 .531 2 .000168 .20 Tops .5238 17.72 1.77 .5847 .5592 16.04 16.88 1.01 Eoots .6115 .3057 22.70 .188 3 .000352 .20 Tops .2732 20.42 2.78 .3204 .2918 20.61 20.51 3.28 Eoots .1774 .0887 22.40 1.76 4 .000712 .20 Tops .5910 .3810 .4860 19.01 19.01 2.85 2.15 Eoots .5053 .2526 31.30 1.06 5 .00142 .20 Tops .5427 16.56 1.06 .6353 .5890 17.79 17.17 1.33 Eoots .6238 .3119 29.50 .350 6 .00356 .20 Tops .2715 14.76 1.32 .2632 .2673 17.09 15.92 1.23 Eoots .2196 .1098 20.03 1.13 7 .0058 .20 Tops .1792 20.03 2.37 .2514 .2153 20.92 20.47 2.08 Eoots .0636 .0318 29.30 1.68 8 .0168 .20 Tops .3293 17.03 .584 .4393 .3843 19.08 18.05 .460 Eoots .3565 .1783 28.04 .505 «4H ^ 1.42 1.39 3.03 2.00 1.19 1.27 2.22 .522 Grown December 9-January 19, 1916. Tron determined colorimetrically in this series. .164 .137 .222 .194 .101 .229 .256 .255 .692 .212 .262 .026 .064 .051 .010 .097 .133 .018 .256 .297 .020 .246 .175 .049 .150 .147 .255 .237 .058 .115 .276 .210 .100 .080 .090 .03 .085 .370 .400 .38 .120 .50 .40 .02 .36 .50 .16 .45 .43 1.18 1.02 1.10 .98 .800 .900 .200 .148 .120 .06 .134 191S] WaynicJc: Antagonism and Cell Permeability 195 1.4- 1 .3 1.2- 1 .1 1 .0 .9 . 8 .7 0.6- \ / \ . 5 .4 Fig. 14 Ferric Chloride + Calcium Chloride (See Table 14) 106 University of California Publications in Agricultural Sciences I Vol. 3 Table 15 Ferric Chloride + Magnesium Chloride Solutic A in Dry Weight 03 13 a jo n 03 C be 03 O C 03 Ah c3 No. FeCl s MgCls 3 1 .0000S9 .20 Tops .9650 18.48* .182 .874 .091 .7444 .8547 18.70 19.59 .163 .172 .833 .853 .111 .101 Eoots .6126 .3063 25.05 .204 .547 .047 2 .000168 .20 Tops .7875 18.12 .181 .189 .210 .9525 .8700 18.67 18.44 .164 .172 .191 .190 .210 Eoots .6816 .3408 21.40 .229 .553 .033 3 .000352 .20 Tops .8787 15.17 .172 .218 .067 .7365 .8076 17.15 16.16 .137 .154 .310 .264 .073 .070 Eoots .6525 .3262 22.95 .176 .515 4 .000712 .20 Tops 1.0114 16.77 .324 .116 .093 1.1515 1.1464 17.57 17.17 .311 .317 .119 .117 .094 .093 Eoots .4837 .2418 21.86 .258 .503 .057 5 .00142 .20 Tops .6648 17.40 .209 .069 .233 .7927 .7287 16.22 16.81 .185 .197 .103 .086 .209 .221 Eoots .4627 .2313 22.12 .258 .093 .058 .00356 .20 Tops 1.1639 16.50 .176 .153 .147 .9476 1.0557 16.10 16.30 .109 .142 .153 .128 .137 Eoots .3298 .1649 20.28 .433 .936 .066 7 .0058 .20 Tops .6608 17.08 .340 .218 .040 .6189 .7398 16.07 16.57 .400 .370 .234 .226 .040 Eoots .4846 .2423 20.09 .235 .788 .050 8 .0168 .20 Tops 1.0927 18.96 .344 1.020 .140 1.0824 1.0875 20.05 19.45 .391 .367 1.100 1.06 .120 .130 Eoots .6358 .3179 21.05 .177 .262 .051 Grown January 24-March 6, 1916. 