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F..¢. \ .. y ‘ . . ... .v 6 . 4 . nrr . u . . . .. -w . J“ . h 1 . . k1” . I . ‘ O ‘ a. . .. .. . . . . . .. . 1. t ..L.. a. LP......_..V.;..“. .¢ .. T. f 1......13322 ..1au.—~z:*:=.ua£~x . _ I Ir r 7... C \ f V 3 ' I ’ V A '1 ' A." Q n, r .. . . x 9.. a..... I s .v . ’s .5..th 5 431g .. . a bra!) Q... . . 3 1293 0080 a“ m sum IIIIIII‘I‘III‘ m m W W m | i n; . a U . .. .f ,.. ,u ‘ \ 8 "t y7‘3‘x: .s‘b PLACE IN RETURN BOX to remove this chpckout from your record. TO AVOID FINES retum on or beta! due duo. DATE DUE DATE DUE DATE DUE MSU Is An Afirmdlve Action/Equal Opportunity Inquuion swans-M SOME QUAETITATIVE.AKD QUALITATIVV STUDIES N r ”JET HORIOITE‘S BY TAIICHI ASAL Submitted in partial fulfilment of the requirements for the degree of Master of Science in the Graduate School, Hichigan State College, Department of Botany June, 1939 ACKNOWLEGEMENTS The writer wishes to acknowlege his indebted- ness to Dr. R. P. Hibbard for his guidance during .the course of the work and for his assistance in the preparation of the thesis; to Dr. E. A. Bessey for his suggestions; to Dr. E. H. Newcomer for his encouragement and valuable suggestions to Dr. W. D. Baton for criticising statistical treatment; and to Dr. G. A. Heppert for his information about animal-hormones. 121503 1. ll. 111. IV. V1. Vll. Vlll. fir LL. m: r- V-‘ ,—1 'A‘l \y‘v- l..-.L_. Or a . . Introduction. Historical review. Katerials and met'ods. EXperimental Exp. 1. The effects of hormone on seedling growth on filter paper. Ex.. 11. The effects of hormone on seedling growth in flats and pots in the greenhouse. Exp. 111. The effects of h rmones in the form of dust on the seed upon the subsequent growth of plants in the field. Exp. 1V. The effect of hormone on plants when applied in the fonn of lan lin paste. Exp. V. The effect of hormone on plant cuttings. Exp. Vl. Ho mone application on colchicine treated corn seedlings. Exp. Vll. Demonstration of the effect of light and darkness on the Exp. Vlll. Discussion. Summary. Literature grotth of seedlings grown from hormone treated seeds. Effect of hornore on catalytic action of ferric salt for the deconposition of H902 cited. Index of tables. Index Index of graphs. of plates. SOHE OUATTITATIYE 2RD CUALITAIITE STUDIES ON PLANT HOIKOTES V - KERODUCTIQE Growth is a mysterious phenomenon of nature. A crystal grows, increasing its size quantitatively. The root and shoot of a germinated corn seed grow at the expense of the plant's own stored energy, while at itmsmaturation phase the milky substances are condensed with a loss of volume and thus energy is stored in the form.of food in the seeds. Then a mosquito bites our hand a swelling occurs, but no zoologist would designate this as growth, but many workers with plant hormones consider hat the swelling of the plant body ceased by biting insects or that due to any other causes, is the result of growth associated with hormones. hany investigators are trying to demonstrate that the complex phenomenon of growth is theresult of the presence of some one or more hormones, even though there has been offered no definite nor satisfactory proof of their existence up to date. It is of course a difficult matter to determine whether or not all the phenomena related to growxh can be attributed to the presence of hormones since growth is one manifestation of life itself. Voluminous reports ujon "plant hormones" aVe been published in recent years. There are two apparent reasons why so many workers are entering this field. jfiany complicated growth reactions such as trepisms, curvatures, abnormal growths, correlations, metaxenia, etc., may be explained by postulating the presence of hormone-like substances. Since hormones or chemical m ssengers have been demonstrated in the animal world, it would be desirable, if possible, to extend the idea to the plant world. The term "hormone" in the vegetable world is used very ambiguously. The word was first suggested in connection with plants by Fitting (25) who found that a substance present in orchid pollen caused swelling of the " as gynostemium in the orchid flower. Starling (54) defined "hormone "any substance normally produced in the cells of some part of the body and carried to distant parts which it effects for the good of the body as a whole." Eoogl (50) preposed the inclusive term "auxin" for growth sub- stances that bring about cell enlargement. In comparatively recent years a number of terms not well defined have been referred to by various workers as growth hormones, phytohormones, auxins, plant hormones, etc. However, Boysen-Jensen (ll) wrote that to determine the hormonal function is more i portant than to define hormone itself. In this paper the term hormnne is used to include (a) laboratory synthesized chemicals which stimulate growth or cause bending of the coleOptile and, (b) chemically unknown substances extracted from plants which stimulate growth and produce curvatures. The present paper is chiefly confined to the studies of the first type, for example, indole butyric acid and naphthalene acetic acid. A limited number of experiments have been performed both in the greenhouse and the field to determine the practical value of using these hemicals in the form.of dusts on various seed and plant parts. a ’1': .b. EDI: I)...“ .e r). . 2.4! F it... N {maul ,i’yr .. Nia.«..mu but s 3.1:... .. ”Digit.”fi.gll u. .13..»enueE! A... .33.: hi“... .uFf...ll§..:l-Ii it .1 «.7 o. HISTORICAL REVIEW Fbr the sake of clarity an effort has been made to arrange this review according to functional evidence and only those papers considered pertinent to the present problem will be discussed. Plants resyond to certain stimuli such as chemical, thermal, mechanical, gravity, light, etc. Among these the reaction to light, known as phototropism, has stood out as a well known example since the time of Darwin. Darwin (21) car- ried on experiments with the coleoptile of Phalaris canariensis and concluded that the phototrOpic stimulus must be transmitted from the tip toward the base. Even today his demonstration of phototropism appears in various text books of general botany. Recently, Boysen-Jensen (ll) demonstrated that the transmission of a stimulus from the ligated side to the shaded one in the Avena coleoptile could be stOpped by an incision. In 1938, Went (63) reported that an agar diffusible substance from other plants caused bending of the Avena coleOptile when unilaterally applied. he also demonstrated the existence of a quantitative relation- ship between the concentration of the substance and the degree of bending. If the function of a hormone is to cause curvature in certain parts of a plant body, then Went must be given the credit of discovering "phyto- hormones." Rothert (47) reported that the removal of the coleOptile tip reduced rowth in the stump. After 10 to 14 hours growth began :2 L.) OD gain and the suggestion was made that this might be due to regeneration of the tip. Theorizing from this observation Boysen-Jensen (11) con- cluded that a substance is dispersed from the tip which promotes growth in the basal region. The writer believes, however, that it is probable that the teaporary cessation of growth was due to the direct effect of injury by daca;itatic . Weij (62) showed that if two agar blocks containing a growth-pro- moting substance were placed on eitherend of a cut coleoptile cylinder 2 mm. long, a decrease in growth substance took place in the upper block, but no increase was demonstrated in the lower block. "The most likely explanation of this and numerious similar obser ations is that it is con- sumed in growth," wrote Boysen-Jensen (ll). The present writer believes it is just as probablethat the hormone can be, so to speak, consumed through chemical reaction. In 1934 Want offered the "Pea test method" for measuring the con- centration of growth hormone in the solution form, and announced that it was just as reliable as the Avena test. Du Buy (15) reported that growth in the coleoptile is gradually retarded when the endosperm is removed. Some hormone workers (11) emphasized the importance of the supply of growth su stance from the endosperm, and when this was absent, through being cut out, growth was retarded. The author suggests that the retarded growth may be due to a direct effect of injury rather than the removal of any hormone. Although Avena curvature occurs with the use of many synthetic chemicals and agar diffusible substances from living plant tissues, the response is, how ver, not always of the same nature nor always constant, and therefore "the use of Avena curvature values is not a reliable cri- terion either for the action or for the transgort of growth substance in the tissue of other plants," wrote Skoog (50) recently‘lQBB). Very little is known of the chemical nature of hormones found in plants. At the present time the compounds which are obtained from plants and which stimulate Avena curvature are c. y three in number, Auxin "a", Auxin "b", and heteroauxin. All of them were found first b? Koogl and his co-workers. Auxin "a" (Auxentriolic acid) was isolated in 1933, from urine (51). It is characterized as follows: molecular formula clanégog, molecular weight 258, c ystals hexagonal, and sta la in acid. The name A min "a" vas given after Auxin "e" was found. Auxin "b" (Aflfienolonic acid) was isolated from.maize germ oil (52), with a molecular formula 18H3004, molecular weight 310, melting point 185, and is unstable to both acid and a kali, and easily decomposed by peroxides. Both Auxin "a" C and "b" have recently been ex racted from higher plants; for example malt, maize, peanut, sunflower, mustard, and linseed oils. Hetroauxin (5-indole- acetic acid) was prepared from urine (55) and also later found in yeast. Its molecular formula is ClOHQ 02E. The presence of hormones in living tissues is at the present time chemically undetectable, and no quantitative or qualitative chemical tests are therefore available. For the determination of synthetic hormones, when diffused or injected into plants, Hitchcock and his co-worher (29) in 19e8 report d that the Avena test or other seedling tip test were not reliable but that the Winkler and Petersen colormetric method (indole group test) was quite suitable. Identification of indole-butyric acid by the spectrosc0pic method is most apglicable for such small quantities applied (59). A mechanism for the fornation of hormone-like substances has been offered by several workers. In 1952 Sakamura (48) reported that the for- mation of some growth substances was accelerated by temperature, and also by feeding amino acid in the c'se where lower plants were used. In higher plants, however, very little is known of how the hormones are produced or the conditions which facilitate their develOpment. In 1957 Avery (3) reported that hormone content was proportional to light intensity and also to concentration of 002. It is else that some substance which causes Avena curvature is increased, but whether the substance obtained by agar diffusion is a real hormone or merely some chemical by-product is not yet clear. In 1934 Donner (7) reported that mineral acids alone would cause curvature or would accelerate normal growth in.Avena coleoptiles under certain conditions. Hitchcock (29) recently concluded that "There appears to be no Specificity of action for plant hormones." In 1937, Bonner (8) carried out a very interesting experiment. Decapitated riot tips of the pea, growing in nutrient solution, were cut and transferred into new cultures to eliminate the original thiamine (Vitamin Bl) that remained in the tissue. After repeating this transfer several times cell development was nearly stOpped. Upon the application of Vitamin 31, growth again star- ted. This experiment was designed to give some idea of whether or not any other hormone was necessary for the growth of roots. However, Thimann and his co-worker (53) concluded, regarding this work, that there was no doubt that Vitamin Bl was a growth hormone, "because it stimulates root growth." If one is led to believe from this that Vitamin B1 is specific in its action on roots, it must be emphasized that up to date phytohormones have not been shown to be specific. Cell division by hormones has been reported by several workers. In 1935, Snow (52) reported that the application of Auxin and B-indole acetic acid (heteroauxin) to the upper ends of decapitated Helianthus seedlings caused growth in thickness through cambial division. But other workers have postulated the existence of other special hormones for -7- cell di ision ($9). The role ofehertain substance, bios, which "appa- rently does not belong in the same category with auxinsfi has been des— cribed by many investigators (43).- The suggestion that hormones may act upon cell division has not yet been satisfactorily demonstrated (11). fMost investigators agree ugon the inhibiting effect of hormones upon the growth of lateral buds.- In 1958, Albaum (1) reported that, when a l per cent indole ac\tic acid in lanolin paste was applied to the apical cut surface of certain prothallia it inhibited adventitiOus outgrowths, while removal of the lanolin paste by excision caused the resumption of growth. This experiment is very interesting, because the inhibition by indole acetic acid in lanolin paste could be removed by the excision of the part of the plant upon which it was applied. This hormone seems to retard budding without much apparent diffusion into the tissue. Thus even if diffusion occurs at all, it must be very small and mainly limited to the point of application./ One hormone worker reported by Boysen-Jensen (11) mentions the presence of a hormone, produced by the terminal bud that promoted its own growth but inhibited lateral buds. If this is so, then one may ask how is it possible that a single hor one can serve dual purposes? Hitchcock (28) has shown that lateral bud inhibition can be brought about by the use of indole acetic and indole propionic acids, as well as with ethylene and pronlene gases when applied to decapitated tobacco plants. Therefore bud inhibition is not due to a Specific hormone because the inhibition may be caused by many other chemicals. Zaize seedlings growing in darkness usually show significant elong- ation of the first internode. Inge (56) reported that heating to 51°C one hour stopped the elongation, and from that he co cluded gr wth inhi- 5 I 7 If in tion was due to the destruction of the hormone. Such conclusions, often found in the reports of hormone workers (11), are inadeo Luate and unjust- ified from t‘1e experimental evidence presented. Some apdarently do not take into consideration the fact that conditions such as vam ations in ten1perature or treat ent with x-rays have their direct effects upon pro- toplasm and must therefore be evaluated. In 1935, Thimann (56) observed inhibition of root elongation and at the same time the initiation of net roots by the a plication of the ana- 10 ozs corp01nds such as indole acetic acid, 3-1 ndene acetic and l-couma- ryl acetic acid. In 1935, Zinnennan and his coénorker (67, 68) reported that several ho mones, beta naphthalene acetic, S-aceto naphthalene acetic, indole butyric, phenol acetic, fluorene acetic, anthracene acetic acid, and alpha naphthalene acetonitrile, nd ced the formation of roots. Amen? them, sip? a nPWhtbilene acetic acid and indole butrric acid were the most effective root prolucing substances. Zimme mann (66) showed that ethylene and propylene were effective in causi ing curvature of ’.vena coleOptile. Carbon Inonoxide and ethylene also inruced roots on stems in numerous species of plants than a plied in lanolin paste. This further -"1orts the non-spec1ficity of hormones for root initiation. Several ivorkers have reported that r at rowth was also increased by decapitation. Cholodny (16) concluded the Yaize root tips produced a growth substance which inhibitel its develop eat and taat if the root tip ‘?as removed, increase in length of the stump t ok place. The root and shoot grow in symbiotic rela H10 slip on «J one benefits fron the other. The root is parasitic on the shoot for its carbohydrates and possibly 5 Vitamin El (53). On the other hand it might be hindered by the presence of the shoot. Roots are easily on tiva ted on special media and root trowth without shoots coili e stimulated under ideal Conditions. Whether shoot growth could be stimulated under ideal conditions is pro lematical but theoretically pos ible. Shoot growth without roots and root growth without shoot when the inhibiting effect of the other is removed may be stimulated although this may only be temporary. In 1 54, Laibach (57) reported that callus fornation appeared after the application of lanolin paste containing an ex ract obtain d from orchid pollinia or human urine. There are many reports concerning callus or tumor fe:mati ns and her one enthu31as ts have been trying to show that such ab.ormal growth phenonenon are connected with the presence of a horn ne, especially in the case of tumor forma tion of leiumizous plants. These abnormal growths, be we fer, are not due to a definite hormone for such growth may be obtained