1918] Waynick: Antagonism and Cell Permeability 197 1.2 - 1 .1 1 .0 o . y o . 8 0.7- .6 0.5 - .4 0.3- . 2 . 1 \ 7 ^ All y.\ \ & Fig. 15 Ferric Chloride + Magnesium Chloride (See Table 15) 198 University of California Publications in Agricultural Sciences [Vol.3 Table 16 Copper Chlori de No. Solution CuCl 2 Drv Weight 3 CD CS U 54_| eg 3 fcJO Ph CD 3 CO bfi #? ° 3 03 M S3 Pn 3 CD h£ "S pi Ph c6 CO 1 .000038 Tops .4611 25.60 .425 .194 .120 Boots .4692 .4651 23.60 24.60 .420 .412 .191 .192 .201 .160 2 .000079 Tops .2492 20.20 .455 .716 .155 Eoots .3242 .2867 24.12 22.16 .563 .509 .723 .719 .221 .188 3 .00015 Tops .3887 24.50 .312 1.10 .114 Eoots .4198 .4042 25.30 24.90 .361 .336 .90 1.00 .0261 .070 4 .00031 Tops Roots .2744 .1372 18.45 18.45 .150 .150 1.43 1.48 .040 .040 .001 5 .00047 Tops .2581 22.00 .176 1.35 .214 .002 Roots .2344 .2462 18.10 20.05 .102 .139 1.02 1.18 .094 .154 .002 .002 6 .00063 Tops .0785 19.10 .254 3.77 .003 Roots .0700 .0742 18.21 18.65 .425 .339 5.10 4.43 .248 .248 .005 .004 7 .00079 Tops Roots No growth. 8 .00198 Tops Roots No growth. 9 .00392 Tops Roots No growth. Full Nutrient Tops 1.1234 19.20 .311 .213 1.0268 1.0751 21.00 20.10 .241 .276 .199 .206 Roots .7210 18.99 .299 .216 Grown March 9-April 20, 1916. 1918] Waynick: Antagonism and Cell Permeability 199 1.4- 1 .3 1 .2 1 . 1 1.0- 0.9 - .8 0.7- / \ \ / / .6 0.5 - 0.4 - .3 .1 Fig. 16 Copper Chloride (See Table 16) 200 University of California Publications in Agricultural Sciences [Vol. 3 ueaj^ UB8J^ © © lO CI OC CO © CI "fH ©' o i— i ^0 J° to CI Oq T— 1 ^ © o © © oq to • O © OO © © © eSu^ueaaad 1 — I r— 1 CO © © 1^ to lO to b- -. to to © © ^ e> © © ""1 •"j M . © oq oq oq cq oq oq r-i oq oq to b- oq >^ b- i— 1 + ueapf CO b- 00 © CI © b- © © © OC ■ a l ►H 1 T-l rH oq CI rH H^ PQ a < En ^ rH Fh oq o lO © © © © © © © IC © *H O qsy jo tq o © # CO rH © # "*. CO oq to 00 L-- **. CI oq Ph s aSe;uaDjad; © oi t^ ©' 00 00* oq" © ©' rH c* oq' rH or rH <1 Q ci I—. i-H oq H 1—1 oq oq oq oq r— rH oq i— rH P5 m Ph co QO © t^ ci © CI >o O to o ueapi -h 00 © oq >C © >o © to oq O o b- © CO © >o HH OC' © r— to O -+ oq © # I— "*. >q r " 1 - -t © -f ©_ r^ CO t^ © © tO to c ,_! © >o ,_, © © © ■g CO ci © i— i © to © © 00 to HH CO to • O to "3 CO 00 to no b- oo to CO © b- © CO L— to oo ~+ HH to rH © © tq >o oq > HH H- rH rJJ cc rH i— I 02 GO co m CO an -t— 03 — OQ +* co co +-- Q Ch o P_, o Ph o Ph o Ph O O o o o o o O o O O EH W Eh « Eh Ph EH Ph Eh K b- co — 00 CD Oq 00 o Ol H< to Ol O o © © © oo C © CI © b- H- o CM rH CO r- © © auaj\[ © l~ X to CO — CO DC b- CI Cl b- CO o rH „ o CI 35 -f Cl Tf4 to l~ 1- CO b- CO b- GC rH ~ to © © © Cl to © © S H J" 35 CO 35 t- H- -1 re Cl >c CO o Cl -f 35 CD 35 to CC -f C5 © c r— I— I aSinueo-ied -t H- Cl h- X Cl IO ~. -h CO r JZ -t I- CO CO 1- 1- "°. to rH ©_ rH "t ~_ Cl 35 c T— < 5 35 Cl >c a Cl co to CO b- ©' r=: r-H ft Cl 1— 1 0Q r— «0J« rH to 35 CO -tf ir\ cc — X CXJ CM 1- Cl cc co -h X o © X Cl X 35 X ^ >o X H< Cl 1 — 1 ^ Cl 35 r-\ 71 r— { 1- cc 35 i — 35 35 35 C5 © © Cl X © co o aSu^ueojdti rH © «-J •"I p "i *-[ Cl -1 ""j CO CC H-_ to IO to 1- >C 31 b- X CO •-J ""I X h. O 03 N g + c >c co — © c © Cl 1 UB8J\[ 31 t- CO 1— j r-i b- to H- CM ep -t-s rH ci ci ci CO H-' -t* to © CM Cl Cl Cl Cl Cl Cl Cl CC rH rS" 1 O — C o o O i>_ o O o o t^ o to © © o © © © © o ~v © c3 "3 qsyjo CO CI X CO Cl Cl CO cc C5 CO Cl co cc 35 cc rH CO >C 35 Cl © CO Cl cr GQ e3im.i90.iati \ — 1 -+' —J CO ci C5 ci ci ^ - ci CO CO CO -H •* CO CO 35 CO -t © ex ci -+' CI Cl I— i Cl Cl Cl Cl Cl Cl Cl Cl Cl Cl Cl Cl Cl CM CC Cl = rS ft o ft CO X rH X b- r— j CO b- CO r— ( X 35 , — 1 Cl Cl © O o UB8J\[ >c Cl O to o rH cc X -h H" -h 35 X CD r- 1 — 1 Q -H -. CO CD Cl to cc H- © HH to" CC to 5D Cl to Cl to Cl -tl *"! to r-l "t •"l Cl © © CO t- b- CO CO CO ^H rH CM ^ to to C5 cc CC 35 X 35 b- CO HH -t © X '5 le- to to X rH ,— 1 -+ b- Cl -b Cl t^ to ■CC X 35 © 35 Cl I~ to co t- rH X Cl re ro b- CO o cc oc cc C5 cc Cl 35 C5 CC © © C5 to -H © to b- to CO -f to CD CC to "t -+' C| H- to Cl "t CO Cl Cl CO © re CC rH £ 02 »2 02 02 02 92 02 K DC +3 02 ■JJ QQ 43 02 +a X +3 02 ■+3 m +3 Vj +3 r* Ch o ft o p^ O r^ o P_l o ft O Z-, ft o A o o o o O o o o O o o o o o o o EH « H Ph H p^ :- X EH W r- P5 Eh Ph Eh X r CI X CO to CC 6 CO 1 - ,— i >o o o cc b- to © ji o o O o o © CO 3 © o o o © © © N o © o o o © © © .2 J" m © © o o © © © c_ ^ T* Cl Cl X to CM -v "+ Cl -t X 1 - -H ^ Cl o o o CO 35 X /. o - o o © © © , — 1 o c D o © © © © C o o o o © © © © © o o o o © # © © 6 rt CM CO •* to © b- X K 191 S] Waynich: Antagonism and Cell Permeability 20^ 1 .4 1 .3 1 . 