with the use of many kinds of chemicals (ll). Parthenocarpy induced by hormone application has been reported by several workers. In 193;, Gustafson (27) resorted that definite chemical sub- stances, which are not specific, caused the ova1ry of a flower to develon into the fruit Without fertilization. He thotught these suest shoes were closely related to the auxins. It was an; ested that if hormones affected metabolism in the plant body, they might affect the processes other than growth, such for exam le as reapiration. In 1968, Pratt (45) reported tin t a concentration of heteroauxi1 var; ing from O. 0000?? to 0. OEN maikedly accelerated t1* e res- Hiir tion of the Triti ~um embryo and strongly depressed growth. It seems th t heteroauxin is capable of markedly acceler ting some metabolic reactions of Aants but decreases others, th Mefore the cause 0: uld jist as well be related to some Ling other than ~r».th promoting substarces. Further, Boysen-Jensen and his co-wo:ker (10) studied the effect of it -lg- n 1veha coleoptile* and F). decapitation on the intensity of respiration finally concluded that there was no effect of grthh substances upon respiration. Thus higher 0 ncentration of hormones may cause chemical injury while loser concentration of hormones exhibits no effect on res- piration. There are also numerous reports on the effect of horhones in normal plant growth. In 1956, Loehring and his co-worke (59) published the only report, so Thimann and his coworker (59) say, that showed anr marked increase in the elonhation of the stem due to heteroauxin application. They demonstrated that intact stock seedlings could grow in aqueous solution of a growth substance (0.07fi) or in the soil, treat~d with certain growth substances. They also reported slight increase in top elongation of_nvena seedlings grown in solution culture with heteroaurin. In 1957, Grace (24) observed increases in growth of wheat and seVerml other young plants tr ated with hormone i. the solution or in the powder fo1.. Farmer (41) applied a wide range of concentrations of hormones. her results showed no stimulation in growth for whee: seedlings, but a decrease of primary roots and an increase in number of secondary roots. Thimann and his coworker (60) have recently report d that auxin treat- ment (indole acetic acid 0.01fl solution) hastened the develOpment of the photosynthetic area at a given time by 50-90;. The conditions under which this stimulating effect was obtained in the o'se of tomato plants, were not fully described, for in other series of the same experiment no effect upon growth was apparent. The variation might have been connected in some ‘ay with water relations. Finally, Hitchcock and his coworkers concluded that Tent's axiom, "Withou gromth substance, no growth," is -11- dif icult to prove 'ecause of the limitation of experimental methods. To quote furth r from these authors, the above axiom is essentially no different from that proposed by Avery, "Io nitrogen, no growth," and could be said of other im ortant substances such as P, K, Ca, Fe, etc. The effects of hormones upon normal plant growth have been reported with conflicting results, some positive and_others negative. Several workers have attempted to ex lain these variable results. Greenfield (26) prOposed the hypothesis that "auxin satiated plants" have already sufficient auxin, and therefore an additional sup 1y causes injury and consequently no growth. On the other han "auxin unsatiated plants" have not enough hormone, and consequently could take up an additional supply and increase growth. Wheat seedli gs were taken as an example of "auxin satiated plants," as their growth was not stimulated over a wide range of concentrations of several different growth substances. Cho- lodny (17) presented the hypothesis that growth substances promoted the rate of development of growing cells but shortened the length of their individual life cycles. In the case of the cells in the root these matured quickly without increasing in length, and therefore resulted in retardation of growth. In the stem, on the other hand, the zone of cell stretching was greater, and growth continued for a relatively longer time. However, there is insufficient evidence to support this hypothesis at the present time (11). Growth in plants, induced through the use of so-called plant hor- mones, has not indicated that any of them are specific, and therefore it naturally follows that the results are due to stimuli of chemical, electrical and mechanical, etc.,nature. In 1957, Leonian (58) concluded that heteroauxin was a growth inhibiting rather than a growth pro oting -le- in) substance. Ea interpreted many reports, including Avena coleOptile curvature, as plant responses to irritation which are equivalent to me— chanical injury or to parasitic invasion. Purdy (46) reported for Avena that the initial curvature_caused by wounding was positive and then was negative and for a second time became positive. aarotta (40) reported that Sprouts of Zea mays rrww readily in nutrient solution. If, howev r, the sprouts were slit (l mu.) new roots would then appear. Also on ad- dition of a Pb-salt to the nutrient solution, a retardation in gro th occurred for the first few days, but after ll days root growth was ac- celerated. Recently, in 1938, Thimann and his coworker (60) reported a very in- teresting study. They vernalized Avena and Triticgn seed by the cold treatment and compared their growth with hormone treated seedlings (0.01fifindole acetic abid in solution.) The result showed similarity of growth between the two treatments. Reviewing the literature and evalu- sting their results, they concluded that at first root inhibition was doubtless a general prOperty of all auxin activity, but that later, growth was accelerated. Avena plants from hormone treated seeds, these authors found, would flower from three days to one week earlier than the controls. The writers believed that the growth of a plant especially in the early stage is dependent on the amount of water availabie, and there- fore any increase in the amount of absorbing urface would naturally cause an increase in growth. In the case of Eriticum, auxin treatment at an early developmental stage produced marked reduction in the size of the first leaf. In general it was found that the gr ater this reduction in size the greater was the subsequent acceleration of growth in the older leaves. The authors tentatively suggested that the vegetative effedts of -15- vernalization were due to the prolonged e posure of the seed to its internal auxin supply. if . 71-71“ '1 a“ $.'T‘!r‘_.“"v."‘_"~ n NAZanIQLo AnJ ui'hUfio LJLL- Katerials used in these experiments are lis.ed in table 1. h‘ormoue dusts were made Ly mixing fineir grourl Lerrcie 13'sitl: fii: ”ale powder. fixing was facilitated by the use of a rotating machine run usually for a period of 20 hours at a time. Two concentrations of 3-indole-butyric acid (1 per cent and 20 per cent) were first made up and from these all the other concentrations were prepared (table 2). When very low concentrat- ions were desired, a relatively high concentration was selected and the weaker ones made by dilution with talc powder. Lanolin pastes were pre- pared by mixing finely powdered crystals with lanolin. .A ten percent paste was first Lade and from this a three percent and a 0.1 percent were then prepared. For practical purposes, it is important to know the Optimum.amount of dust adsorbed by seeds before using the seed dusting method. The amount adsorbed, as it is seen in table 3, is not constant, and therefore merely adding an excess amo.nt of dust and shaking this off would not give the exact amount taken up by the seeds. Bernburg (5) reported a method for determining the Optimum dust adsorbing power of 5 gram quantities of seeds. His method is not applicable for larger amounts of seeds, as for example a 100 gram sample. After several trials the following method was -14- adopted. Various am unts of talc were added to various 100 gram samples of seeds. These were then mixed in a bottle, shaken vigsrously by hand for five to ten minutes, and then transferred to a sieve and the excess powder removed with violent shaking for one minute. The results of this test are given in table 3, and the curves are shown in fig. 1. The prin- ci la is this; the Optinmn point of adsorption is assumed to be that point where the amount added (Y) equals the amount adsorbed. Thrre is therefore no excess (X), consequently the value of‘Y at X - O and this natirally is the Optimum adsorption point. In the case of buclwheat the Optinum.point was calculated by solving the normal equation, which is given in table 4, and run in; through the calculations which a.e found in the subsequent paragraph. All the reults for optimum quantities of dust adsorbed by 100 gram sample are given in table 5. From table 4 the following data are outained for buckwheat n = 5 z y = 11.57 2 x? - 51.48 x x 3 8.78 2 xy = 24.57. The predicted equation is xzaiby, where a and b are constants and x is excess dust remaining after mixing. The normal equations are n a + bziy : x ~ 2 82y+oiy :va. Substituting known values from the table, the normal equations are 5 a 4 11.57 b = 8.78 11.57 8 § 31.48 b : 24.57. Solving above equations alternately, the constants for the predicted equation are found, a : - 0.279 and b = 0.878. The theoretical equation must be x = - 0.268 4 0.878y. when dust is completely adsorbed there will be no excess amount remaining after mixing it with the seeds, that is, at this point x : 0, and the equa- tion becomes 0 = - 0.268 4 0.878y. From this, the Optimum amount adsorbed (y) is solved, y g 0.505 g./100g. seeds. In the case of corn, large petri dishes (l5 cm to 25 cm in diameter) covered with bell-jars, were used as moist chambers for germinating purposes. Filter paper was placed in the dishes on round glass plates adjusted to an angle of about 50°. This afforded more normal conditions for root growth. Wet cotton was placed around the glass plate to main- tain moisture conditions. Ten or 15 seeds were placed on the filter paper near the upper edge. The bottom of the dish was covered with tap water, about one centimeter deep. Soon after germination had occurred only five to ten uniformly germinated seeds were lined up at definite distances on the filter paper. The whole set, arranged in an orderly manner, was placed on the table under difdused light. At a later date it was found convenient to use a much larger moist chamber, which is shown in Plate 1. The inside of this glass box or Wardian case (llx70x90 cm.) was lined with cloth to retain moisture, and the roof part was so hinged that ad- justment could be nmmmafor temperat‘re. The case was set upright in a galvanized iron pen in which about 5 centimeter of water was maintained. r51- ' animos- .1 -15- 01 one end of the case there is a swinging door. Seeds were mounted on filter pa er resting on tilted glass plates placed in a snall pan (50 x 50 x 10 cm.) shown to the left of the Wardian case in Fig. 1. This small pan was kept in the moist chauber during an experiment. All measurements of whatev r nature were mdde in his chamber to prevent drying of young roots or root hairs. The case was placed in the botany greenhouse. Further details for other methods or modifications of the same will 1e given under each experiment. Waj-—q-q ‘ m~ v'v , .V l r -'I ‘4 L -AJA. ._J-~ -..7.._4_' A-- Gr ce (24) and others have secentaly advocated the use of hormone dust on seeds and have reported increases in gro th of roots and tops of certain species of plants. Hormone in the powder or dust form is said to be better than the liquid form especially on soil or sand cultures since the supply is available at greater dilution and over a longer peri- od. A study of the literature fails to reveal the most effective hormones or their best Oper ti g concentrati ns. It seems, however, that naph- thalene acetic acid and indole—B-butyric acid are two of the best sub- stances fOr root development (ll). In 1958, Hitchcock and his coworker (29) reported that species of the same genus did not respond alike to a given treatment, and the same might be said of different varieties of the same species. However, hey reported that in most cases the Optimum concentration of indole butyric acid in solution was 0.05 percent or -17- less, and this was eagecially true for root production. According to Grace (24) wheat seeds dusted with 2:l,OO0,000 of indole-3-acetic acid (2 parts of hon;one per million parts of seeds) increased root growth 65 pe-cent at the end of a 50 day period and that soy bean seeds treated with lO:l,OO0,000 naphthalene acetic acid yielded greatest tOp growth. In a preliminary experiment the grotth of barley and corn roots was studied carefully in petri dish cultures every day for several days, but no significant differences could be detected at low concentrations such as 0.05-4 p.p.m. (prrts per million parts seeds) when indole butyric acid was used. These studies showed the necessity of adhering to certain definite procedures in such experimentation. The seeds are very sensi- tive to variations in external conditions, such as moisture and tempera- ture, ani seemingly more so than to honnone treatments. When seeds are placed too far from or too close to the water level, genuination is retardel. In the case of corn there is a day gained in germination when the seed is placed with the embryo next to the wet filter paper rather than in the reverse position. It was found necessary to adopt some definite procedure for observing root growth. Equal time periods for observations, and photosraphic exposures, etc. were selected. The cultures were a r n'ed according to concentrations, or according to age, etc. By such means other workers might easily be able to repeat the experiments. Root heirs of young seedlings are particularly sensitive to moisture changes. A 20-minute: exposure to dry air is sufficient to ruin seedlings for experimental purposes. Seedlings which have a deli- cate root system such as wheat or barley are not very suitable for experi- ments on root growth under the conditions obtaining. For such reasons most of the experiments were conducted with corn which seemed relatively mere resistant to changes in.moisture conditions. . H- ‘*u..-. - " (‘- The relatively few seeds used in any one e perinent may seem insuf- ficient from which to draw conclisions, yet the nature of the experiment precludes the use of larger nimbers of corn seeds in any one experimeit . To increase the number of seeds, for example, would increase the time to make the necessary measurements and this in turn would produce greater variations in temperature, moisture and other conditions and would lengthen the time period for the study of each culture. Consequently - a ‘. T. -‘. ‘1 ..._ -.‘ ...‘r .n 4.: , 7-, ..;- : ., LAKJU dual-”J; U- u_ir.‘v..¢ __L. 1-4.3 Genoa H k n (u i" t g. S the experiments here the experiments with corn seedlings on filter paper were repeated twenty times or more. The use of any chemicals for disinfection was avoided for fear of secondary reactions arising. The ef”ect of these might be difficult to distinguish from those due to hormones. Seeds were carefully selected and brushed with a clean dry cloth. If any molds appeared in the cultures the cultures were discarded. All filter papers were sterilized in dry heat and all glassware and dishes first cleaned with formaldehyde and hen treated to dry sterilization. ...'.. “-47. T, .1- '5" r“ . . .— .—.‘-.—,.I—q—qlfi .1 /‘ -,_, - _. ._- -,’\I\“Tfl. O a, T n‘fion LI. n -. h'fip ~-- ~ -.,., q-.. -—~,-v- w.-. .1—x fl .5 7’3““? , '1 ,. . , I v . . . . . , it » V r ,4. J a ‘ ' ‘.. Q L- up) .9.) Late 1-.-? Lu M lL‘ . \ ‘11:.LJlL. . .A . - T . v _. . ‘LJJ-4‘L.‘.