2 1 .1 1.0- .9 .8 .6 . 5 .4 .3 . 2 . 1 Fig. 20 Copper Sulphate + Zinc Sulphate (See Table 20) 208 University of California Publications in Agricultural Sciences [Vol. 3 UB9J\[ CM CM CO O o CO o o o *M3P HNH H CM CM t- Id OOO OOO OO OOO OOO OO QTTr> OOOO ^!DO0 NH(M ©CO© NHH COhO IQIOS HNM HCOI> l>W i— I y il i° CO (M C5 LQ LO © lO ^ rf lO © «D CO i* W CO ^ a (M (M H a ^ t^ C H M CO N OS eSBJuaoaed OOOOOOOOOOOOOOT-HOOrHOOrHrHrHOr-irrCOCOCO o UU0J\[ co co o CM 00 rH OO CO CM CO 1—1 CO "cH CO a CO LO CO H ffi ^ OO 0] W O H I.O ■* 03 S O ICOlMNClNlCSOOOHS CO •°H J° co l^ oo i— co co co b- o lo co r« t-h io co co : i— i o -* oo o b- to i— i r« os >o GO 00 eSu^ueOjafJ rH rH CM rH rH CM rH i-J CM ,-h t-H CM t-h CM CM CO # I rH tJH Tti CM CO CO CM O CM CM CM r-i rH r-i r-i CM CO CM o o CM CM =3 7J u CM ph W CD rH + H CO OS UB9H rH v^v ln CM b- b- HHC3 b- rJH CO LO b- CO O0 b- CO ■* (M O CO 00 b- b- CO b- e J t° H N IO h © ^ H O O M (M CO I- h LO G r- N O C H © © L- eau^uaoaej ^ r^ r-i rH t— I rH r-l r-l i-l rH r-l rH rH rH CM HtMIO CM rH CO N CO H co^c. aco o CM T-l O) LO CM O r-l r-i t-t CM CM rH ueaj\[ 2 .a CM O CM O O CM LO r-l OO rn qsy JO CM <* b- LO CO rH © CO CO rr, aSBluaoaafT CO t-h' t}H CM' O" OO* t-H rH* o* cm' o* co" -J m +a GO •H OQ -^ M VI m ■M CO +3 50 -1— w J2 oo +3 CO ^J '<-> Oh O Ph o Ph o Oh % Ph o p o p_l o Ph o Ph o _ o Q O O O o o o C O o o o o o o c o o o o EH M Eh pp Eh M Eh r^ b^ w Eh H= H ^ H pq En rP b^ w H IO o »o co ^r co o -* OO o to b- 72 o a o £S3 O c o © i — i CM I- o rH oo o XII o o o o o o CM o o o o o o o o o o o o o o o o o o o o o ©_ © © o o o o o o o 1- -H CM Ci 00 rH •V -h CS TjH oo ^ CM os OS o O o rH rH CO CS T* oo oo o /. o o o o o o t-H IO o o o o o o o o o o o c o o o o o o o o o o © °. o o o o o o o o 6 ,_, CI CO rH IO CO \ l^ CO o o fc r-l L918] Waynick: Antagonism and Cell Permeability 209 1.4- 1 . 3 Fig. 21 Copper Sulphate + Ferric Sulphate (See Table 21) 210 University of California Publications in Agricultural Sciences [Vol.3 Table 22 Ferric Sulphate No. Solution Pe 2 (S0 4 ) 3 Dry Weight a a 3 3 3 « c 1 .0000014 Tops 2.2399 17.09 .107 .098 .246 1.3062 1.7730 17.55 17.32 .099 .103 .113 .105 .281 .263 Roots 1.0626 21.81 .122 .763 .315 .0000028 Tops .5313 20.50 21.81 .119 .122 .070 .763 .264 .315 2 1.9198 1.5226 1.7210 20.21 21.35 .115 .117 .072 .071 .201 .231 Roots .8731 25.50 .013 .145 .220 .4850 .6769 26.50 25.85 .020 .016 .158 .150 .286 .253 3 .0000070 Tops 1.4150 24.00 .095 .393 1.5247 1.4698 22.62 23.31 .080 .087 .102 .102 .395 .394 Roots .6984 31.10 .032 .263 .223 .7624 .7304 30.61 30.85 .041 .036 .224 .243 .277 .250 4 .000014 Tops 2.3544 20.57 .032 .089 .358 2.3461 2.3502 18.90 19.73 .041 .036 .106 .097 .378 .368 Roots 1.0361 27.20 .022 .184 .172 1.1861 1.1111 29.20 28.31 .011 .016 .250 .217 .178 .175 5 .00007 Tops 4.2000 14.80 .018 .072 .236 3.6033 3.9016 15.15 14.97 .017 .017 .073 .072 .268 .252 Roots .9813 25.81 .056 .134 .103 .9168 .9490 26.00 25.90 .077 .067 .118 .128 .129 .116 Full Nutrient Tops 1.5682 18.33 .293 .231 1.5775 1.5728 19.40 18.86 .312 .302 .279 .255 Roots .9776 20.40 .271 .279 Grown April 22-June 3, 1916. 1918] Waynick: Antagonism and Cell Permeability 211 1 . 4 1 .3 1 .2 1 .1 1.0 - .9 . 0.7- .6 .5 .4 .3 .2 . 1 \ Fig. 22 Ferric Sulphate (See Table 22) 212 University of California Publications in Agricultural Sciences [Vol.3 Table 23 Zinc Sulphate No. Solution ZnS0 4 Dry Weight 3 M 1 « K ° 3 bfl a h 1 .00000767 Tops .6718 23.00 .324 .7541 .7129 24.04 23.52 .341 Eoots .6138 .3069 17.88 .191 2 .0000131 Tops .7164 25.30 .326 .8350 .7757 24.65 25.07 .284 Roots .6009 .3004 17.20 .187 3 .0000395 Tons .6142 21.19 .372 .8607 .7374 22.50 21.84 .316 Roots .5864 .2932 15.90 .166 4 .000153 Tops .5109 21.90 .372 .3800 .4454 19.40 20.65 .437 Roots .4628 .2314 20.50 .128 5 .000395 Tops .8208 Lost .4104 20.38 20.38 .200 Roots .4432 .2216 22.30 .053 Ful 1 Nutrient Tops 1.0992 20.17 .310 1.0750 1.0872 19.12 18.69 .297 Roots .8120 20.00 .271 .332 .305 .344 .404 .200 .303 C 60 .374 .560 .810 .268 .228 .233 .46^ .0622 .0587 .310 .121 .386 .348 .101 .358 .101 ,324 .0708 .320 .322 .0644 .886 .660 .097 .760 .710 .981 .089 .423 .074 321 .073 .321 .073 .01' .224 Grown January 24-March 6, 1916. 1918] Waynick: Antagonism and Cell Permeability 213 1 .0 .9 . 