—-.4-I-'J .L. -ofllJ .J. L—JJJ— V‘— L-~ :Ll‘V“ J Before dustirc all seeds rere fanned cleaned and ~*olished in a 9 i *“o’ cloth sac. Samples of 100 grams of seeds were m'xed with dust in small 8 to 12 02. bottles and kept thereaft r in the same containers. Dusting hormone powder on the filter papers was accomplished by spreading the stock powder on as evenly as possibly with a spatula. fl The data for the grof th of Dent corn on filter paper trea ed with hormone are shown in table 6 and p ate 7 illustrate the condition of growth. It is seen from the table that germination iis sli2htly retar- ded at the higher concentration, for example 0.2 mg/paper, and als tEat root gre th was inhioite; significantly at a higher concentrati; than 0.02 mg/panar. theoretically, hormones increase gro th, conse- quently there should be a certain prOportionality between ;ro th increases and increasing concentrati but no such evidence wa: apia1er it. It would appear then that one might just as wellattrioute the slight varia- U} tions to or :ance and not to hormone treatnen s. Hormone tr ated seed do not show any - arhed tendexcy to: ards inc: arsed growth at low concent- ration, such as 0.05-2 p.p.m- (table 7). Re atively yo n er stag s of growth are presented in these cases, because with older seedlings second- ary factors w.uld enter in, such as a imitation of stored food in the endosperxgz, for e;-;ar:1ple. In table 8 can is found the data of another corn ernohiieit where in this case the re is definite evidence of growth acceleration in both tops and roots who re the con centratio: s of hormone were low. At high concetnrations growth as sr~ulfl0antl re arded. Fig. 2 shows the curves plotted from data in table 8. The graph shows that the inh it Jition in root growth is greater than that of tot growth. This was repeatedly observed in the different experiments. In table 9 will be found the data for seeds treat-d with hormone con centr tions varying from 20-200 parts of hormone to a million parts of see's. TOp growth is not much affected but root growth is significantly inhibited at higher concentration. At hiehor concentrations (2000 p.p.m.) as shown in table 10 both A‘ both tops and roots were siflnificontlyinhihited vhiLe at 200 pop-mo only the roots were inhiLited. It was repeatedly noticed that When root growth was inhibited n nerous secsn ary roots dexelOped from hear the seed. It was also observed that numerous root hairs dexelo ed on retarded primary roots. It seems that the root hair is more resistant to chemical injury of hormones. Kecsters (25) reported, "The root hairs are less sensitive to the growth substance." The production of dense root hairs may be accounted for through the fact that hormones are not believed to affect cell division but will modify cell elongations. Others affirm that hor- mones inhibit both cell division and elongation, and if this is so, root hair growth might then be stimulated. Without furth r anatomical study other suggestions would have little value. Various stages of de- velOpment in certain 8 eat corn seedlings can be seen in plates 5,4, and 5. Table ll shows the effect of ight upon the Sweet corn seedlings growing on hormone treated filter paper. Karked differenc.s can be seen in the length of the first internode. Darkness seems to favor elon- gation. Inhibitions of top and root growth thrsu;h hormone activity is less in darkness than in diffused light. These reactions could just as well be due to effects of darkness or to several other possible favorable or unfavorable co ditions. fiany observations have led the author to conclude that hormone treatment has not significantly modified the growth of the first internode, whether in darkness or in diffused light. The length of the coleOptile was constant regardless of hormone treatment or ight variation. It is snggeste‘ that retardation of germination while in darkness is due chiefly to slowering of the temperature. -31- The effect of hormone on the rowth of buckwheat is shown in table 12. Buckwheat roots are also sensitive to variations in moisture, etc., and if exposed for even 20 minutes to room temperature growth will have ceased at the end of twenty-fiur hours. Later experiments with buckwheat were conducted in the Iardian case described above. Buckwheat seedlings show considerable variability in their growth habits. It was very dif- ficult to obtain seedlings of uniform leneth and apparent equal vigor, as was possible to do with wheat or barley. The results of such experi- ments are plotted and shown in Fig. 3. At lo er concentrations the curve indicated stimulation but this was found not to ”e the general rule vhen tie experiment was repeated six tmmes. At the higher concentrations there was always a retardation of top and root growth in the younger plants. At higher concentration (300-5000:l,000,000) both roots and teps were retarded; the former much more so. At a later date growth retar- dation of both tops and roots was greatly diminished, and complete re- covery finally resulted. Plates 8 to 11 show the various stages in the develOpment of buck- wheat seedlings. It followed inevitably that at higher concentration when primary roots were inhibited there was a heavy proiuction of se- condary roots which develOped from the base of the stem. As compared with check plants, the total functional root mass was theSame or in some cases better. This can be seen on a study of plate 12. Exjeri- ment 1 is summarized in table 15. L_ -. 7'1 "‘Dfl‘f‘f“: T'T‘ '3 1- Tfjwj Tvmflufif‘ x TH (3T7! " T"? 0‘?! (37'1“? LI~ TC m '3 uvwwnv I“? f"? ‘1 rrt'fi D “ah. :41. L...‘ “:11... l1 I-L . A- _4 AJ-L ._ (JV .LJ J} HV|¢ I.OLI_J -1. gush-LI -. f ‘ r- UV . ‘LLl 4“: .L .LJ-‘\ ; D ;L on me 171 err-o 77*“ “U TTC‘T,‘ ‘b,-~3 AI i—l—J ‘O‘AJ—é—‘L; Inigo-LJ'. Plants were grown in pots or wooden flats and placed in concrete benches in the greenhouse. Watering was carefully done from below by running water into the bench until it reached a height of 5 to 10 cm. At the end of a few hours the water was drained off. Water was never applied on the toy of the soil. As a result of preliminary studies it was found wise to dilute the hormone dust with 100 to 200 :ram of dry soil and to distribute this as uniformly as possible over the surface of the soil. Seeds were then placed directly on this layer of soil and covered with approximately 2 cm. of sand. The soil used in these ex- periments was a Hichigan san y loam. More eXperimants were conducted with higher concentrations than with lower ones, since the latter had not been shown to have a stimulating effect. In one of many experiments with buckwheat it was observed that growth of seedlings was greatly inhibited at 20 mg/pot concentration, but that when the flower stage was reached the rate of growth was mark- edly increased. At maturation the height of the plants was slightly more than in the case of the controls. Some of the possible explanat- ions for this effect may be found by (a) assuming as some enthusiast would, that it is a direct result of hormone treatment; that (b) jaro- vization (10), is a likely cause; that (c) injury of a mechanical (45, 40) or chemical nature is operative; or that (d) it is a matter of chance. In view of the fact that stimulation effects are not usually indicated, this would point to the conclusion that natural chance might be the explanation. If jarovization is effeczive, then in the Earlier stages of growth iniibition should be evident, and in the later 3"‘1 In.“ ”- .)1‘" - “’0‘ stages accelerations of growth would obtain. In this buckwheat experi- ment it appears that jarovization might afford the explanation, but 0108;? study shows that this is not probable, since the most signifi- cant characteristic feature of this type of vegetative modification is the fact that it hastens maturation. In the case of these buckwheat seedlings, maturation was retarded. A further consideration of jaro- vization leads one to wonder whether suchtreatments as implied would be conducive to normal growth. It was orfinally claimed that yields of winter varieties were increased but in a number of recent tests this Les not For: Tutli: d. Stimulation of growth hrough mechanical, chemical, and electrical injury, is admittedly possib e. In some cases the mechanism appears to be one of enzymatic activity or oxidation while in other instances it seems impossible to learn the cause. Jarovization might also be clas- sed under theheading of injury since the nature of treatment is one similar to “echanical and chemical injury. It has often been observed in biological experimentation that unfavorable conditions of temperature and chemical proportions, etc.,do induce accelerated growth either directly or indirectly, providing the injurious factors are not very strong. Living organs or tissues some times show an increased resist- ance a ainst a poison and later not only recover their original activity but in some cases become more active. But these examples are scarce, and it is rarely possible to find in nature all the optimum conditions at one time in one place to bring about the desired effects, or even to produce them artificially (ll). If the unfavorable stimuli are not severe, one will first notice inhibited growth which will soon be replaced by accelerated growth. 'SVH 1 I :‘II Elf- Ioii.’fl.4n . . ah." Now if this is continually maintained to maturity then obviously we have a true case of accelerated growth. Howey r, conditions are rarely so c mbined that an increased growt rate can be continually maintained. Normal plants can sometimes be made into dwarfs by drastic changes in environmental conditions but this is not common under the general run of conditions for there are certain genetical limitations. To conduct further experiments, it is important to determine whether or not hormone is essential for normal growth. If hormones are necessary then Tent's (63) statement "Without auxin no growth," is corre t. On the other hand if hormones are not essential for normal growth then Leonian (58) has the right interpretation. Sweet corn seeds treated with hormone dust and grown in the green- house showed marknd inhibition of growth at the early stages out later normal growth was attained. This is shown in table 15 and plate 14. The results with buclvheat are shown in tables 16 and 17, and summa- rized in table 18. The experiment shows that buckwheat at earlier stages of develOpment was markedly inhibited by the hither concentration (100 mg per pot) while at a later date growth was only slightly checked. Low dry weight yields were obtained at high concentration. Plate 15 shows the appeareance of the plants at the beginning of the flowering stage. The conclusion drawn from this experiment is that buckwheat in the seedling stage is retarded in its growth approximately 40 per cent when compared with checks. The concentration of hormone dust in this case was 0.1 gram indole butyric acid per pot. After the flowering st ge and up to maturation there was a marked increase in growth. At maturation the height and vigor of the plant was like that of the check. Yet there was a decrease in dry weiéht, which decrease is believed t» be due to the decrease in growth during the seedling s age which in turn was due to the hormone treatment. Growth of the bean seedlings was markedly inhibited at 2000 p.p.m. of seed as shown in table 19. Germination was delayed two days behind that of the check in the field, while in the greenhouse it was delayed for seven days. It is suggested that the eXplanation of this lies in the mechanical removal of hormone in the field experiment. Grace (24) re- ported marked increase in growth of been plants at 10 p.p.n. naphthalene acetic acid. In this experiment at the same concentration as in Grace's eXperi ent only three percent increase in length and dry weight was obtained and this is not considered significant. This is within the li- mits of natural chance. In the case of high r concentration (200 p.p.m.) inhibition was marked and significant. Results for t mato seedlings treated with "Rootone" (naphthalene acetic acid) are found in table 20. The tomato plants were started from seeds planted in 8 cm. pots. When these were about 15 cm. high they were transplanted (October 4) into 25 cm. pots containing sandy loam. Root ne powder was added to the soil around the r 0.3. Two weeks after, marked inhibi-ion was observed in the Rootone treated seedlings. This inhibi- tion was observed in the Rootone treated seedlings. This inhibition consisted in a reduction of growth in height and number of branches. The photograph (plate 17) was taken one month after transplanting. About 6 weeks later, on December 20, the treated plant had not yet re- covered. . In table 21 can be found the data for growth of wheat and millet in wooden flats 50x35x10 cm. in the greenhouse. Rootone treated seeds were sown on the soil in the flats. Then the plants were about 2 cm. rows 35 cm. long and 10 cm. apart. The U) tallthey were thinned out to results at the end of a period of one month indicated that millet at higher concentrations was markedly retarded in its growth, while wheat showed an increase in dry weight in the lower concentration amounting to about seven per cent. The root system of the millet plants was f:und to be different from that of the control. The roots were increased in number but reduced in lenrth in higher concentration (60). In the field experiments with wheat it was noted that tille;ing was better in treated plants. In this experiment increased tillering was noticed and it was determined that the number of tillers is also correlated with the dry weight. At the end of a period of One month 40 plants from each grouj were classed according to the number of tillers. The great- est n mber of tillers was three, but this high number occurred in fewer plants. The larger number of plants had one or two tillers. The data re shown in table 22. The n‘mher of tillers are found in column headed (n) and the number of plants in each class under the column headed (N). The surmation of these values (2 n3) will give the total number of tillers 1 in each group. The value of CE n3) eipressad in percentages, eased on check is the relative value of total tillering for the group. This is given in the last column (table 22). It is seen that the number of tillers is correlated with dry weight. The dry weight figures are found in the lust column in table 21. This experinent shows hat increased dry weight through hormone reatment (0.33. Rootone per 1003. seeds) resulted because of the inc- -97- \J roses in the numb.r of tillers. In other words, hormone treatment inc- reased th dry weight of wheat seedlings through increasin: tille -‘ The writer fines no reference in the literature on this particular f atur . It is sugfiested that this increase of dry veight can be ex- d) (D plained in the following way. at first 3ro th is retarded due to horrone treatnent and then this in turn stimulated tiller formation. These results were obtained on one month old plants. If the plants had been selected at a much later date and the data collected, the above results might not have ‘een obtained. It is quite probable that then the dif- ference mi3ht not be so apparent. Cases are plentiful .:here the stiiu- lating effects of hormone treatments wear off, so to sneak, and the plants are no better than the checks. In the case wher inhiiitions have been reported later test w.:ul1 indica te recov:ry and both treated and check plants would yield alike. ‘V'W’ T. ‘V“'P:- v-“Y'-rf" I-unI @7' '~. ~ =f—fiwwf‘r‘ ‘ff‘t " TF“"{‘ ‘-.‘ f—j , -——~ - h.-. . fi._.__, ._ /, 71—fim "f wafi fl ‘% .' .. .. ' . . , . l ‘ . I. u . . . . ,J . I I) ,3 ‘3 .4... )LI.L.»;.-..v L ._ O 1...} .J H- ._'.'J Lu) b- A-ba. . ,U-‘1.4 -- .1.-. i ._u ... u ~L... ‘J .1? \ )J. -'-\' 4|... ._4 ‘J , .41) ‘ w - r r 1 p: 7 w- _ —-1. —" fl ’ aI-w-tv 9 AT, 131‘s}... m —n - m v r‘T q 1T ‘- ofui. .. . a - . .......L -‘t....‘..L-L V- .1 ...- .J..‘- .. .1. -uJ... c From his results in the greenhouse, Gr ace (2%) sugjested the use of dusted seeds for field ilantl ‘s. This led to our field experiments with dusted corn seeds. The seeds "ere planted by hand in rows 3 feet apart. The hills in each row r=re also 3 feet apart. Each row was about were 165 feet long. Bean, buckwheat, and .he at Aalso plan ted in the field. Data vere collects: in SLCh form as to be utilized for statistical treat- ment if necessar". 63" ’ .‘ - H‘- The data in table as shuw . at Dent cein was slightly retarded in germi ation and significantly inhioit d in its W‘ m'th in hci_1t at 200 parts of indole butyric acid per million parts of seed, but not affected at lower co- entratiens. The results for Sweet corn (table 24) treated in the same mztner show vary plainly that at earlier stages, of growth inhibition is signific nt at the hi her conce: tr ti n. Observaticns ‘ 8 weens later shore d that t e effects of inLioition had disappeared. Table 25 contains the data relat1‘ve to the gro Lh of bu ckcieat up to flo ering time. At the higher concentration (3,000 p.p.m.), inhibi- tion is apgarent. A ter this por riod and during maturation no dif between check and treated plants was noticeable. At concentrati ns 0.15 to E p.p.m. and at the tine the experi;ent was completed, there vas feind to be an increase in height amounting to an average of 5 percent and an increase in dry weight of about 4 percent. These differences in favor of hormo:.e t eatnent a1re of no practical si nific.nce. Fisher's "analysis of variance" was applied to he data for hormone treated corn as found in table 26. From the data in tahle 26 the follow- ing table of am.al sis of variance has been prep ca.red. AnClysis of Variance for Testi13 Gr wth in Length of Corn. Source: of Variar ce Degrees of Sun of Vari- Siandard freedom squares ance de iation T0 tal 93 02le .o -- -— Between means of treat. 4 1606.8 401,450 .. Betm een lines. 19 lOl4.0 56.568 -- Error 76 1597.2 47.352 6.89 ’1 ta do The. The standard egror, S, of the difference betveen any pair of means is l s a 6.89 (2/20):: .-. 2.14021. variability between treatments: the value of F in Fisher's table, cor- responding to the degrees of freedom.for 4 and 76 is, F = 5.56, while that for the sample is F a 401.45/47.55 = 8.47. The value, 8.47 demonstrates hat variability between treatments is highly significant. To determine the variability between line means or location means F. the following procedure is given. From isher's table, F = 1.67, and F sanple = 63.368/47.53 = 1.12. This shows that variability between line means or location neans is not significant, since the calculated value F is smaller than 1.67. The test for significance between two means is also given. For 76 degrees of freedom the values of the 5% and 1% levels are reagectively t a 1.99 and 2.65. Hence a difference between treatment as large as 2.17 x 1.99 a 4.32 is significant, and a difference as large as 2.17 x 2.65 = 5.76 is highly significant. This shows that any mean difference goeater than 4.32 is significant, and any greater than 4.76 is highly significant. 