8 0.7- 0.6- .5 0.4- /\ .3 .2 0.1- \ \ Fig. 23 Zinc Sulphate (See Table 23) 214 University of California Publications in Agricultural Sciences [Vol. 3 Table 24 Zinc Sulphate + Ferric Sulphate Solution Xo. ZnS0 4 Fe 2 (SOJ 3 1 .0000019 .0000035 Dry Weight Tops 1.0594 1.5294 Eoots .10444 2 .0000038 .000005 Tops 1.5364 1.4864 Boots 1.0716 3 .0000057 .000007 Tops 1.2300 1.6150 Eoots 1.1250 1.2944 .5222 18.50 18.70 18.60 29.80 18.60 1.5114 16.10 1' .5258 31.00 6 .0000379 .000070 7 .000076 .00014 Tops 1.6164 1.7228 Eoots 1.7611 8 .000152 .00028 Tops .9850 18.35 1.0268 1.0029 20.00 Eoots .6987 .3493 27.30 .35 18.10 1.4225 16.90 17.00 .5625 32.30 4 .0000076 .000014 Tops 2.3088 16.40 2.5100 2.4094 17.00 16.70 Eoots 1.5623 .7812 28.00 5 .0000152 .000028 Tops 2.3429 15.70 2.0129 2.1774 15.70 15.70 Eoots 1.5133 .7566 30.30 Tops 2.0533 18.10 1.5544 1.8038 16.20 17.15 Eoots 1.4194 .7097 25.30 18.40 1.6696 19.70 19.05 .8801 27.00 19.17 On ~ .096 .098 .045 .149 .170 .042 .172 .124 .068 .083 .098 .040 .154 .147 .050 .160 .160 .025 .157 .144 .019 .179 .176 .006 .097 159 .140 .090 .150 .160 .150 .177 O) Ph 3 .215 ,230 .149 .222 .031 .029 .037 .030 .262 .231 .184 .247 .036 .030 .082 .033 .288 .277 .130 .282 .076 .064 .010 .070 .110 .111 ,139 .110 .065 .062 .061 .063 .151 .158 .103 .154 .052 .046 .069 .049 .122 .151 .243 .136 .042 .049 .141 .045 .201 .317 .0412 .0412 .128 .043 .219 .282 ,367 .250 .045 .042 .043 Grown April 24-June 6, 1916. 1918] Waynick: Antagonism and Cell Permeability 215 .5 .4 0.3- - - - -^K X V / v / \ / \ \ o .1 ^:\>< Fig. 24 Zinc Sulphate + Ferric Sulphate (See Table 24) 216 University of California Publications in Agricultural Sciences [Vol. 3 Table 25 Mercuric Chloride 4- Ferric Sulphate Solution A SO Pm No. HgCl 2 Fe 2 (S0 4 ) 3 Dry Weight ^ Pn ° g 1 .0000047 .0000035 Tops .2039 22.40 .233 .2868 .2553 16.70 19.55 .179 .206 Roots .0518 .0259 18.70 .840 2 .0000094 .0000070 Tops Roots 3 .0000189 .00005 Tops Roots 4 .000047 .00014 Tops Roots 5 .000094 .0007 Tops Roots 6 .000189 .00105 Tops Roots 7 .000378 .00210 Tops Roots .2569 .3579 .0869 .2042 .2996 .0896 .2362 .2300 .0596 .2396 Lost .0237 .2288 .2988 .0784 .2184 .4272 .0496 21.75 .3074 21.90 21.82 .0434 18.90 16.70 .2519 19.40 18.05 .0448 18.13 17.36 17.32 .2331 17.41 .0298 19.01 16.10 .2396 .0118 15.50 16.70 .2638 17.20 16.95 .0392 21.30 .123 .138 .230 .222 .172 .231 .201 .271 .192 .183 .187 .221 .420 16.10 420 .151 .201 .131 .166 15.61 .3228 17.40 .0248 18.10 16.50 .190 .102 .232 .146 ££ g Ph S .723 .513 .618 2.670 .327 .515 2.95 .421 .371 .412 .391 .312 .416 .511 2.110 .100 .100 4.580 .731 .807 .769 2.170 .895 .550 .738 .463 ft .722 o a o .101 .157 .222 .129 Grown March 22-May 3, 1916. 1918] Waynick: Antagonism and Cell Permeability 21' 1 . .9 0.8 - 0.6- . 5 . 4 . 3 . 2 0.1- Fig. 25 Mercuric Chloride + Ferric Sulphate (See Table 25) 218 University of California Publications in Agricultural Sciences [Vol.3 Table 26 Mercuric Chloride O ( hi) i Solution .as S*S a £*£ c3 Xo. HgCl 2 Dry Weight ^ Ph ~ S P^ 1 .0000135 Tops .4904 17.20 .141 .4967 .4935 16.71 16.95 .098 .120 Eoots .1493 .0746 19.31 .478 2 .000066 Tops .2200 15.70 .332 .2236 .2218 14.24 14.97 .380 .356 Eoots .0239 .0119 10.50 4.31 3 .000135 Tops .1514 9.12 .421 .1421 .1467 8.95 9.03 .399 .410 Roots c fan a 3 .91 .248 .91 .248 .893 .296 1.69 .010 1.77 1.73 .017 .013 .775 .013 1.23 .012 1.33 1.28 .013 .012 Grown March 14-April 24, 1916. 1918] WaynicJc: Antagonism and Cell Permeability 219 1 .4- 1 .3 1 . 2 1 .1 1 . .9 . 6 0.5- 0.4- .3 .2 .1 / ./ \ I I s Fig. 26 Mercuric Chloride (See Table 26) EXPLANATION OF PLATES PLATE 13 Appearance of plants as mounted in corks at expiration of the six weeks' growing period. [220] UNIV. CALIF. PUBL. AGR. SCI. VOL. 3 [WAYNICK] PLATE 13 1 PLATE 14 No. 1. .24 M. MgCl 2 .004 M. CaCl 2 No. 2. .24 M. MgCl 2 .01 M. CaCL No. 3. .24 M. MgCl 2 .02 M. CaCl 2 No. 4. .24 M. MgCl 2 .04 M. CaCl 2 No. 5. .24 M. MgCl 2 .06 M. CaCl 2 No. 6. .24 M. MgCL .08 M. CaCl 2 No. 7. .24 M. MgCL .10 M. CaCl 2 No. 8. .24 M. MgCl 2 .12 M. CaCL No. 9. .24 M. MgCl 2 \l6 M. CaCL No. 10. .24 M. MgCL .20 M. CaCL No. 11. .24 M. MgCl 2 .24 M. CaCl 2 No. 12. .24 M. MgCL .30 M. CaCL No. 13. .24 M. Control. | 222 UNIV. CALIF. PUBL. AGR. SCI. VOL. 3 [WAYNICK] PLATE 14 ; \ w - . J ■ill 1 PLATE 15 No. 1. .30 M. MgCl 2 .004 M. CaCL No. 2. .30 M. MgCl 2 .01 M. CaCl 2 No. 3. .30 M. MgCl 2 .02 M. CaCl 2 No. 4. .30 M. MgCl 2 .04 M. CaCl 2 No. 5. .30 M. MgCl 2 .06 M. CaCl 2 No. 6. .30 M. MgCl 2 .08 M. CaCl 2 No. 7. .30 M. MgCl 2 .10 M. CaCl 2 No. 8. .30 M. MgCl 2 .12 M. CaCl 2 No. 9. .30 M. MgCl 2 .16 M. CaCl 2 No. 10. .30 M. MgCl 2 .20 M. CaCl 2 No. 11. .30 M. MgCl 2 .24 M. CaCl 2 No. 12. .30 M. MgCl 2 .30 M. CaCl 2 Control. 