1.ese data show that hormone treatm nts 20 or 16 are significant in causing growth inhibitions when compared to checks. A study of table 27 shows that in general hormone treatment has little or no effect except in the case of soy been at the concentration 0f 2000 parts per million parts of seeds, and wheat seedlings which indicated slightly more tillering finder hormone treatnent. In table 27 a general summary of several preceding tables is found. Yarked secondary root developnen was indicated in the laboratory and greenhouse ex1eriments and it was noticeable in the field echri- ment. This is not due to hormone treatment necessarily but is a natural degelopment in the jrowth of corn. The original main root system of the li - soon dies and a large numb r of roots develoy from the crown V L): see and these are responsible for the nutrition of the corn plant and its later development. This loss of seminal roots and the heavy production of crown roots was studied for the writer's own satisfaction b: dig~iag up at the end of the season sheet 50 Sweet corn and 20 Dent corn plants. Later, phot03raphs were taken and same may be seen in plates 19 and 20. EXPERIHEII‘IV. ThE E7HTCTS O? HCnIOlE UN FLA TS WHEN APPLIED IN THE roar cs LALIOLII-T Pot Naphthalene acetic acid mixed with lanolin paste was injected below the egidermal cells on stems, petioles, and leaves of buckwheat plants at various stages of growth. These experiments were conducted in the greenhouse. In most all cases roots were produceé at the point of in— jection. The effects were not so different except at the highest con- centrations where cracking or bending of stem or petiole ocurred. After 10 days the wounds were healed, and after 2 weeks marked swelling ap- -31- peared. Then several epider;al cells would form knobs, showing that the roots would soon push thraufh. Consequently after 3 weeks, espe- cially during the sumter period the roots forned wvuld be about 2 cm. 10-2. In the winter period the elongation of these roots was very slow. However, root initiation followed application of hormones. Significantbud inhibition was observed in hormone treated plants growing in the gre nhouse, and plate 25 shows this. Tomato plants growing in 25 cm. pots in the greenhouse, with the temperature varying between 20-27OC., were treated with the lanolin paste (0.1” naphthalene 'acetic acid) on November 1. The average height of the plants at that time was 55 cm. (plate 22). To study coleoptile bending, Zea gays was selected be ease this was found (3i . l) to be more resistant to moisture and temperature variations than Avena in its early seedling stage. In this experiment naphthalene acetic acid mixed with lanolin was applied on the coleoptile with a needle. At first the experiments were carried out in the dark room but the re ults were so irregular that the method was revised. At the higher concentration (10 percent) the bend was negative. The same was true at the lower concentration (5 percent, table 29). Later it was found that moisture and temperature conditions were very important for coleoptile curvature. The new method of procedure was as follows. Glass vials were filled with moist sand and one day old germinated Zea .seedlings were planted in these (plate 20). Twenty-five vials were placed in a dish containing 2 to 3 cm. of water. They were covered with an inverted flower pot. This afforded the preper moisture conditions. When coleOptiles were up to 2 to 3 cm. in height lanolin hormone paste was applied at various heights upon the coleoptile. The temperature (25—27O '---r:¢..g. C) :as higher tn;n that in the dark room and more f vora ‘10 for coleo- ptile growth. In ta le 50 the results are shown. Applications at the node are more effe t ya and for a longer period, than those on any other part of the coleo tile. isually after 24 hours the cole O3tile returned to its ori3i 31 position if the hormone paste, had not beén too concent- rated or the humidity too low. The cha;:ge in the dir ction of curvature and the degree of curva- ture was found to be controlled more or less by age of part, the ano nt hormone paste, variations in m isture conditio. s, and changes in ten- pcrat re. on a consideration of the above facts the author believes that the phenomenon is one of polarity rather than hormone activi This will be discussed more in detail later. The writer has noticed that under certain conditions the rate of growth of the coleoptile and seedling leff er closed is concurrent, but under other conditions just the Opposite is true so that th -e coleoptile sometimes is empty. The lat er condition obtains when hormone p ste has been apjlied. This piste aff*cts the W'o st; uctures differently. The enclos d leaf is sensitive to the paste and its growth rate is reduced while the coleOQtile is more resistant (table 31). It is more resistant also to ch e ical i jury tlian the plunule (plate 25). feet of varying co centrations applied on Zea coleOptile H: The e tudied. The results are giten in table 52. Higher concentrations U) was often caused more curvature but the urvature due to age was more signi- ficant than thev var ”at ons caused by concentration differer c s. Th si~nific ant effects at hi _he r can Ht Mt on were more probably after a; n'vn ' ' ‘ N w *- -s 0v -“ —~ ‘~~ . r~ ‘ N i oats. .t hi,3her conce: nation the cur.a iris renaincu lon er and recovery was SlDTET. It :as fouhd also that r cover: ras slow r raen Sfmnetrically on intact coleoptile: it resulted in frorth irhir the tip of the plu.ule and r esultei in empty coleoptiles (plate 24). Znner igents "ith d -cfi\itatei coleoptile? are reported in table 0 “v “ .3.’ " 3 O : ion was carefully eon, ‘j c tuin3 the coleoftile tip H. d (,3 (+— k L a a ') d- (D Q U) 'd H J O (D C l: O H (0.? to 0.4 cu.) with a razor blade. Ear on C) We :3 1+ 3‘ k U *3 l ) “‘ 0 d- }, .J ( J ( I d. 0 L7 0 t‘5 I the cut surface unilate rall". The dif”ere< none paste hot ass intact and decapitated coleoptiles was one of degree. Decapitated coleiptiles r scted_quic:er. it the end of a fOlT hour -‘ perio on as had 0 rvature oecurred in the egspitatca coleoptile: While in intact cole ptiLer it was fficult to see any curvature. Decapi- tation anpears to wane the cole itile sensitive to her one t; eatment. The opti;un time for curvature depends upor; co:ditlons, an? a to 1? hours in that of decaxitated coleOptile s. Old coleoatiles do not react at any 0 ncentration of hormone While young ones Till. Since syr1etrica1 application of hormone did not cha mge the growth of the coleoptile, it is an import ht matter to decide whether or not Tendinr 1s ‘ue to fro th. Reports are nu ‘.ero is that the bending of the coleoptile is due to the activity of a phyto- hornone. Few are the reports as to the direction of the curvature. This to the writer appears more impor tznt thai the ce3ree of CJTV ture. 1 ”ant and his coeor-:e E) resorted t“at usually in the case of *3 A {:3 th'.‘.t is towards th3 light ’% 0 d- 0 d- ’1 O H. U) 1 I) S C) O [3 <: (1. CT L5 C) '3 O (n I" J k <1 (D U source, but if the light intensity were nefetive. They conclud;d that this di ”fe'e ea in reaction was due to the concentration of hormone. In the first case the horuone concen - ration was ligner on the shtded side while in the second case it was r.) V (}rani ; and his co orker (25) reported that it was ii ticult to pre iict the direction or dejree of bending of various plant structur:s when a ne p3rcent J-n- nro ionic acid in lanolin past was ap lie d. 4 study of be: din3 and cell elongation of various sturcctures indicated that it was difficult to predict the direction or the degree of ben- ding. The resp:nse to hormone trestuent Viried for the same tissues under different cc“aitio s of tem_ watuve, light, etc. In 93“9ral, the younger and more rapidly gru in; parts proiuced cerutures sooner. The press t exn -riment is in arreenent with this conceptio . In gsnsral it is oeli e"ed hat th3 nejative curvature (inva“c} due to direct injury and not to hormone tr ;tnent, since it has been I mpeate dl;r o s; rv d the t the coleoptile Tgei placed in 10'3 H . “Afi {Wt- -u‘J‘g? Lt1.e czrvatmre ani especially so when {3‘ (1) are conditions ex ibits a at higher concentration. A shrinkinr of the tissue on the concave side is ap;a;:ent. The author concludes therefore that n gative cur- vature is due to the injury of the tissue on the concave side. In the case of so: itive cur"at;.e, it is sugéested that this is related to chemico-physical polarity. This will be considered later. A summary of the eXp r1n,nt w*ith the Zea coleoytiles lS given below. "q l. Humidity is one of the i port nt factors in causing 8 Chen a in the direction of curvature. LOv humidity often produces ne;ative c rvatures, which the author ‘elieves is due to injury, probably chemi- cal in nature. 2. The after-effects arising from the curvature are ireater when the honn ne is ap lied on the node than on any other part of the coleo- ptile. 3. Small variations in the amount of lanolin paste used, had no . visible effects an c;rvature, but when the amount w.s increased to ten or more times, then the direction of curvature was changes. 4. The age of the coleo tile is an important factor in the type of curvature made. Ioun; parts react better than older parts. 5. When the concentrationof the hormone was increased, recovery was that much more retarded. 6. The decapit ted coleOptile is slightly more sensitive than the intact coleoptile when the hormone is applied in the form of lanolin paste. 7. Symmetrically ap-lied hormone on decapitated g§3_cole0ptiles had no effect on the elongation of the celeOptile but often inhibited the elongation of the leaf within. The effect of a 0.1 perc'nt naphthal.ne ecelic acid in the form of a dust (Rootone) on the rootin; of cuttinc; was stulied. Stems of cartsin C.) plants tgre cut in lengths of 5 2 lO 0:. and all leaves W re strifiped .— I. ,2: “t L.) (I) :3 .6 c+ C2 ' 5 (D 1 ed (0 off except a f J. Ihc cut ends were i_mcr <0. and the excess 30 der shaken of?. The cuttings were then planted i1 sand ... in flats to a depth of foir or five on. on October 4, and placed in the greenhouse. In some cases 0.3 :. "iootone" was scat cred alon: the .4 rows (o6 cm. lone) close to the cuttiigs and then covered with Sand. .13 Taterin; was always done from below and nev r from the tep, in order to prevent any pos ible reuov l of the hormone. At the end of forty days (Luv. If), all the plants were washed out of the sand and classified into groups (platcs 2;, 27, an; 28), according to the abundance of roots. — , All cutti;gs vith man; well developed roots were placed in froué A. Then a fewer n mber of roots but more than 5 were present and only no- ‘ aer'te de eIOpment was indicated these were placed in group B. Those 'J . cuttings with less than 5 roots, were put into group C, and in the final 3:4ip, I, 7332 31st the Jittings that were alive but had no roots. The results of the study of the grou‘s are expressed in pe centages and are found in table 34. Percentages are based on tot.l ninber of plants aliveo. Some of the pleats died but their death was due to causes other than the harmone treatment. 1he plants treated were Chrysanthe- u h" " *WA w rr" V V. “ IT! q 5‘ . A A": *4 q '1' ‘ .um1,;-esencruantneuuu, Kazan.ea, azaloa, Kleinia, and Crassula. ga~ (a, T- r" 7. ‘\‘ .\ yu - 1w . w ‘- v'- —\ v‘ ,- - s . if 1" I J- -. ,3 .0 . Meoemtryanthemim save a Letter root ddhelbpmdnt out none or the 0th rs ”are significantly be elitted Ly the her one treatment. Thether rootins in this case 7a: ate to the horuoie treatment. or so.e fortuitous \.J a b' factor should be tested further by epea in; the enneriient. It appears he ever that the treatments increased the neuter of roots in class B but did not accelerate root grotth ex est in the case of Tessxbrfanthemum (table as). Cuttings whose ronts were extremely agent, such as in many hardwoods or in Azalea, were “02828117 little improved with his particular uormons tregtuent \00). Rootone has not been Widely used to date and its ,--. ‘v «4 ;~--- W“ \TITI“ TT’“, r—T‘w‘p ‘ W * 7 " " 7‘” ' . ' * v P - ,- -. . .. - — " " *I M ' T “'I -' l‘ .4 ;~ .‘ “t w w . at; H '1" m.‘ *w T f1" W“... /.L.. . . .7 J, I .. 0 any -‘A‘~ .3L .._4 .Ja‘i-‘U$\ \J;\ w'V—J sA* J .LI-—J —>.~J“-‘ m- '6! ' . “J Uv.~.u 'Jtha’d.~A O lings usually form bulbous ins on all A. ( L) (I: Colehicine tre;ted Zea ser q - q 7‘ the roots, and root elongation is marked~y retar ed. This stalling of the TOOt is due to the formatiun of new polyploid cells. It was sugéested that hornone {pglication might stimulate gr nth of the pjljleid tissues. One dar old Dent corn seedli.gs were soak d for 20 hours in Colchi- cine. These treatei seedlings, thich at this time showei ma Red 3 ellings, were then washed in runnin; tap tat r for 34 heirs. They were now trans- ferred int; varying concentrations of naphthalene acetic acid in solution. They remained in this for 20 hours and then were W she; in run in; tap water for 4 hours. Their further gr; th was stulie: in petri dish cul- ‘1 tures. 0n observation the next da; no growth had occurred in 0.4 percent concentrati n, and no marked change in concentration of 0.004 pe cent. That is, the former was too strong and the latter too weak t2 cause growth or root navel; ment. At 0.04 bercent concentration, horev r, ew roots developed from t? e primarv root re r t 9 seed. The firOper concentration then stimulated secondary root develspnent on the prifiary roots. The treatment, honevcr, did not induce any further chance in the polynloid tissue as was hoped, but perhaps this mi3ht have been due to terninatin3 the exneriment too soon. Treatments and observations are tabulated (table 36). ‘7‘”? 3"3737-1 “*1 ‘“'“"‘."“-'r,'*3‘-" ‘ ' n3 "1"“? Trista-ohm '3‘“ “r rurr'i HI) .tn33iv'ro A" “$-41— luv-1.4001 (.1. O l -.A .b’..sJL.-._'A ll. vi. CL A..-LJ ‘4_.;..de. U: A.IJ...L HAI— D.—.a.‘.\.-.‘. -.__IJJ-‘) V'A'! fTT'T“ ’7""",'TLT 1"? fits-$3? orfvofs-“r‘fi on “:7 3r')\" 'jwfi-w M .3rfi'3-m-m arms L-“ C-§'\J.._-_ k A...- ‘Uklnxin p'w—J 41...); JM‘ rt: slxv 1L..- - ‘. ob \..- -4 ..-~.4 «J. .Q 'd “J‘ I. The advantages in the use of a box with a glass side for observation of seedlir~ arowth are a parent. Cne day old seedlings, germinated in moist L“? V oh i ers (petri dishes), as re placed on the $18 3 side of the box (60 x 30 I 15 cm.) and covered with moi: sand. Seeds had been previ- ously treated with indole butyric acid in dust form. The left side of the glass face of each box was covered with black paper to procide , 11 V, o darhnes s. as other Sid (D was eIposed to the li ght of the :reenhouse. Fhotor wf.s were taken diii lr until the pl‘uu le 3 had jronn above the top of the soil surface. Three such photoéraphs are presented (slates 00, 31, and 32). The 3*"rar1, of this experiment follows: 1. Darkness favors t‘1e elongation of tops, and esjecially the internode° for root dronth darkzzess is sliéhtlv better. 3 <__. a 2. Light retards top :rorth of young seedlinjs. inis is mainly die to the preventisn of r ‘th of he firs takes place to Mgre t: r extent in tted 5. hormone treat: nt (indole but ric primary root fronth and stinulzted the deve roots and 150 root ha" ir s. Yaxinum root develojment of vounje about 10 ays. sfter this period crown roo now function in the nutrition and developno 5. Retardation of growth in 2000 p.p.m.) was observed as early as tnen rj-n\ was started (plate UK: ; o 1"";‘7‘ 9171‘ “In V'TII “firm/«Tr '5‘ "w"? “w- n p ma “K“J-K fi'hhii A 0 fig. 4-1:} HEP :lv LIL .V. -~J Lid Jib .LQ. \fi FTr_ "7 "’! "2" $7,? F'TIM"‘ O‘fi R“ .L .._J :A.J\J .. \v'aIi '4'..- 2 The effect of naphtialene acetic acid I“ gen peroxide by ferric salts was studied. ‘ o ’." the hi 1es t internode. “longaticn t an in the light. acid 2000 p.o.:.) inhi it:d leXent of many latera r seedlinf mas attained in to start develoyinz. These nt of the corn plant. t concen ration of hormone ty a"s after the e: We riment LZTIC ACTI;K 0F FERRIC TLLT on the dec mposition of hydro- The importance of the cata- lytic effects of ferric salts in biochemistry have recertly bee n noted (2 ). lne possible effects of hormones on catalytic reactions in gene- rel have not been studied, as far as the ment describedb ere was car ied on accor and his co-workers (20). 0.4 per cent n8”htli ene acetic acid :w dissolvins 0.89. of the crystals with 5 CC V .J . writer is aware. r1 1 Y‘~ ufi‘-é The experi— to the procedure of Daniel solution was nrepared by L of 95 percent ethyl alcohol, and finally dilutin: this with distilled water. A 0.04 percent solution was prepared by diluting 5 cc of a 0.4 percent solutian with distilled water. The excess alcohol in the s;r3;ger solution was driven off by q gentle boiling for 10 ninutes while in tne weaker solution ooilin: was C”) continued for 10 minutes longer before dilutions nith water were co p- leted. The solutions used ii this experiment are indicated below. 1. 0.5 H Fe013 + 0.5 I HCl. 2. (l) + 0.04% naphthalene acetic acid. 0. (l) + 0.4% naphthalene acetic acid. 4. 0.0éf naphthalen ac-tic acid. The first contains Fe as the catalyst. The second has the Fe catalyst and a weak solution of the hormone. The third is the same as the second but the hormone.is ten times as concentrated. The last solution contains the honnone Without the catalyst. In preparation for a series of tit- rations, ocumercial H202 was diluted approiimately to a 0.5 percent con- centration, and concentrated H2804 was diluted (1:4). Distilled water was made ready at hand. Five minutes before titrations were started 10 cc of H202 were mixed with 15 cc of the various solutions to be :ested. From each of these was withdrawn 5 cc which was pipetted into Erlenmeyer flasks each containing 15 cc of diluted (1:4) H2804 and 10 cc of distilled water. At the proper time periods indicated in table 68 titrations were made. These data appear in table 58. Curves ere plotted for each solu- tion as well as for the check."0" and are shown in fig. 4. Curve "0" shows the amount 0 enn04 used to titrste the H303 solution alone and the value is found to be constant throughout. Curve "1" shows the de- composition of 1202 at succeeding titration periods and here the effect of the ferric salt is clearly indicated. Curve "2" shows that the rate of decomposition is retarded and this must be due to the press ce of na- phthalene ace»ic acid. Curve "5” ShOTS that decoxpOsition has almost stOpped. In this case ten times as much naphthalene acetic acid was used as in the solution represented in urve "2". Curve "4" represents what happens when no ferric salt is pre ent. No decomposition is apparent. If there is no chemical reaction between K n04 and naphthalene acetic acid, then all the curves should meet on line "0" at zero time. While curves "1", "2”, "5", and "4" do nojneet at zero tine, this may mean that either the alcohol used gs a solvent, or the impurities in the naphthalene acetic acid, or the latter itself, may account for the slight discrepancies. Since the effect is greater the -igher the concentration of naphthalene acetic acid used, it is believed that this is the cause of the result obtained rather than the little alcohol or the minute impurities. Further studies are planned. The data plotted on legarithmic paper show that curves "1" and "2" are of unimolecular form, that is, the rate of change dC/dt is prOpOT-‘ ‘4 tional to the concentration "C" of H909 at a fiven time, "t" - dC/dt {)0 C or " dC/dt = If. C. u Differentiating betueen initial concentration CO and concentration C c -j C/Cdet c O at time 13, —42- or Log (CO/C) : K , If where n is the reaction constant which varies with the type of curve. The value of K was calculated from the above equation, and is shown in table 89. In thesecalculations initial concentration "CO" was deter ined for each catalyst from the curves at zero time respectively. “~I 'J’Y 7? I 3" .2 -14 .4 _‘. _—_—..__..---. _ Germination of cirn was not significantly affected except when hormone concentrations were too high (2000 p.p.m.) and then it was reduced. These results do not confirm those of other workers. Corn embryos seem to be more resistant to hormone treatment than the seedlings. Soon after germ- inatisn has started inhibition begins to appear. Thinann (60) reported that Avena seeds treated with indole acetic acid solution (0.501 percent) for 24 hours showed an inc ease in germination. The author has noticed that under ideal condition (temperature, moisture, and freedom from microorganisms, etc.) good seed of corn and wheat will nearly always germinate 100 percent. This is possible when the embryo side of the corn kernel is kept moist until germination takes place. This is more im- portant than keeping the endosperm always net. When concentrations were higher than 100 p.p.m. (indole-butyric acid) tte growth of the primary roots of both corn and buckwheat we reduced. During a period of two to seven days this reduction amount d {1' to fifty per cent bot it was accompanied by an incrersed prod? of seccn‘ary roots and root hairs. it the end of a week the quantity of root growth was often remarkable. it this time regardless of the reatnen t, crorn roots developed and the further gro: th of the plant depended upon these roots and not on the primary seminal roots. In the case of corn the primary root system he only a temporary function and, therefore, exerts very lit ule influence on plar t growth (fig. 19 and 30). Hormon application to cuttings Lay be saitaole for the succulent type of plant but not for the woody kind aCCering to these studies. aoh Nth lene acetic acid has very little effect on rooting (table 55). Hormones increase rooting on young, succulent cuttir 5a in the non- doznant conditions, but produce very little or no effect on root elongation. The initiation could be attributed to chemical siiupla- tion of cell division. To determine this morphological irzvestigations are ne essary. If root systems are benefitted by hormone treatments, hen natural y top growth should also benefit, and increased growth should follow. This is the Opinion of many workers (ll). The writer agrees with the statement that hormone tre tnent fill initiate secondary root det elOpment but will inhihit primary root growth (59). be re :ds to. pro th the author believes that in most if not all of these cases the explanation is due to some other cause or groui of causes. :ost of the hormones are more or les difficultly soluble and of low diffu- si ility. Tany hormone treatments have been performed in solutions and the effects on the root are more direct while the effect on the top is indire t. The I ”(s H ‘ I concen rations r aching the tops are probably too weak to be effective. It is well recornized that increased growth can be brought about by chemical, thermal ani mechanical stimuli, and even by jarovisation. The latter is of limited value, however, ani can have lut little pra- ctical significance for the farmer. The writer has never heard of far- mers making money by utilizing the method of jarovisation.‘ In the wheat eXperiment (Exp. 11) it will be remembered that til- lering was increased and some might attribute this to hormone treatment, but such effects can be accomplished by chemical and mechanical means. In the northern part of Japan certain experiment station workers have advised farmers to tread upon seedlings of winter wheat in the field, in the hone that the pressure of the foot would induce tillering later. This proved to be a costly Operation and never practical. There are many factors (genetic, nutritio a1, soil, climatic, etc.) that can ex- plain beneficial results on normal growth other than relying on honnones to bring about normal growth. The most important studies of phytohorm nee has centered around the study of the effect of these growth substances on coleOptile curvatures and growth in length. If hormones do not increase growth of coleoptile (4, 54, 41), then there is very little to support the theory. In x- periment lV (table 53), it was found that a 0.1 percent naphthalene acetic acid in lanolin paste applied symmetrically on decapitated Egg coleOptiles resulted in an elongation of 50 percent after 13 iours, but that in the case of the check it was 90 percent. Instead of acce- lerating growth, it refuced it. If these results are correct, as the writer believes, then hormonesare of do Rtful value. In the case of plants, Smith (El) insisted "growth is the normal function of cells. They are always multiplyinfi when they are not inhibited by one thing or another." Eis conception gives us a very good idea of what normal growth is. Even auxin a, and b, and heteroauxin are absent from green tissues of higher plants (29). Uany experiments with higher plants show that a supply of hormone for normal growth is not necessary. There is n conclusive proof that phytohormones are specific, but Specificity of animal hO‘nones is established witho t doubt. It is well known that root inhibition, root hair develOpment, top groth, bud inhibition, internodal growth, coleOptile bending, and parthenocarpy, etc., 0*n be induced by various chemical, mechanical, thermal and electrical means. It is necessary to make clear now what is meant by the term hormone. Reasoning from known facts in animal physiolosy one would exyect phyto- hormones to be specific, but they are not. Consequently the term phyto- hormone is not justified. Kuogl (as) has recently complained tiat "the term must be limited to biological catalysts of organic nature which are used by the organism itself to bring about the various physiological effects." If plants possess hormones these must be quite different from the 'animal ty e, since the functional variations between animals and plants are great. It might be asked, can lover unicellular animals possess any or all of the homrones found in the higher animals? This is doubtful, since honnones in higher animals are not merely by products, but secretions from special organs auctioning in a definite way to con- trol the highly organized animal body. Without hormones the maintenance of life is impossible. For higher animals the nervous system acts as a telegraph, the circulatory system serv~s as a traffic system, fhile the hormones function as.mess ngers. Tithout these hormones the various s'stems are not properly or:anized and the living animal can not exist. Could life be expected in a man's legs or head picked up from a battle field and stuck into a nutrient solution? In certain conditions, cer- tain parts of plants can live and grow in a nutrient solution, and so k.) te tissues from hi 3n er animals be cultured. P- al;o can ch ain def in Growth promoting sulstances might be similar in their action to enzymes, salts, ugars, or sons polar CamgOundS which affect the free energy of surfaces. Kany nhvsiol 3ical processes in plants ar= carried on throu h surface b undaries which are well known sources of enerjy for the plants. The writer feels that the :osition taken by those advocating the existence of ohytohormones 18 not well f unded. From a functional ‘- D) and morphologiCol view poi t, the higher plant i a much more simple organism than t‘e animal and the correlations bet een the different (I) s if an? ax: also of a simpler type The plant has .1 parts and its hor_on no nervous system, or a circulatory system like the animal, and no che- mical messengers se m neces 3' ry to regulate and coordinate its various more or less si mp1 e reactions. The author has made a list of those substances most often reported exhibiting gromth promoting Characteristics, so as to make a study of their cherr cal structure. These are presented in table 40. They pos- sess active groups, -COOH, -OE, etc., which have high affinity for water, and also nonactive groups, such as carbon chains or rin s I.hi ch have high affinitfikn7 fat solvents, but not for water. The su3ges ion is offered that wlen such polar c npounds are applied unilaterally, they penetrate into the cell. ?olar compounds by nature change the free energy -47- of the surface and thus indies va riftions in nsrxesbility, chanjes in concentration, mOdifiCStl;RS in turgor pressure, etc., on one side of the coleogtile "here “r*lieu, and thus brin3 about curvature. In addi- tion to this function of polar compounds in modif yin 3 surface tension throu3h molecular orientatiin, they also rerulote and ori~inate electro- motive forces (e.mwf.) in the plant body, sir es the total e.n.f. of a plant is the sum of the e.m.f.'s of the unit cells (18). Since a plan is mane up 0 many cells, a snail chcn3e in e.m.f. in a unit cell will brin3 about a large change in the whole plant. In penetration of these polar con; unds must be very slow and when they do penetrate, their effects are lar33: y limited. 'Ile f rst effects are near or on the polar compound an e.m.f. is created, it is proballe that some polar compounEs and ion ns already present will migrate from one side of the coleoptile to that which was chemically treated. In this way curvature would esult through modification of tir3idity From.the time of Vochting (Went an Thimann "on03raph) (1878- 1908) to the present the existence of polarity in correlation phenomena has been definitely proven. This is not onlv so in the case of whole organs like one of the higher 3reen plants, but for its parts, such as leaves, stems, root and fruit. Even each separate cell exhibits polarity. In more recent times electro-polaritr has been demonstrated and is non 3enerally conceded. A number of possible differences in potentials existing in living plants togeth er with their probable causes, are listed below: 1. Concentration p tentials exist between solutions of different concentrations. Such are possible in the cell sap of different tissues. 2. Diffm mien potentials are set ip when miscible solutions diffuse 3. Liquid juncti n potentia 3. When liquid A is iicsolved in a mixed solution of t.o in issible st strnccs ant C, tn-n 1 is i-lly distri«nted in F an? O and co ‘ien l a potan ti.”l diffcieice 1" ceind t e:-:' at . 4. Xe brene potentials are prifalent in plznis s. They may arise thr; 3h (l) are: us I c‘otrib-tion of the ions on either side of the .1. .- v‘r 'X ~ » «A. r) '.» a ,- ' ‘ .. nenbrane and (u) by tie unem;'r pezetza tion or diffusion of t; thrOLnfii the :naulrane.. 5. Injury poaent uls. I. some case 3 the mechanisn is unknown. *— I H. O “-1. P C?" (+- L Li 9 5 $3 (+- O k—ub In other cases it is susp cted to be due to stimu injury the potential is high and tlen fades a a,. iie injured part is egative to the uninjured, due, indirectly, as so 9 th 1k, b0 increasin3 e. Oxidation-reducti n potential. In tissues netaoolic processes are occurring at differgnt rates ard are rot uniform. In co.seiuence potentials rise. One can observe this under the microscope. Dyes such as methyleneblue can be injected into tiesues and their further cnanges observed. Yethylene blue changes to the colorless f»rm then reduced. Some livi-: u) tiss J83 or ci3ani3ns contain pigments that chan3e to the leuco form on reductiOn. Quantitative tests can thus be :ade. 7. Electro kinetic potentials. These potentials arise from certain surface henoflenc, S"Ch as selecti"e ads orption an ionisation of pro- teins, etc. 8. Potential variatigns mithin a cell are very well lznown. Since the nucleus is more ne3ative than the cytOplnsm the cell as a whole is -49- Protoplarn is said normally to ie cl~ctro negative an? thrau n stigu- e in the part excited and therefore in- creasing the potential and nakin: the protOplasm or cer ain parts of the plant mor active. Polarity might be affected in this nay. The presence of a Sth or s-'13 {59) nov also affe t polarity and by analogy the hem ‘ n3 of the coleo tile. For, as is ell 110.“, salts change the ionization rate and form, an: sa3ar stabilizes the ollcidxl micelles (Ff). Inlirectly then, polarity 's affects d 0y electric potentials. Thimann and his coworker (33) s ggestgd that the activity of a honnone is due to its double bond. But double bonds appear to be of little immortunce in physiological groces:es. i-Counaryl acetic acid, has is een reported (56) to affect root initstion out not- Arena curvature. SiiCe this compound has two active groups, -CC;K and ~CH in the struc- u11c, its solubi;ity must be too high to be polar. It diffuses too easil“ in all directio:1s an ’ hence can not bring about the elJn°atio 41 e of the cole: tile. The compound L7 iis effects must be L i 9 FJI of gne 3 less polar. V 1 -. .,_. _-‘, , ° 9 3 ,1 ,.° Farther explanation 01 the use ani U) m of polarity and of polar compounds is beyond the scope of this pahrr bit the author would like to introduce here several brief reports ahich affirm the polar theory and the existence of electric potentials. Erauncr (lb), in l 33, reported that in th: case of horizontally placei Jlant parts the lo er side sees me electro posi ive to the apper side. The shaded side of the .w f". ssedlin? stem :23 cl. tro-posi t11ve to the illu1;Hirated side. Photo cngrc” it may h1 said, has lean transfo med i1to foteztial c :r3y. rather and his co orher (14}, in lPoC, reported th ”.11331 coleoptiles carved to 2r} the n sitivc pole in an electric field. Cheye- fore the convex side must be positiveto the concave side. In lso7, Clarl (K ) stidied th ejolarity of ;lant parts in ccnxid- ssified, (a) into acid dyes, (D r. : -,-- "I ,. .14.- ,,.- . ‘- eraole detail. 1hc polar also an 3 cl light 3ree n, acid green, methyl oran e, etc.,and (3) into basic dyes, Sa nin, methyl violet, nedtral red, gentian violet, etc. Then Vicia iected with basic dyes (positive 1; char'“?), the dyes U _233 roots were i accum1lat ed on the concave side. These dye“ are :iotaoly toxic and it n.uld seem tlat the stimulus and high potentials Wiuld negative any results. Clark also re orted that the intaCt.