224] UNIV. CALIF. PUBL. AGR. SCI. VOL. 3 [WAYNICKJ PLATE 15 V 4 PLATE 1( No. 1. .04 M. KC1 No. 2. .06 M. KC1 No. 3. .08 M. KC1 No. 4. .10 M. KC1 No. 5. .12 M. KC1 No. 6. .14 M. KC1 No. 7. .16 M. KC1 [ 220 | s ; < - i. fW PLATE 17 No. 1. .00331 M. A1C1, No. 2. .000331 M. A1C1 3 No. 3. .000132 M. A1C1, No. 4. .000066 M. Aid, No. 5. .000033 M. AlClj No. 6. .0000165 M. A1C1 3 No. 7. .0000033 M. Control. AICI3 228 I -J PLATE 18 No. 1. .0168 M. FeCl 3 .20 M. MgCl 2 No. 2. .0058 M. FeCl 3 .20 M. MgCl 2 No. 3. .00356 M. FeCl 3 .20 M. MgCL No. 4. .00142 M. FeCl 3 .20 M. MgCL No. 5. .000712 M. FeCl 3 .20 M. MgCl 2 No. 6. .000352 M. FeCl 3 .20 M. MgCL No. 7. .000168 M. FeCl 3 .20 M. MgCL No. 8. .000089 M. FeCl 3 .20 M. MgCL Control. I'M) PLATE 19 No. 1. .00331 M. A1C1 3 .20 M. CaCL No. 2. .000331 M. A1C1 3 .20 M. CaCL No. 3. .000132 M. A1C1 3 .20 M. CaCL No. 4. .0000662 M. A1C1 3 .20 M. CaCL No. 5. .0000331 M. A1C1 3 .20 M. CaCl 2 No. 6. .0000165 M. A1C1 3 .20 M. CaCl 2 No. 7. .0000033 M. A1C1 3 .20 M. CaCl 2 Control. [232] PLATE 20 No. 1. .000094 M. CuCl 2 .00082 M. FeCl 3 No. 2. .000067 M. CuCl 2 .000058 M. FeCl 3 No. 3. .000047 M. CuCl 2 .000042 M. FeCl 3 No. 4. .000028 M. CuCL .000026 M. FeCl 3 No. 5. .0000094 M. CuCL .0000089 M. FeCl 3 Control. [234] ^ PLATE 21 No. 1. .000378 M. CuS0 4 No. 2. .000189 M. CuS0 4 No. 3. .0000945 M. CuS0 4 No. 4. .0000755 M. CuS0 4 No. 5. .0000567 M. CuS0 4 No. 6. .0000378 M. CuS0 4 No. 7. .0000188 M. CuS0 4 No. 8. .0000094 M. CuS0 4 No. 9. .0000048 M. Control. CuS0 4 [236] PLATE 22 No. 1. .0000047 M. CuSo 4 .0000035 M. Fe 2 (S0 4 ) 3 No. 2. .0000094 M. CuSo 4 .0000070 M. Fe 2 (S0 4 ) 8 No. 3. .0000142 M. CuSo 4 .0000105 M. F'e 2 (S0 4 ) 3 No. 4. .0000189 M. CuSo 4 .000014 M. Fe 2 (S0 4 ) 3 No. 5. .0000378 M. CuSo 4 .000028 M. Fe 2 (S0 4 ) 3 No. 6. .000094 M. CuSo 4 .000070 M. Fe 2 (S0 4 ) 3 No. 7. .000142 M. CuSo 4 .000105 M. Fe 2 (S0 4 ) 3 No. 8. .000189 M. CuSo 4 .00014 M. Fe 2 (S0 4 ) 3 No. 9. .00058 M. CuSo 4 .00028 M. Fe 2 (S0 4 ) 3 Control. [238] PLATE 23 No. 1. .0000014 M. Fe 2 (S0 4 ) 3 No. 2. .0000028 M. Fe 2 (S0 4 ) 3 No. 3. .0000070 M. Fe 2 (S0 4 ) 3 No. 4. .000014 M. Fe 2 (S0 4 ) 3 No. 5. .00007 M. Fe 2 Control. (S0 4 ) 3 [240] PLATE 24 STo. 1. STo. 2. STo. 3. .000135 M. .000066 M. .0000135 M. Control. HgCl 2 HgCl 2 HgCL [242] UNIV. CALIF. PUBL. AGR. SCI. VOL. 3 [WAYNICK] PLATE 24 i / ///