£:EE? tip is electro ne- Ih 195* Czaja (19) discsssed the mechanism of membrane polarit Cell walls of lo er and higher plants sho ed affinity for basic dyes. In Spirogyra cells, the cation of hasic dyes was first adsorbed by the cell wall and then as a d e so It pass ed into the cytOplasm. with dilated solutions of basic dyes, nearly the whole of the anion rem ined in t11e exterr al soluti»n. Adso T:Wti n of apprOpriate salts into the basic dye solution retarded the adsorption of the cation and inhibited its passage into the cell. Thimann and his coworktr (69) st ted that the formation f the growth pro oting substances which affect égegagurvature is limited to the tip, 1 I bdt when the tip had been remove d, a new zone of auxin formation w-s produced at the apex of the stlmp after two to three hours. This (N1 regenerated tip showed no hist0103iczl differentiation, so that auxin formation was not necessarily associated with special cells. This fact it seems would indicate that the coleOQtile itself exhibited electro polarity, and accumulated some chemicals which affected the bending of the coleogtile. However, curvatures produced by external application of chemicals a are temporary, asArule, at loner concentration, but at higher concent- rations permanent injury results. Electro polarity will not entirely account for curvatures, yet it appears to be the best explanation for many tropisms. fivw-P hm? “ I . ' I l. TAB germination of certain seeds was not affected by the "phyto-horhones" used unless the concentration was very high and then it was markedly r duced. Two authorities have claimed that germination is speeded to such an extent as to be of practical value. 2. Primary root develOpmert has significantly reduced when higher concentrations were used but was in no way affected at lower concent- rations. At concentrations high enou3h to be effective but not too high to be deleterious secondary root growth and root hair develOpment Was stimulated. In corn the seminal roots were highly accelerated in their growth by hormone treatment but the crown roots which replaced these later were not affected. Consequently the plant snowed no later improvement. TOp 3rowth of young plant was not affected by hormone treatment unless concen rated solutions were used and then inhibition occurred in the roots as well as in the teps. T en the high concentrations did not kill the root, but both the root and tops were only inhibited there was later a recovery so the no significant difference could be detected in yield of fruit, dry weitht, or aipearance when compared to checks. This was true of plants raised to maturity either in the croon- house or in the field. This is not in agreement with the studies of Grace. Treatment of seeds with hormone in the farm of dust before plant- ing in the field will not give results of any practical value to the farmer. 3. When the hormone was applied to petioles, stems, and leaves (buckwheat) in the form of lanolin pas e, roots were produced. Such root initiation has been observed on other species. Significant bud inhibition was also obtained in the case of tomato. This confirms other workers' findings in the case of other species. ”nilateral application of hormonized lanolin paste on £33 coleOptiles at different levels was more effective on curvatures when applied at the node than at any other place. Hormone lanolin no te applied to the coleoptile has little or no effect on elongation but does inhibit that of the leaf within. After such tr atment the coleOptile appears empty. No report of such a reaction by the plunule has *een observed by the writer. Certain parts of pla1ts seem to he more resistant to hornone'treatnen . The order below starts with the most resistant; First internoda coleoptile>r;>ot hair> 131131111167 secondary root? prihary root. 4. Hormone treatment of needy cuttings has little or no practical value. hormone tr: tnent of vegetative cuttings produces in most cases few roots which do not elongate to any extent. 5. Laphthalene acetic acid inhibited the deconposition of H202 by a ferric salt. 6. Chemical me sen;ers (hormones) of the type occurring in animals have not yet been demonstrated in plants. 7. The most plausible explanation of coleoptile curvatures and rel.ted phenomena is that of (a) chemical etc. injury and (b) electro- polarity. IT m‘ra "rvyv-fx-rfi J.L 4...; ~Ao; .. .4 l. Albaum, H.G. Inhibition due to growth hormones in fern prothallia and spore hytes. Am. J. Bot. 32: 124-132. 19351 2. Avery, G.S., P. G. Burhholder, and _ . Creifihton. Growth hormone in terminal shoots of Nicotiana in relation to light. Ibid. 24: 666-673. 1937. 3. Avery, G.S. an 0.0. Larue. Growth and trepic responses of excised §3§n§_coleoptiles in culture. Bot. Gaz. 192; lee-199. 1938 5; Bernburg, L.3. Beitrage zur Frage der Dosierung von Trocken- beizmitteln fur kleinste Nengen feiner O‘merien. Z. Pfl. Krankh. g2; 596-603. 1937. 6. Blum, H. and K. Scott. Photodynamically induced trepisms in plant roots. ?lant Dhysiol. E: 525- 37. 1933. 7. Bonner, J. Relation of hydrogen ions to the growth rate of the .izena coleOptile. Protoplasma El: 406-423. 1934. 8. . Thiamin (vitamin Bl) and the growth of roots: The relation of cheriical structure to physiolonical activity. Am. J. Bot. 5: 543-549. 1938 9. Bose, J.C. The motor mechanism of plants. Longnans, Green a Co. New York. 1926. 11. Boysen Jensen, P. Growth hormones. Trans. “very .S. and ?.R. Burkholder. HcGrew-Fill Book 00., law i'ork. 1936. 10. Boysen Jensen, P and J.Iielsen. Studien fiber die hormon- alen Peziehzngen znischen Spits e und ?asis der Avena-koleOptile. Planta 1: 321-331. 1935 4; fi.... 0 o — A- _ A... 0.. “u N. O V. a w... h. L e, e o 1..» LL .1 .w :L D . m... Lu . V.),V 1% 1 _. cain may C Q 3 .m. C .51— «J +L .L b a,“ C a I as e ‘ . 1 n H... .. Wt. 7A 2 “i. 3 4.. .. v Q 4 +. y. C n). H. +.. 3 “I” r . i C 1. .Tv. "I O + « hi. 0 X L a . t 9 ‘ w. a. . a. O T. 3 ”a a .. . x‘ , . D . as; 7 it.' n... C n... o his 1-. HQ C l r o e ; Mu 1L ween .L- '4’ O éAW "’ U'. .- ---\. 0 an ~I 1‘ do I‘ a Q J9... rut- ,. .--~4-’~ “1 :7; “ '1 V.» ‘1 “ "J ‘ . A U .fi- 1‘. “’\h‘! .J A ~/_ - ot-‘ v ‘ L. r 3 -A' v V‘- T" \ ls. l \— 5H .A- v "an. .-QJ- 9'9, 1 .AL\-A\.L . us”, M r 75“- .1 .L 1'7. PP?- N_. y‘ +r. 1 . I ‘45 1’“ O T‘lv .-+ U “-9 .b‘v‘-4 \ . al. 3"” Wavy” 1.}. .A—s'... . -‘ O 7‘ c AAA ." A’J Q 3 .L‘. A“ +3fi . \ fl- J H._ a a .4 Hi. .1. o o a; a as a q..* r _ x; A 1“ Fr «fl 3... .- m.- n __ n~ '0 h. v. +» Tn w 7 l “u , . m a... .1” k. 11— L. a. .4 f. by s w . o _. Th .1 Y- .. ’ .1 o 11 H... .r-. ’ L. 1; .. m. 4‘ xi. .. “A ‘4'» o 0“ 'Y‘ 1 -5-"‘ 1 ' .LJ-- 0" ~.- -.r~n fl 9 -O .4" 3‘er v.- 0‘ A..-/ I . -.\ ”.Ov‘ 1‘. «a '3 \ A mi ‘7“"‘ ‘ * né (1 .-n-v (.4- ,5. " —l" ‘N n” n... . .. n4 . V‘ r‘ 5 " '; 'JeooaJu-VQL .13. f‘ I. :4. Grace, Y.K. Physiologic curve of response to phytohormones by seeds, growing plants, cuttin7s, and low2r plc nt forms. Canadian Jour. Res. 15; 538- 546. 1937 25. Granick, 3. and £7. Dunham. Growth responses of various plants to n50 1e 3-n-pr0pionic acid. Pa 3e rs of? ichigan Acad. Sc. Arts and Letters-gg: 69~77. 1936. 26. Gustafson, F.G. Inducement of fruit develOpnent by growth prom ting chemicals. Proc. 1.:t. Lead. €c._2§: 338-636. 1936. 27. Greenfield, 8.3. Reaponses of stock seedlings to hetero-auxin ap_lied to the so: 1. Am. J. Bot. 24: 494-499. 1937. 33. "itchcoc:, A.2. Indele-3-pr0pionic acid as aggrowth hormone and the quantitative measurement of plant respor ms . Contrb. Boyce- Thompson Inst. Plant. Fes._z: 87-95. 1933. 29. , and 7.3. Zimmerman. The use of green tissue objects for date iining the phvc1o o ical activity of growth substances. Ibid. 30. K0051, F. fiber 71 hsstoffe der Auxin-und der Tios7ruppe. X17} fiitteilung uber p lans lich Wachstu nsstoffe. Ber. Deut. Chem. 988. Hft. A. 68: 16-38.1935. 31. , E.J. Hangen Smit, und H. rxleben. V. 'Kitteilung. '1 fiber ein Phytohormone der Zellstreckung. Reindarstellung ées AiJins aus menschli chen,Harn. Heppe-Seyl. Zei ts chr. 7hysi01. Chem. 214: 241-261. 1933. £24 , , und . VII. Hitteilung. " Studien uber das Vorkonmen von AuXines 1m.nenschlichen und in tierischen Organismus. Ibid. 220: 137-161. 1933. .Uq 1 33 O , , und 0 X1 . fiitteilung. Uber ein neues Auxin aus Earn. Ibid. 398: 90-105. 1954. 2'4 0 ’ ’ und 0 Jill O Mit‘eilung. Uber den Einfluss der Auxine auf das Wurzelvachstum und uber die che ische Ea ur des Auxins der Graskoleootilen. Ibid. 298: 104— 112. 1934. . 30031, P. On plant growth hormones. Chem. and Indus. (London) :6. Inge, Eh D. and W. F. Loomis. Growth of the first internode of the epicotyl in Kaize seedlings. Am. J. Bot. 2 * 543-547. 1937. 38. Leonian, L. H. and V. G. Lilly. Is heteroauxin a growth promoting substance? Am. J. Bot. g4: 135-139. 1937. 39. Loehwing, W. G. and L. C. Bauguess. Plant growth effec s of heteroauxin applied to the soil and plants. Science n.s. 54: 46-47. 1936. 40. Iarotta, Luisa. (EXperiments on root sprouts cut off and gmmtn in nutrient solution). .Atti. Acad. Lincei_§§: 515-517. 1936. 41. Earner, D. R. Growth of Wheat seedli gs in solutions contain- ing chemical growth substance. .Am. J. Bot. E3: 169-145. 1937. 43. Feesters, L._“. The influence of Agrostemma githago. Proc. Akid. Sci. Amsterdam, 39: 91-97. 1936. 43. fiiller, W. L. Bios. Jour. Chen. Educ. 2: 256-237. 1930. 44. Paal, A. T“oer phototrogische Reizleitung. Jahr. Wise. Bot. 58: 406-438. 1919. * 45. Pratt, R. Influence of indole-3-aéetic acid on the respiration and growth of intact wheat seedlings. Am. J. Bot. ‘3: 389-392. 1938. 46. Purdy, H. 3. Studies on thu path of transmission of photo- t20pic an; geotrOpic stimuli in the coleoptile of 3:333, Biol. Tedd. g ('6): 3-29. 1921. 47. Rothsrt, W. Uber HeliotrOpismws. Teitr. Biol. Pflanzen Z: 48. Dakamura, T. und T. Yanagihara. Zur Bildung des Tuchsstoffes bei Aspergillus niger. 1’roc. Imp. Acad. Tokyo 8: 597-a99. 1952. 49. Shog, F. The effect of x—irradiation on auiin and plant growth. Jour. Cell. Con . Physiol. 7: 237-270. 1935. SO. . Absorption and translocation of auxin. Am. J. Bot. 25: 361-372. 1936. H O Saith, E. F. Kechanism of tumor 7ro: h in crowngall. Jour. figr. Res. E: 163-1EE. 1917. 52. Snow, R. Activation of c mbial growth by pure hornanes. Ken ~*‘hytol. 34: 547-3QC. 19J5. 53 Soding, H. Uber die Tachstursmechanik der KaferkoleOptile. Jahro. 7iss. Bot. 29: 231-2; . 1964. 54. Starling, E. 3. Die chemis he Ioordination der Karpertfitig-> :eiter. Véih. Ges. Deutsch. haturf. Afzt., Stuttgart 7F 91): 246-260. 55. Theodor, S. at D. Constantinesco. Action de l'acide b-indolyl Fcetique sur la germination et le Development des Grins. Compt. Bend. 492-494. 1937. U 0 O to F). O H O |.._l {0 n3 56. Thimann, I. V. On an analysis of the aétivity of two growth- pronoting substances on plant tissues. Proc. K. A ad. Wetensch. Amster- dam 58: €9"-912. 19u5. 57. . On the nature of inhibition caused by auxin. 58. and C. L. Schneider. The role of salts, hydro- I. arena coleoptile sen-ion concentr; iJn, ani agar in the response of the » t0 83:31:18.3. {1.31. J. 201;. BE: 270‘2500 19-)70 59. and J. Bonner. P1a.c growth hormone. Physiol. JO. and 3. K. Lane, After-efifects of treatment of seed with auxin. Am. J. Bot. 35: 535-545. 1952. —-——- 51. Traib, E. P. Growth substances with particu;ar reference to subtrOpical fruit plants. Proc. Amer. Soc. Hort. Sci. 3 : 4JE-442. 19J8. 62. neij, E. Van.Der. Der Techanismus des Tuchsstofftransportes. Rec. Trev. Bot. Ieerl. fig; 379-496. 1952. 65. ?ent, F. T. finchsstoff und Vachstum. ITid. 2 : 1-116. 1928. 64. . On the pea test method for auxin, the plant Tetensch. A sterdan 7' r:17-575. 1934 *5 U 0 O 4 1 C b“ I! L, O hormone, P O ‘- "F 65. , and n. V. Thiuann. Phytohormones. Tacmillan Co. ‘T u. ‘r w- n z :3” rorn. 1957. f‘ H. v-u- ‘w« 1" 9" fl ‘F"' ‘ ‘ fl 7... ‘n 1A" ‘,‘ . 4-. . I ." o7. ulmmelmhn, P. .., h. UTOCWUr, 8R1 n. A. nitcscoch. Zululatlon and sti ulation of roots from esposure of plants to carbon monoxide res. Contrib. Boyce-Thompson Inst. Plant Res. a: 1-17. 1935. cc. ~nd T. Tilcaxon. Several chemical growth suastances which cause initiation of roots anl other responses in plants. .1EE IL v F',. T7413]. e ()1 ( 0 Description of chemicals ans “H V n‘. osst nrm er, hormone, - ., Optinnii Calcilation‘ of 'ptlirld o‘ (irowth Growth Growth Growth (H buckwheat s of of of of of of O 01 vfi“ LLDU I parts of hormone per mi1_ion o? )ercentase of ’5 ‘4 seed and quantities of in rt dust used to make up mixtur-s. pst adsorbing capacity of 10 trans of seals. (D (D ed'. adsorbing capacity of 100 gram seed as files. Dent corn roots on hormone dzsted filter paper. dusted eht corn seeds on filter paper. Dent corn on hormone dusted filter paper. hormone dusted tant corn on filter paper. dusted Sxeet corn seeds on filter paper. Sweet corn seeds on dustel filter paper. dusted buczwheat Wheat seeds on filter paper. Summary of experinent 1. Some preliminary obs rvations of plants growi g in hormone treated ,- o “ __ _‘. soil in the greenhcise. Gr wth of dusted Sweet {Er 3‘I1I't“ Gro th Growth f" f n. 1.‘ err-J" on Calculation for the siyzificance of antone of‘ V corn seeds in the frenhouse. Growth of buckwheat in dusted soil in greenh'use. bucks eat on dusted 5 i1, sun.ery of tabl s 13 and 17. in greenhouse. a o Rootone dusted wheat and millet seeds in greenhouse. treatxent on tiller .J to x] O F; [\3 C) C) (.3 ()1 O] (J! C»! (J proi.ciion of wheat. Grinth of Dent corn dis ed seede in the field. Growth of Sfeet corn dustei seeds in the field. Growth of dusted buck Lest seeds in the field. “ea°.rciei.s of height of p en's from di:ted 9zee t corn seeds in the field. with hor..one dust. p. os-rvati:n on oif“erent seedlin s trgg ‘ -. . o t . L' n . . fl '1' — ~ :- “ 9 ~r Le res and direction 0; coivature 0. rec coleoptiles at lob HamlilfJ. '5‘ " A r ‘ ‘r‘x f P 4 “- r . " r '0 y ' “(- Sifects of naphthalene acetic ac_d in the iorm of lanolin paste ‘ ,: - ',—.. ' a .1 P7- .',.‘H:.-,-. aojlie at vario.s points on noa seeeii“ s. of curvat‘ire . "1‘... .3 , 4. n 1..6 e1 1930 U3 0 ~. Various cone ntration of hornone in lanolin paste The effects f norm-ne in lanolin piste on decapitate Zea coleoptiles. affects an cu tinjs through the use of Rootone. ln‘ Rooting ef¢zcts of c ttihgs die to Rootone, c lculated in percen - ages based on total cettings alive. Effects of hormone on corn seedling previ0151y treated with colchicine. Solution miztu es with and without catalyst. Titrations for decent osition of .32-00 v-l. Rerction velocity constant for de: QOL osition of H...On ‘44 ~40 Clemicals report 1 TS horzi one or as hormone-l ike in activity. .Insrnlru P, .. . .. II LIE.‘ “‘7 r r. ‘\ n AWj'rrn I...“ .41 4. J'RAL Ila.) Weight of dust added and excess for the determination of Optimum by 100 gram.seeds. Concentration paper. Concentration paper. Decomqosition of hormone and percentage growth of Dent corn on filter of hormone and percentage growth of buckwheat on filter of 3gOgtitrated with 0.1 N KYnO4. (7‘ o 10. ll. 12. 13. Large moist chamber and pan for ~V..-1.‘, ) . f -..4 .‘JL Porous pot moist chamber. Seedlings fro; hormone paper. Seedlings from hormone paper. Seedlizgs from hormone Sweet corn seedling on light. dusted dusted dusted sweet corn seeds growing hormone dusted filter paper under diffused L V T t F1170 ' 1.118 .L..u.) jeitdnri tion. Sweet corn seeds growing on filter Sweet corn seeds Sweet corn seedlings freeing on hormone darkness. from hornone Seedlings from hormone Seedlings paper (Aug. 15 paper (Aug. 19). Seedlings from horxone paper (Aug. 23). Seedlings from hormone paper (Aug. 23). Seedlings from hormone paper (Aug. 25). Seedlings from hormone ). paper (A g. 2. dusted dusted dusted dusted buckwheat buckwheat buckwheat buckwheat buckwheat buckwheat dusted filter seeds seeds seeds seeds seeds seeds growing growing growing growing growing rosin? on filter on filter paper in on on on on on on filter filter filter filter filter . 13F) ‘14» at. Y: ‘r .IM 27. 28. 29. 51. 32. Seedlings from hormone dus ed 8 set corn seeds growing in the greenhouse. r g. Euckwheat growing in hormone dusted soil. Soy been plan 3 grown from hormone dusted seeds growing in the greenhouse. Ro tone treated tomato seedlings growing in pots in the greenhouse. Root system of millet grown from Rootone treated seeds. Root system.of Q ~...4 :eet corn srown in the field. u Root systmn ofIDent corn grown in the field. Aerial roots produced on buckwheat stems and petioles by application of hormone in lanolin paste. The e‘fects of The effects of ptiles. The effects of coleOptilos. The effects of The effects of The effects of The effects of Seedlings from solution. Dent corn seedlings Dent corn seedlings Dent corn seedlings hormone in lanolin paste on tomato plants. quantity of lanolin paste apglied and the are of Zea coleo- various hormone Rootone Rootone Rootone concentrationsof hormone in lanolin paste on Zea in lanolin paste on decapitated Zea coleoptiles. on roo.in* of cuttings of Chrysanthemum. on rootin; of fiesemhryanthemum.andiKleinia cuttings. on the rooting of Hydrangea and Craesula cuttings. colchicin treated Dent corn seeds retreated with hormone growing in boxes with one side provided with glass. growing in boxes with one side provided with glass. growing in boxes witi one side provided with glass. \. “II-liklulnfih. RIF-IN HE .41 . fl‘w TABLE 1-40 Itinel'll,1 W5 Table l. Ibscription of Chemicals and Plants Used. Substances OSHONES AND CHEXICALS: B-Indole-butyric acid (5g.) Naphthalene acetic acid (lg.) (5005-) (0.5g.) Colchicine 4% solution Talc powder Colchicine Lanolin paste Rootone(naphthalene acetic acid) SEE; D8: ' Zea mays (Ibnt corn) 1937 I! Triticum vulgare (wheat) FaEOpyrum esculentum(ouckwheat)" Panicum miliaceum (millet) " Phaeeolus sp. (kidney bean) Pisum sativum (pea) " Zea mays (sweet corn) Soya max (soy bean) " Lactuca sativa (lettuce) " CUTTINGS: LyCOpersicon esculentum (tomato) Chrysanthemum Hydrangea Opuloides Azalea sp. Mesembryanthemum crystallinum Kleinia repens Crassula arborescens From Dr.R.P. Hibbard, Bot.EEpt. Dr.E.H. Newcomer,Bot.Dept. Stock room, Bot.Dept. American Chem. Paint Co. Dr.R.P. Hiobard, Bot.Dept. N H N ' N I! ll Farm crOp dept. H Ferry-Horse Seed Co.Detroit. Dr.R.P. Hibbard, Bot.Dept. Kr;fl.A. Frost, Hort.Dept. Prfifo COE. Y'Vildon, " I! H Prof.H.C. BeeSKOW, Bot.Dept. H H H II . . I i .I-4.B r ‘5...“ l‘ix - ‘ Table 2. Dust Number Parts of Hormone per Million of Seed, Percentage of Hormone and Quantities of Inert Dust Used to flake up Mixtures. Wt. dust 1 Dust No. Hormone e 00 . l'ormone page%s g in dust p.p.m. g % 0083* l 0.00 0.2 0.0 2 0.05 0.0025 3 0.10 0.0050 4 2 0.1 5 6 " 0.3 6 10 0.5 7 20 1.0 8 40 2 9 50 3 10 80 4 11 100 " 5 12 120 6 13 150 7 14 160 8 15 180 9 16 200 10 17 400 20 18 1000 0.1 100 19 1600 0.16 100 20 2000 0.2 100 BUCKWHEAE* 1 0.0 0.3 0.0000 2 0.075 0.0025 3 0.15 0.0050 4 3 0.1000 5 30 u 1 6 300 10 7 3000 100 30! BEN-4'4““ 1 0 0.2 0.0 2 2 0.1 3 10 " 0.5 4 20 1.0 5 2000 100.0 * Indole butyric acid. ** Alpha-naphthalene acetic acid. Table 3. Optimum Dust Adsorbing Capacity of 100 g. of Seeds. Seeds n Y X Adsorbed (added) (excess) g. g. 5. Dent Corn 1 0.507 0.245 0.262 2 1.053 0.785 0.268 3 1.557 1.269 0.288 4 2.065 1.775 0.290 5 3.082 2.736 0.296 Soy Bean 1 2.001 1.750 0.251 2 2.554 2.301 0.253 3 3.215 2.958 0.257 4 3.754 3.500 0.260 5 4.790 4.496 0.294 Buckwheat 1 1.054 0.571 0.483 2 1.990 1.505 0.484 3 2.010 1.560 0.483 4 2.510 1.858 0.652 5 4.010 3.270 0.720 Table 4. Calculations for the Determination of Optimum Dust Adsorbing Capacity of 100 Gram Samples of Buckwheat Seeds. Actual weights Weights from equation n (edged) 4(e:cess) 1 V 1 X 8 ' 8 8 8 1 1.054 0.571 0.505 0.0 2 1.990 1.505 1.050 0.654 3 2.010 1.560 2.010 1.490 4 2.510 1.858 4.010 3.250 5 4.010 5.270 -- -- Table 5. Optimum Dust Adsorbing Capacity of 100 gram Seed Samples. Wt. of D0st per 100g. seeds Seeds Optimum Applied 8 6 Dent Corn 0.26 0.2 Soy Bean 0.24 . 0.2 Buckwheat 0.305 0.3 Wheat 0.3* 0.} * Approximate value. E. v. 8‘ Table 6. Filter Paper. Growth of Dent Corn Roots on Hormone Dusted Seeds previously soaked 12 hours and set on dusted filter paper (120 cm?) with 0.02g. indole butyric acid, April 23. 27°C. Mean of 10 plants. Germination Apr. 25. Apr. 26 for N0. 7. Room temperature 22- Hormone Concentrations Root length N0. mg/paper Ag;.27 ($1 45;.25 L3) 432-29 L41_ 1 0.00 7.3 100 13.5 100 16.0 100 2 0.0005 7.2 99 11.3 53 13.5 84 3 0.001 7.2 99 10.5 78 11.0 69 4 0.02 6.0 82 8.0 60 9.5 59 5 0.06 4.3 59 6.5 48 9.2 58 6 0.1 3.0 41 6.2 46 8.8 55 7 0.2 3.0 41 5.5 51 7.2 45 (2) = Age in days, after germination. Table 7. Growth of Dusted Dent Corn Seeds Set on Filter Paper. Hormone used, indole butyric acid. Set in petri dish cultures May 7. Germination May 10. Room temperature 22-25°C. Mean of 10 plants. Conggzigggions Length Of TOD £T)(%)§20t L?)o) May 13 (3) may 15 (5). m_1_1§ May 20 110. 5p.o.m. T R T R T T cm CH] cm cm CH] CH] 1 0.0 6.7 14.1 12.9 19.1 18.5 20.1 2 0.05 7.5 13.5 13.1 18.0 17.0 18.5 3 0.1 5.5 12.9 12.2 16.5 19.0 22.2 4 2 6.0 13.0 12.4 19.0 17.0 17.5 6 10 7.3 14.0* 12.3 18.7 16.0 i 21.1 I 20 6.8 11.0* 12.2 16.2 18.8 21.3 " """""""" 72 """ 72 """ 7's """" i """ 7 """ 7T" 1 0.0 100 100 100 100 100 100 2 0.05 112 96 102 94 92 92 3 0.1 82 91 94 86 103 110 4 2 9O 92 96 99 92 87 6 10 107 100 95 99 86 104 7 20 102 78 94 85 102 106 * Secondary roots appearing. ** Root branching on primary root. (3) = Age in days after germination. .7158‘lb.l5!uflfifivu! r‘ . . Table 8. Growth of Dent Corn on Hormone Dusted Filter Paper. Seeds previously soaked 4 hours and set on paper May 7. Germination May 10. normone, indole butyric acid 0.02 g. on filter paper (1200m2). Room temperature 22-2500. Mean of 10 plants. Hormone Length of T0p (T) & Root (R) Concentration May_13 (3) May 15 (5) Eégil§ ‘Méifgg No. mg/paper T R T E T T #_ cm cm cm cm cm cm 1 0.0 7.5 12.3 13.6 16.1 20.3 26.2 2 0.0005 7.1 10.5 13.5 14.2 17.2 --- 3 0.001 7.0 9.5 14.1 14.5 21.1 26.5 4 0.02 4.5 5.2 911 7 5 19.0 26.1 6 0.1 5.2 3.2 9.2 3.9* 16.2 19.5 7 0.2 5.1 2.5 8 3 3.7* 13.9 15.0 --- """""" 7 “““ 52. """ 7: """ """ 7 """ a; " 1 0.0 100 100 100 100 100 100 2 0.0005 95 86 99 88 85 -- 3 0.001 93 77 104 90 104 101 4 0.02 69 42 67 45 94 100 6 0.1 67 26 68 24 80 75 7 0.2 68 20 61 23 68 57 * Many secondary roots (3) = Age in days after germination. Table 9. Growth of Hormone Dusted Dent Corn on Filter Paper. Seeds treatud with indole butyric acid dust. Planted May 29. Germination May 30. Room tempe- rature 24-2700. Mean of 10 plants. Length of T0p (T) & Root (R) (11) (15) Concent. June 1 (2) June 2 (3) June 3(4) Jun 10 Jun 14 Hormone N0. p.p.m. T R T R T R T T cm cm cm cm cm cm cm cm 1 0 1.0 4.2 3.2 7.3 5.2 10.3 26.9 29.5 7 20 0.8 4.3 3.1 7.4 5.1 10.8 27.2 30.0 ------‘---’-------------------’-----‘-------------------- 20 80 102 97 101 96 116 101 102 12 120 80 55 100 69 87 51 -- -- 16 200 100 52 87 55 96 59 86 92 (2) = Age in days after germination. Table 10. Growth of Dusted Sweet Corn Seeds on Filter Paper. Seeds treated with indole butyric acid. Set June 30. Germination July 4, and July 5 for number 20. Room temperature 20-22°C. Mean of 10 plants. Length of T0p_(T) & R00t(R) Hormone _ Concentration July 6 (2) July 7 (3) July 8 (4) N0. p.p.m. T R T R T R C121 C271 CL". CM CM CM 1 O 2.8 7.0 8.1 11.8 12.1 16.2 4 2 2.5 7.1 8.3 12.3 10.5 14.6 7 20 3.0 6.9 8.7 8.6 11.6 12.8 16 200 2.6 2.6 8.6 3.2 11.8 6.6 20 2000 1.7 1.5 4.5 1.8 5.9 1.8 ' """"""""" :2 """ 5 """" c2 """ i """" 7 """ 32" l 0 100 100 100 100 100 100 4 2 89 101 102 104 87 90 7 20 107 99 105 73 96 79 16 200 93 37 106 27 98 41 20 2000 61 21 55 15 49 13 (2) = Age in days after germination. Table 11. Growth of Sweet Corn Seeds 0n Ensted Filter Paper. Germinated seedlings eXposed t0 dif- fused light (A) and darkness (B). Indole butyric acid (0.1 g.) dust on filter paper (250 cm2). Set July 28. Covered with bell-jar (A), and inverted pot (B). Ger- mination for (A) July 1, and July 2 for N0. 16 and 20; for (B) July 2, and July 3 for Number 16 and 20. mean of 10 plants. Hormone measurement (July 17) of Concentration sixteen day old seedlings Inter Coleo- No. mg/paper T0p Root node ptile cm % cm % cm cm (A) 1. 0.0 31.0 100 30.2 100 1.1 4.2 4 0.1 27.8 90 27.0 89 1.0 '4.5 7 1.0 30.2 97 17.8 59 1.2 4.3 16 10.0 16.0 84 17.1 57 1.2 4.2 20 100.0 5.8 19 1.0 3 1.3 2.8 Av. 112 .2 TB) 1 0.0 33.0 100 20.4 100 6.8 4.5 4 0.1 31.5 91 20.0 98 6.2 4.4 7 1.0 28.8 32 19.1 93 6.0 4.4 16 10. 0 29. 0 85 18. 1 88 7. 3 4. 3 20 100.0 15.9 44 0.5 3 __7_-_l .1115 Av. 6.7 4.4 Table 12. Growth of Dusted Buckwheat Seeds 0n Filter raper. Seeds treated with indole butyric acid dust. Set Aug. 15, in moist chambers in greenhouse. Temperature 23-270C. Humidity 99-100 percent. Germi- nation Aug. 17. Number 7 slightly poor in germination. Hormone T0p length. Root length No. p.p.m. Aug. 21 Aug.23 Aug.25 Aug.21 Aug.23 Aug.25 cm cm cm cm cm cm 1 0 6.7 8.7 11.0 15.6 16.7 20.7 3 0.15 7.1 9.8 11.6 16.2 17.6 21.3 5 30 6.5 8.8 10.3 14.6 15.1 15.8 6 300 6.2 8.8 11.5 7.0% 9.9*** 15.9# 7 3000 3.4 8.3 11.0 3.4** 8.6 10.3## ' """"""""" “““ 7: """ """ """ """ 7 1 0 100 100 100 100 100 100 3 0.15 106 102 105 104 105 103 5 30 95 101 94 94 91 77 6 300 93 101 105 45 54 77 7 3000 51 96 100 22 52 50 * Primary roots about 0.70m. 4easured longest sefiondary r00 t ** Primary root completed grwoth. " *** Some primary roots completed growth. # When primary root is short more secondary roots develOpe. ## Number of secondary roots 6 to 15. Table 13. Summary of EXperiment 1, Effect of Hormone Treatments on the Growth in Length of Seedlings. Hormone cmoncentration & per- Plants Table Age Part cent 01 growth based on check. 2 y. 20 .120 200 2000* Dent Corn 9 2 T -- 80 80 100 u 4 n -- 96 87 96 n 2 R -- 102 55 52 .u 4 n -- 115 51 59 Sweet Corn 10 2 T 89 107 -- 93 61 n 4 u 87 95 r- 98 49 g. :2 I}: :1.()']L SE95; ""' 35,7, 22:]. 4 " 90 79 -- 41 13 0.15 30 300 3000* T 106 95 93 51 .. 105 94 105 100 104 94 45 22 " 103 77 77 50 0.0005 0.001 0.02 0.1 0.2** Buckwheat 12 (ID->0)? m Dent Corn 8 3 T 95 93 69 67 68 u 5 N 99 104 67 68 61 " 8 n 85 104 9 4 80 68 " 3 R 86 77 42 26 20 " 5 " 88 90 45 24 23 0 . 1 1. 0 10 10 0*”- Sweet Corn Light 11 16 T 90 97 84 19 Dark n n n 91 82 85 44 l. II 0' Light 0! n R 89 59 57 3 Dark n n n _9 8 9 3 88 3 Hormone, indole butyric acid. Age in days after germination. T = top length. R.= root length. * pep-m0 2 ** hormone in 0.2 mg. on 120 cm paper Table 14. Some Preliminary Observation of Plants Growing in Hormone Treated Soil in the Greenhouse. Seeds planted after soaking 4 hours, June 18; sand was added on tap to the depth of 2.5 cm., after roots were out; pot covered with glass. Greenhouse temperature 27- 3200. Plants per pot, 5, but 10 for millet. Treatment duplicated. Date of Hormone emergence Date of Measure. Plants No. Conc. Root Tgp, Flgwering Sep. 17. P.P.m. June June July* T0p length Sweet 1 o 21 23 22 ' 55 cm. Corn 4 2 " " " 54 7 20 n N n 55 15 200 u n u 53 20** 2000 " " " 38 mg/oot Air dry wt. Buck- l 0 20 22 16 45.5g. wheat 4 0.2 " " 17 38.2 7 2 " " 20 41.0 16 20 u 23 22 21.2 2044* 200 u 29 -_ __ we/pot 4 Soy Bean 1 O -- 22 27 4 0.2 -- " 25 7 2 -_ u 25 16# 20 -- 28 26 20## 200 -- 3O -- me/pot uillet 1 O 20 22 4 0.2 " " 7 2 II II 16*# 20 n u _) 20 200 " " * Date tassel appeared on corn. ** Dwarfed at maturation. *** Rapid growth after flower formation. # Growth inhibited. ## Dead July 10. *,¥ Younger stage only inhibited. Table 15. Growth of Dusted Sweet Corn Seeds in the Greenhouse. Seeds with indole butyric acid dust. Planted Oct. 1, in 25 cm. pots. Germination began Oct. 7, but later in number 20 which was very poor. Temperature 20-2700. Mean of 10 plants. Hormone b‘ilk Height No. p.p.m. Oct. 21 Nov. 23 cm % cm % 1 0 Nov. 23 35.2 100 61.5 100 4 2 " 38.8 110 61.7 100 16 200 " 32.5 93 60.8 99 20 2000 24 30.7 87 59.6 98 . [I'll-'81:»... . .01.. n (I (I111. Tables 16, 17. Growth of Buckwheat in Dusted Soil in Greenhouse. Surface of the soil in pots (250m in dia.) was covered uniformely with the 200 g. air dry soil mixed with dust containing certain concentrations of hormone; seeds were on the surface of this soil, and then covered with dry sand, to a depth of 2cm, Oct. 1. Temperature 27°“. Humidity 66-84%. Tops, ovendried at 85°C on Nov. 19. Jean of 10 plants per pot. Hormone Height and date of measurement. go. mg/pot Oct. 212 Nov. 19 7 Oven dry wt. POT A cm 75 cm ;e g 7;, 1 0.0 13.5 100 59.2 100 3.89 100 2 0.1 18.5 101 55.3 94 3.34 86 3 1.0 ‘ 18.2 99 59.0 100 3.54 91 4 10.0 17.0 92 54.1 91 3.06 79 5 100.0 11.7 63 50.5 85 2.11 54 R 0.3 13.4 99 56.8 90 3.64 94 a‘ 3.0 12.1 66 52.1 88 3.22 83 1366'; """""""""""""""""""""" l 0.0 19.2 100 51.5 100 3.46 100 2 0.1 18.7 98 51.7 100 3.52 102 3 1.0 15.4 96 52.1 101 3.48 100 4 10.0 17.2 90 45.3 89 3.02 88 5 100.0 11.1 58 44.7 87 1.63 49 R 0.3 18.8 98 51.5 100 2.58 75 R 66 49.8 97 2.54 74 ' 3.0 12.6 Dust No. 1-5 = indole butyric acid n = Rootone 0.3 g. per pot ‘x' = II 300 E. per 1301;. Ceraination began Oct. 6, except for N0. 5 and R which germinated on Oct. 7. “J Table 18. Growth of Buckwneat on Dusted Soil. Sumnary from tables 16 and 17. Average percen- tages in height based on check as 100 percent. Hormone Height - Dry Wt. .I_0 . mg) 1:- ct Oct . 21 ‘1‘! 0V . 1S: 0V; 1L )0 70 2'0 1 0.0 100 100 100 2 0.1 99 97 94 3 1.0 98 100! 95 4 10.0 91 9O 84 5 100.0 61 86 51 R 0. 3 99 93 35 R' 3. 0 66 9 3 79 Table 19. Growth of Dustcd Soy Sean Seeds in Greenhouse. Eaphthalene acetic acid mixed with dust. Seeds planted in 250m pot, Oct. 1. Germination Oct. 8 and Oct. 16, for number 5 (Ho. 5). Temperature of greenhouse 20- 2700. Harvested Nov. 10. Boots carefully washe ; roots (.1: and tops dried toEether in oven. Kean of 10 plants. Hormone height and date Oven dry wt. K0. p.§:7. Oct. 21] Nov. 10 fl» Nov. 10 J c m yo c m 79 g 70 l O 19.5 100 56 100 3.38 100 3 10 20.7 106 58 103 3.48 103 4 20 16.1 83 56 100 2.84 84 5 2000 4.6 23 * -- 1.06 31 * EEad Nov. 1. Table 20. Growth of Rootone Treated Tomato Plants in Greenhouse. Rootone powder (0.2 gram) per plant placed around roots. Flowers appeared (50 percent) Nov. 3, in check plants, and Dec. 15 in rootone treated plants. Temperature 20-2703. Height measurement to tip of leaves. Kean of 6 plants. Date Check Rootone (0.2g/p1ant) cm % cm 9% October 4 15.6 100 16.7 107 " 21 48.3 100 36.8 76 rev. 29 135.3 100 93.2 70 .Table 21. Growth of Rootone rusted Wheat and lillet Seeds in Greenhouse. Lean of 40 plants. Temperature 20-2700. #- Rootone Late of Heig TOp of plant Ho. g/lOO Emergence Oct.2l ov. dry weight Begin SOw on S cm % g % Wheat: 1 0.0 Oct.6 Oct.7 33.3 100 42.6 100 6.11 100 2 0.3 " 35.1 105 45.3 106 6.28 102 3 3.0 * 33.5 100 44.1 104 6.08 99 Iillet: 1 0.0 Oct.7 Oct.9 11.1 100 26.9 100 4.62 100 2 0.3 " 10.8 98 30.1 112 4.70 102 3 3.0 * 10.9 99 27.9 100 1.96 43 * Germination, poor or a/lOO = grams of Rootone per 100 grams of seeds. nearly one day behind. Table 22. Calculation for the Significance of Rootone Treatment of Tiller Production of Wheat. llor {Lone n N nN mil 5% conc. Number of Number of £1100 tillers* plants g. 0 1 20 2O " 2 12 24 " 3 8 24 08 68 100 0.3 1 13 13 " 2 17 34 n 3 10 o 77 77 103 3.0 l 14 i4 " 2 2O 40 u 3 6 _1_8 72 72 99 * Total number of tillers per plants Number of plants for each group 40. nN : Total number of tillers in 40 plants. g/lOO = Grams of Rootone per hundred grams of seeds. Table 23. Growth of DEnt Corn Dusted Seeds in the Field. Hormone , indole butyric acid. Planted June 3. Lmergence June 9. measurement to the t0p of flo- wering tassel on Aus. 8. Mean of 20 plants. Hormone ‘ lasselsfl Flower Height No. p_.p_.m. in full cm % 1 0 Aug. 3 Aug. 8 212 O 4 2 " " 215 + 1 7 20 " " 215 + 1 10 3o " " 218 + 3 12 120 ‘ " " 206 - 3 14 ' 160 " " . 207 - 2 16 200 '" " 195* -8 « Statistically significant. # Tassel appeared in 80 % of plants. % Bercentage difference from check. Table 24. Growth of SWeet Corn Dusted Seeds in the Field. Hormone, indole butyric acid. Planted July 5. Emergence from ground July 10 except for number 20 which was one day behind. Tassel appeared in 80% of plants Aug. 8. Floter in full Aug. 18. Height measurement to the toe of flowering tassel, Aug. 20. Mean of 20 plants. Hormone Height No. p.p.m. July 18 Aug. 8 Aug. 20% 1 O 21.7 0 75.7 0 107.2 0 4 2 21.2 -2 79.8» +5 106.3 -1 7 '20 21.5 -1 74.1 -2 105.6 -2 16 200 21.5 -1 68.t* -10 106.5 -1 20 2000 17.6* -12 71.4% -7 109.3 +1 * Statistically significant. *" " highly significant. # Variation within the row was statistically significant. Table 25. Growth of Dusted Buckwheat Seeds in the Field. Hoemone, indole butyric acid. Planted June 30. Emergence from ground Aug. 6, except number 7 which was one day delayed. Flower start Aug. 23. mean of 20 plants. Hormone Height No. p.p.m. Aug. 8 Oct. 3 Dry wt. cm % cm % 8 % l 0.0 5.5 O 35.2 0 192.6 0 2 0.075 5.4 -2 82.8 -3 185.1 -3 3 0.15 5.8 +5 90.0 +5 200.2 +4 4 3.0 5.3 +4 90.2 +5 202.; +4 5 30.0 5.3 -4 88.1 -3 185.5 -4 6 300.0 5.4 -2 81.5 -5 195.6 -4 7 3000.0 5.1% -7 86.1 +1 194.4 +1 * elongation retarded significantly. Table 26. heasurements of height of Plants Grown from Dusted Sweet Corn Seeds in the Field (Aug.3,Table 24). Hormone conc. and measurements Lines 1 4 7 16 20 cm cm cm cm cm 1 69 86 72 53 59 3 70 38 73 68 68 5 33 91 74 72 53 7 87 90 80 72 63 9 ' 85 9O 80 73 73 ll 82 89 78 55 7O 13 81 82 77 54 73 15 71 67 77 71 82 17 72 83 74 64 84 19 75 73 73 7o 78 21 77 74 72 7o 81 23 77 76 71 64 68 25 80 75 76 66 65 27 60 65 82 72 81 29 75 84 78 67 51 31 74 88 65 58 78 33 64 69 67 69 69 35 67 7O 67 68 68 37 75 71 89 76 62 39 64 87 58 68 72_g_ AVG. 75.7 79.9* 74.1 68.0** 714% * Statisticaly significant. ** " highly significant. —rm Table 27. Plants Soy bean '31"!- h e a t 351 .1th Pea Observation on Different Seedlings Treated with Hormone Enst. Seeds planted in the field, Aug. 20. Hormone Germination Remarks concent. Start 80% ‘ p.p.m. 0 Aug.27 Aug.28 2 u n 10 " " 2o " " 2000 " 30 Germination poor. Check 24 26 R " " Tillering appeared Sep. 15. Cheek H N R u _ 11 Check 27 29 R II N Kidney bean Cheek 26 28 I‘3t3tdace \ R u u Check 27 30 R N 80y bean seeds were dusted with napthalene acetic acid mil-ted with talc. R : Seeds dusted with Rootone and exce es shaken off. Table 28. Summary of Exp. 11. Percentage differences based on check Dent Plant corn Sweet corn Buckwheat Table _—23_- 24 28 Age gdays) 3o 8 3o 40 2 40 4o P-pmm. ii: 77 75 75 $5 76 % 2 £0.15 +1 -2 +5 -1 -4 +5 +4 20—30 +1 -1 a2 -2 -4 +3 -4 200-300 -8* -1 -10** -1 -2 -5 -4 2000-3000 --- -l2** -7* +1 -7* +1 +1 * Significant. ** Highly significant. . i . 1.3%..11‘ neatlvr‘flylr.’ Table 29. Degrees and Direction of Curvatures of Zea ColeOptile at Low Humidity. Degree Number Of Hormone oi‘cur— tip empty Plants concent. vature coleoptiles % 6 0 0 Normal " 0.1 +68 Normal " 3 ~38 4 (0.5cm) " 10 -35 5 (0.40m) Place,. dark room. Cultures covered with pasteboard box. Temperature of room 21-2500. Humidity low (22-42%). Coleoptiles were used when length was 2 to 3 cm, and hormone application in the form of lanolin paste at the middle. Measurements taken 10 hours after application. Date of observation Sept. 24 (12:10 A.M). Degree of curvature determined by means of shadOWgraphs on bromide paper. Positive curva- tures are indicated by (+) and negative by (-). Hormone used naphthalene acetic acid. l E..II.|ILH:H~ . ail - . 34...... ' lo ... "III! I - null 0.. ‘ Table 30. Effects of Naphthalene Acetic Acid in the Form of Lanolin Paste Applied at Various Points of Zea Seedlings. Hours after application No. Location 8 hrs l2hrs 20hrs 34 hrs 1 Check . Plumule appeared 2 0.5cm from tip ++ G ++* + bmp. +- 3 Mid. coleop. ++ P ++* + Emp +- 4 Node ++ P ++ ++ ++ 5 Mid. internode ++ ++ + +-+ Experiment conducted in laboratory in porous pot damp chambers. Room temperature 23-2700. Humidity of sefide ling environment high. Hormone apolied Oct. 12 (10 3.x). Length of coleOptile 2 to 3 cm. Number of seed— lings used 40. Positive curvature indicated as (+), negative curvature as (-), and recovery indicated (+-). P. means sharp curvature at point of applicatioa. Empty coleoptiles indicated by Emn. Two negative curvatures recorded here also(*). 5-: 1 .IIWLH. a . aw Table 31. The Effects of Quantity of Hormone and Age of ColeOptiles on Direction of Curvature. Group. A B C (D Av. Length ColeOp. at start (cm) 3.5 3.0 2.7 2.0 Quantity of paste L s L s ' L s L 5 Curve after 12 hrs 0 0 - t+ - + - + - + - + — + Hormone used, 3 percent naphthalene acetic acid in lano- lin paste. Room temperature 23-2700. Number of plants 32. Hormone paste applied at middle of coleoptile. Positive curvature (+), and negative (-). See plate 23. Table 32. The Effects of Various Concentration of Hormone in Lanolin Paste on Zea ColeOptile. No. ‘ 1 2 3 4 5 5 h BothSide fife—Emma Percent Hormone 0 0.1 3.0 10 0.1 10 Cuzflfafter 12 hrs 0 ++ ++ ++ 0 ++ " " 24 " O +- +- + 0 ++ Number of plants and conditions of the experiment similar to that recorded in table 30. Place of hormone application 0.5 cm below tip. Plus signs indicate medium or strong curvature. Plus and minus signs togedier indicate reco- very. See plate 27. Table 33. The Effects 0: Hormone in Lanolin Paste on Ibcapitated Zea ColeOptiles. Hormone Older coleop. Younger coleOp. Hg. Conc. Hggin 24 hpg Begin. 12 hrs 24 hrs cm on cm cm 1 0.0 3.3 3.7’ 1.3 2.5 O 3.5 0 2 0.1 2.8 3.2 1.4 3.3 ++ 3.3 +- 3.0 2.9 3.4 1.6 2.8 ++ 3.5 +- 4 10.0 3.3 3.9 1.7 2.7 ++ 3.5 +-+ 5* 10.0 2.8 3.4 1.7 2.7 ++ 3.7 ++ 6H 0.1 2.7 3.4 1.5 2.6 o 3.4 o Plants and conditions of the eXperiment similar to that shown in table 30. * Lanolin paste applied at internode. ** Lanolin paste applied symmetrically on decapitated coleoptiles. Plus indicates positive, and minus negative curvatures. No effects indicated by O, and recovery by (+-). . ...L' ._c._'_. a Table 34. Rooting Effects on Cuttings Through the Use of Rootone. Temperature of the greenhouse 25-3200. Placed in prepagating bench Oct. 4. Examined Nov. 15. Number of plants_fl Total plants Plants Rootone A B C D Alive Used Chrysanthemum Check. 26 6 7 O 39 40-50 " Tal-S 20 o 8 4 32 " " R 21 16 6 o 45 " " a-e 16 12 8 2 38 " Hesembryanthe- Check. 12 12 8 4 36 30-40 " mum R 15* 15 o 0 3o “ Hydrangea Check 21 8 6 o 35 4o " R 20 12 6 o 38 " Kleinia Stem Check 12 6 6 8 32 30-40 " " R 8 10 5 7 3o " " Leaf Check 6 7 7 9 29 " n n a ' 5 9 6 8 28 ' " Crassula Stem .Qheck 10 12 3 O 23 25 n N R 9 14 3 o 26 " " Leaf Check 12 8 O O 20 " n " R, 11. 10 o o 21 " Azalea Check 0 O 0 0 2O 40 u R o O O O 10 " * Root develOpment significant. Class A = Many roots well develOped. B = Medium develOpement of roots, more than 5, rather short. C = Few roots, 1-5. D = No roots. R = Rootone. 2+3 2 0.2g. Rootone for each row (35cm) in flat. TQl-S = 0.2g. talc powder without hormone for 35cm rows. Table 35. Rooting Effects of Cuttings Due to Rootone Calculated in Percentages Based on Total Cuttings Alive. Data from Table 34. Percentage of plant rooting Plants Plants Rootone A B C D alive 7 Z 73 75 % Chrysanthemum Check 66 15 18 0 100 " Tal-S 63 o 25 13 100 " R 47 4o 13 o 100 " R-S 42 31. 21 5 100 Mesembryanthe- Check 33 33 22 11 100 mum " R 40 50 O O 100 Hydrangea Check 60 23 17 0 100 " R 53 32 16 O 100 Kleinia Stem Check 37 l9 19 25 100 " " R 27 33 17 23 100 “ Leaf 9heck 21 24 24‘ 31 100 8 " R 18 32 22 29 100 Crassula Stem Check 44 52 13 O 100 " " R 35 54 12 o 100 5 Leaf Check 50 40 o o 100 " " R 53 48 o o 100 Azalea Check 0 O O O 100 " R O 0 O 0 100 R = Rootone 8-8 = Rootone on soil. Tal-S = Talc without hormone. Table 36. Lffects of Hormone on Corn Seedlings Previously Treated with Colchicine. 20-2500. Plants in petri dish moist chambers. Laboratory temperatures Treatment & Observation Check Hormone treated. Seeds germinated Nov. 1 Nov. 1 Colchicine 0.2% sol. Nov. 3 Nov. 3 Washing (tap water) 4 hrs 4 hrs Hormone (0.04% naphthalene) -- Nov.4 (20hrs) Washing -- 4 hrs Petri dish culture Nov. Nov. 5 Lateral roots appeared Nov. Nov. 7 Lateral rooted plants 2 12 n " " 7; 10 60 Table 37. Solution Mixtures With and Without Catalyst. No. 0 1 2 3 4 (0.5m) (0.5m) (0.04%) (0.4%) (0.04%) Blank F6013 + HCl 1 + Naph. 1 d- Naph. Napha cc cc cc cc cc F6013+ HCl 0 10 10 10 0 Naph. 0.04% o o 5 o 5 " 0.4% O 0 o 5 o HOH .13 .3 .2 __9. .19. Total cc. 15 15 15 15 15 Table 38. Titration for Decomposition H202. Mean of four titrations with 0.1 N KHnOA; room temp. 25°C. No. 0 _;L 2 3 4 Minutes cc cc cc cc cc 5 15.10 14.50 15.20 16.50 15.85 15 15.15 12.50 14.00 1 6.65 15.90 30 15.10 10.20 12.20 16.70 15.88 45 15.10 7.90 10.95 16.70 15.82 60 16.20 6.35 10.00 16.69 15.80 90 15.20 4.20 8.10 16.65 15.86 120 15.17 2.90 6.60 16.50 15.85 180 15.10 1.25 4.25 16.50 15.84 240 15.17 0.60 2.85 16.43 15.86 Av. 15.14 15.85 Table 38. Titration for Decomposition H202. mean of four titrations with 0.1 N KMn04. Room temperature 25°C. No. 0 1 2 3 4 Minutes cc cc cc cc cc 5 15.10 14.50 15.20 16.70 15.85 15 15.15 12.50 14.00 16.65 15.90 30 15.10 10.20 12.20 16.70 15.88 45 15.10 7.90 10.95 16.70 15.82 60 15.20 6.35 10.00 16.69 15.30 90 15.20 4.20 8.10 16.65 15.86 120 15.17 2.90 6.60 16.50 15.85 130 15.10 1.25 4.25 16.50 15.34 240 15.17 0.60 2.85 16.43 15.86 Av. 71—5714 .1578? u, it! ‘85» I Eq‘ . u Table 39. Reaction Velocity Constant for Decomposition H202. No. l 2 00 15.14 15.85x10"4 minutes K K 5 87.7 78.5 15 84 85.5 30 108 86.4 45 131 82.0 60 172 77.3 90 - 132 73.0 120 133 . 74.0 180 135 73.4 240 139 70.0 Av. 107x10‘4 78x10 e Table 40. Chemicals Reported as Hormone or a Hormone-like in Activity. ‘1 Chemicals Stimulate on Reported by Auxin a Avena test Koogl (1935) Auxin b " " neteroduxin " " l-Iethyl—3-acetic acid " 8-3-indole prooionic acid " -Pyruvic acid 3-Indole pyruvic acid Avena test Phenyl acetic acid ll ‘- Phgyfl pr0pionic acid Isatinic acid " Pea test a some Avena test .3- (b)-Naphthalene acetic u 3—Indole acetic acid 3-3-Indole butyric acid " Indole pr0pi0nic acid Phenyl acetic acid Fluorene acetic acid Anthracene acetic acid " b-L-laphthyl acetonitrile " Ethylenegas Root, Stem. Glutathion Root Sulphanilamide Root 0" -Pheny 1 buty ric ac id Ascorbic acid (vit.C) tea seedling Haagen Suit 8: Went (19 35) Zimmerman & \fiilcoxon (1935) H Croker, Zimmermana hitchcock (1935) Grace, N.H. (1937) Hausen, S.V. (1935) 1-Coumary1 acetic acid(Root initiatebut Thimann,K.V.(l935) not Avendcoleoptufi) GRAPE-IS I 6017011731 r Fig; 2_‘ D NT §CORN l . rthOw-nomiotm‘ are" 011 FlELTER ...... ........ ......... ........ ......... Ii‘S‘wfw'" .. 7 '1‘. 3 days 0131 1! W . ...... ........ ..... > ...... ......... ......... y a >>>>>> ......... ........ ......... ......... ........ ....... ....... .__7.__ +-.. ..... ......... ......... ....... ........ ....... ------- ......... ........ ........ ......... ....... ........ ......... ......... ......... .......... ....... ........ ..... ......... ...... ........ ........ .......... ......... ...... ........ ,.....,L.L ......... ......... ........ ......... l Fig.23 ..... 2354 ‘ , ' ' T461642; . ..,.J im¥CCUCENTRAEIONLaoauonm§Ahp.'*-*“ AGE cRchH‘BUCKwHEAm= ON FILTER I 0 l 8 . >- ; .i l > i 7,. l ‘\ H t ..... ...... ...... 'R8——« ....... ........ 'iTopy ~~n+ LLLLLL ..... ........ ........ ------- ' 4442—2664 "3' ........ ........ ....... ....... ........ ....... ....... ........ ..... ......... -------- ......... ........ ......... ........ ........ ........ ........ ...... ......... ..... ....... ....... ......... ......... ......... ..... ...... ............. ....... ................. a? 1112?. 4 ......... e 75:43‘ h“ foi,:*-'§ =“~'rl- ‘ [2' -»' "r:" ‘3 1'. .: ff£ECOMPOSITION?HQOQTITRATEEIWITH 0:1“MMJin04~~e+%~+~4=efie~~ 6%" 6' .' .. -_ ._a . : i *:.t? "f " 1 ' _.' {Table 38) ..... 3Blan4";” at; i FeCit+iH¢lii‘ ‘!* 111 ~1La.p11..L04‘0_4%1” (1) 11 Naph. (O 4") Naph. (010W ) ”41704 701-40 H N M 3! N ..-.~1 .-L_—v +— _—.- _.__. ...-_4 1.”... v. \ .4. PLATES 1-32 Plate 1 LARGE MOIST CHAMBER AND PAN FOR GERMINATIO” Right: Moist chamber (100x70x90cm). Left: Pan (SOxSOxIOcm) containing filter paper on glass plates for germination of seeds. Plate 2 POFDU: POT T~'OI°'7‘ ”HAMRPR Plate 3 SEEDLINGS FROM HORMONE IIJSTEB SWEET CORN SEEDS GROWING ON FILTER PAPER (July S-Table 10) 0 ~04 “gt Indole acetic acid dust. G l = Check G 4 = 2 p.p.m.* G 7: O G 16 = 200 G 20 = 2000 * p.p.m. = Parts hormone to million parts of seeds. Plate 4 SEEDLINGS FROM HORMONE BUSTED SWEET CORN SEEDS GROWING ON FILTER PAPER The same seedlings as in plate 3, but older. Brimary root growth inhibited but root hairs develOping. Plate 5 SEEDLINGS FROM HORMONE BUSTED SWEET CORN SEEDS GROWING ON FILTER PIPER (July 8-Table 10) The same seedlings as in plate 3, but older. Secondary roots are appearing. Plate 6 SWEET CORN SEEDLINGS GROWING ON HORMONE DJSTED FILIPER PAPER UNDER DIFFUSED LIGHT. (July l7-Table 11) t “I Indole butyric acid dust spread on filter paper (r = 9cm) 1 = Check 4 = 0.1 mg/paper 7 = 1.0 10 = 10.0 20 = 100.0 NI Plate 7 SWEET CORN SEEDLINGS GROWING ON HOEL‘ONE DISTED FILTER PAPER IN DARKNESS. (July l7-Table ll) Indole butyric acid dust spread on filter paper (r = 9cm) 1 = Check 4 = 0.1 mg/paper 7 = . 10 = 10.0 20 = 100.0 Plate 8 SEEDLINGS FROM HORMONE BUSTED BUCKWHE AT SEE DS GROWING ON FILTER PAPER (Aug. lS-Table 12) F\{"\ ‘( ‘ Y7/:"qr'(‘v"7$‘6P-r'§¢‘f‘fr*w‘¢h 0; HHS I M W \' (“77(“3‘ c‘ W (WW 1 ‘ WWW Wm H Indole butyric acid dust spread on filter paper (r = 90m) 1 = Check 3 = 0.15 p.p.m. S = 30.00 6 = 300.00 7 = 3000.00 l Plate 9 SEEDLJNGS FROM HOZMONE BUSTED BUCKWHEAT SEEDS GROWING ON FILmER PAPER (Aug. l9-Table 12) Same seedlings as in plate 8 but older. Indole butyric acid dust spread on filter paper (r = 9cm). 1 = Check. 3 = 0015 p.p.m. 5 = 30-00 6 = 300.00 7 = 3000.00 Plate 10 SEEDLINGS FROM BORE-LONE IIJSI‘ED BUCKWHEAT SEEDS GROWING ON FILEER PflPER (Aug. 2l-Table 12) D‘ame seedlings as those in plate 9 but older. Indole butyric acid dust spread op filter paper (r = 9cm). 1 = Check 3 = 0.15 p.p.m. 5 = 30.00 6 = 300.00 7 = 3000.00 Plate 11 SEEDLINGS FROM HORMONE BUSTED BUCKWHEAT SEEDS GROWING 0N FILTER PAPER (Aug. 23-Table 12) Same seedlings as those in plate 9 but older. Indole butyric acid dust spread on filter paper (r = 90m). 1 = Check. 3 = 0.15 p.p.m. 5 = 30.00 6 = 300.00 7 = 3000.00 Plate 12 SEEDLINGS FROM HORMONE DJSTED BUCKET-{EAT SEEDS GROWING ON FILTER PAPER (Aug. 23-Table 12) Seedlings 6 and 7 from plate 11. Showing primary roots. Left: 300 p.p.m. Rightz3000 p.p.m. Plate 13 SEEDLINGS FROM HORMONE BUSTED BUCKWHEAT SEEDS GROWING 0N FILQER PAPER (Aug. 25-Table 12) Seedlings 1 and 7 from plate 11. Left: Check Right: 3000 p.p.m. Plate 14 SEEDLINGS FROK HORMONE BUSTED SWEET CORN SEEDS GROWING IN THE GREENHOUSE (Nov. S-Table 15) Indole butyric acid. Left: Check Middle: 2 p.p.m. Right: 20000 p.p.m. Plate 15 BUCKWHEAT GROWING IN HORMONE DUSTED SOIL. (Nov. 6-Table 16) Left: Check Middle: Indole butyric acid 100 mg/pot. Right: Rootone (0.1% naphthalene acetic acid) 3g/pot. Plate 16 SOY BEAN PLA‘JTS GROWN FROM HORMONE IIISTED SEEDS GROWING IN THE GREENHOUSE (Nov. S-Table 19) Left: Check. Middle: 20 p.p.m. Right: 2000 p.p.m. All seedlings in right hand pot dead NOV. 1].. Plate 17 ROOTONE TREATED TOMATO SEEDLINGS GROWING IN POTS IN THE GREENHOUSE (Nov. S-Table 20) Left: Check. hight: 0.2g. Rootone spread on soil around roots. Plate 18 ROOT SYSTEM OF MILLET GROWN FROM ROOTONE TREATED SEEDS (Nov. 8-Tab1e 22) Left: Check. Right: 3g. Rootone/lOOg. seeds. Plate 19 ROOT SYSTEM OF SWEET CORN GROWN IN THE FBLD (Oct. 6-Tab1e 24) Showing the point of attachment of the seed. Croun roots have developed, and the primary root has completed its deve- lopment. A = Check. B = Indole butyric acid, 2000 p.p.m. Plate 20 ROOT SYSTEM OF DENT CORN GROWN IN THE FIELD (Oct. 6-Taole 23) Shoning seed attachment from which primary roots develOped. A = Check. B = Roots of plant grown from seed dusted with indole butyric acid (200 p.p.m.) Plate 21 AERIAL ROOTS PRODUCED ON AUCKWHEAT STEMS AND PETIOLES BY APPLICATION OF HORMONE ‘ IN LANOLIN PASTE (NOV. ZO‘EXPQ lV-A) Hormone in lanolin paste injected, Nov. 11. Plants growing in 25cm. pots in the green- house. Temperature 20-2700. A,= Check, lanolin paste only. B = 0.L% Naphthalene acetic acid in lanolin paste. C = 10% Naphthalene acetic acid in lanolin paste. Plate 22 EFFECTS OF HORMONE IN LANOLIN PASTE ON TOMATO JPLANTS 'Right: Inhibition of terminal bud through treatment with 0.L% naphthalene acetic acid in lanolin paste. Lateral shoot growth increased. Left: Tip untreated, but axial treatment with the same hormone in lanoun paste. Lateral shoot growth was retarded. Plate 2} EFFECTS OF WANTITY OF LANOLIN PASTr APPLIED AND THE AGE OF ZEA COLEOPTIIES (Table 23) Three percent naphthalene acetic acid in lanolin paste applied at points indicated by arrows. Older coleoptiles at right. Right hand coleOptile of each group received the heavier application. Photograph taken 12 hours after application of the paste. Plate 24 THE EFFECTS OF VARIOUS CONCENTRATION OF HORMONE IN RLA‘IOLIN PASTE ON ZEA COLEOPT IIES. (Table 32) Photograph taken 20 hourcafter appli- cation of naphthalene acetic acid paste. 1 = Check. 2 = 0.1% 3=3% 4 = 10% 5 = 0.1% (applied on both side) 6 = 10% (applied on internode) Plate 25 THE EFFECTS OF HORMONE IN LANOLIN PAST; 0N DECAPITATED ZEA COLEOPTILES (Table 33) Photograph taken 24 hours after application of naphthalene acetic acid paste. No. l 2 3 4 5 Conc. paste, 0 0.1 3 10 10 % Group on left are older coleoptiles. Group on right are younger coleoptiles. T283195 5881122 5’ ZRBoEEE €83 $28€3i~2nt8§en at later stage and most of coleOptiles recoverd from bendings, yet bendings are seen on younger group (right); No. 5 were treated on inter-nod. Plate 26 THE EFFECTS OF ROOTONE 0N ROOTING 0F CUTTINGS 0F CHRYSANTHEMUM (Table 34) Left: Check. Middle: Cut stem treated with Rootone. Right: Rootone applied to soil, about the cut end of stem. Classification: A - Many roots well develOped. B = Medium developed roots, 5 to 10, short. 0 = Few roots, 1 to 5. Plate 27 THE EFFECES 0F ROOTONE 0N ROOTING 0F MESEMBRXANTHEMUM ANIJKLEINIA CUTTINGS. (Table 34) .. Many roots were developed. Medium developed roots but short, 5 to 10 in number. Few roots poorly develOped and 1 to 5 in number. Wfl> 0 ll Mesembryanthemum, check. w Rootone treated. Kleinia repens, stem Rootone treated. n u leaf Rootone treated. cactrm II II II n Plate 28 THE EFFECTS OF ROOTONE on THE ROOTING 0F HIDRANGEA AND CRASSULA commas. (Table 34) Tap row: Hydrangea cuttings. Middle row: Crassula leaf. Bottom row: Crassula stem. A.= Many roots well develOped. B = Medium develOped roots, but short and 5 to 10 in number. 0 = Few roots, poorly developed and less than 5 in number. Plate 29 SEEDLINGS FROM COLCHICIN TTBEATED BENT CORN SEEDS RETREATED WITH HORMONE SOLUTION (Table 36) All four seedlings treated with 0.1% col- chicine at the beginning. Later the two on the left given an additional treatment of naphthalene acetic acid.(0.04%). The two on the right not treated with naphth- alene acetic acid. Plate 30 DENT CORN SEEDLJNGS GROWING IN BOXES WITH ONE SIDE PROVIDED WITH GLASS ‘TOp: Check. Bottom: Seeds dusted with indole butyric acid (2000 p.p.m.) Left side of each glass face was covered with black paper to exclude light. Right side remained uncovered. Planted Nov. 3. Greenhouse temperature 20-2700 0 Plate 31 DENT CORN SEEDLJNGS GROWING IN BOXES WITH ONE; SIDE PROVIDED WITH GLASS. (Nov. 10-Exp. V11) Dame cultures as in plate 30 but seedlings are 3 days older. TOp: Check. Bottom: Seeds were dusted with indole acetic acid in talc. Left side of each glass face was covered to exclude light. Right side left uncovered. Plate 32 DENT CORN SEEDLINGS GROWING IN BOXES WITH ONE SIDE PROVIDED WITH GLASS. (Oct. 30-Exp. v11) Left box: Sand only. Right box: Sandy loam. Left face of each culture covered; right face left uncovered. For each group plants (10 plants) Left 5 plants: Check. Right 5 plants: Indole butyric acid (2000 p.p.m.) Planted Oct. 10. Greenhouse temperature 22-26°C. - “‘5'“ ROOM USE um \ui759m Sept-u. pi:- .' "6" 9 fig 501319 '41 -. W on“ IES MMMM 8