THE ACCUMULATION of SOME MIN Ok ELEMENTS in COEN PLANTS
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THE ACCUMULATION OF SOME MIN Ok ELEMENTS IN COEN PLANTS GROWN AT DIFFERENT LEVELS OF HYDROGEN-ION CONCENTRATION Dissertation Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By YUAN-PIN CHOU The Ohio State University 1954 Approved byt ACKNOWLEDGMENTS The author sincerely appreciates the help and guidance obtained from Dr. G. W. Volk, Chairman of the Department of Agronomy, Ohio State University, who was instrumental in admitting the author to do graduate work in the Agronomy Department and in arranging the fellowship grant for this study. Deep appreciation is due Dr. J. D. Sayre, Plant Physiologist, U. S. D. A., under whose supervision this study was carried out. Appreciation is also expressed to the Ohio Agricultural Experiment Station for the various aids that made this work possible. i TABLE OF CONTENTS Page Introduction 1 Review of Literature 4 Experinental Methods 10 Method of Growing Corn 10 Method of Maintaining Constant pH 19 Radioactive Isotopes Used 21 Log of the Experiment 23 Method of Sampling and Autoradiographing 2tj Procedure of Analysis 28 Experimental Results and Discussion 35 Experiment in 1951* Gravel Cultures 35 Experiment in 19521 Gravel and Solution Cultures 39 Experiment in 1953 61 Results from C o m Grown in Soils 61 Results of the Corn Grown at High and Low Concentrations of Minor Elements 70 General Discussion 76 Summary 84 Literature Cited 87 Appendix 93 Autobiography 116 ii THE ACCUMULATION OF SOME MINOR ELEMENTS IN CORN PLANTS GROWN AT DIFFERENT LEVELS OF HYDROGEN -1 ON CONCENTRATION INTRODUCTION One hundred years ago, Carl Sprengel (59), a German agricul turalist, evaluated the ten mineral elements generally occurring in plant ash and concluded that they were all essential tor the growth of crop plants. In 1917, Robinson et al. (48) analysed a large number of plants from different regions and reported that 38 elements, mostly minor elements, were found in the plants. In 1950, Sayre (51), using radioactive antimony, wolfram, and iridium (which were not reported by Robinson ejt al.) in a nutrient solution for gravel culture, found that these three elements were all absorbed and accumulated in corn leaves. Jacobson at al. (27) found that a new chemical element, plutonium, could also be absorbed and translocated by plants. Although plants may absorb any mineral element which is available in the medium, not all of the absorbed minerals are necessary for growth of plants. For instance, chlorine is universally present in plants, but it is not an essential element. Sodium is another element which is not essential for all plants, yet is always present in plant ash. Most of any mineral element absorbed by a plant, whether it is essential for plant growth or not, will remain in the plant, though some of it may move out. The distribution of the accumulation of each mineral element is different in different plants and in different parts of the same plant. 1 The location, pattern, and amount of accumulation of the mineral elements may shed light on the study of adequate fertilisation and of nutritional value, as well as of the diagnosis of the disease symptoms caused by the physiological disturbance. Since the begiflfling of the twentieth century, much work has been done on the subject of minor elements. Host of these studies involved attempts to prove that the elements were essential for plant growth. Studies of accumulation and distribution of minor elements in a whole plant have not been numerous, probably at least partially because of the laclc of quick and accurate means of determining them. Since the Atomic Energy Commission released the products of the atomic pile for research purposes, these isotopes have been used as tracers to study the accumulation of certain elements in plants. However, this method is limited to those elements which have isotopes available at the present time. The second best tool for studying the accumulation of minor elements in plants is probably the spectrograph, which is fairly accurate within the range of concentrations at which the mineral elements are generally found in plants. For this work, both radioactive isotopes and spectrographic techniques have been employed. This study represents three years work (1951-1953) at the Ohio Agricultural Experiment btation, Wooster, Ohio, on the accumulation and distribution of some minor elements in corn plants in relation to the hydrogen-ion concentration of the medium. In 1951, a preliminary experiment was performed using Zn®® in gravel cultures of corn. In 1952, Zn®^, Co®®, F e ^ , and brS® were used in both gravel and solution 2 cultures. The latter method was employed in order to provide uniform 6S 60 hydrogen-ion concentration around the roots. In 1953, Zn and Co were used in soil cultures in pots. In addition, corn samples from the field, from pot cultures, and from gravel cultures supplied with high and low concentrations of minor elements were analysed spectrographi- cally in order to get more information about the minor elements which were not available as radioactive isotopes. In all cases, radioactive materials were introduced at the tasselim; stage, and the corn harvested about one month later. No attempt was made to study the total amount of each element in the plant at different stages of growth. The relative concentration (micrograms of element per gram of dry plant tissue) was used to compare the accumulation of an element at different pH levels in each part of the corn plant. 3 REVIEW OF LITERATURE The Recognition of Soae Minor Elements as Essential for Plant Growth Sprengel (59) vas probably the first to maintain that the elements generally found in plant ash, such as potassiun sodium, calcium, magnesium, iron, aluminum, manganese, sulfur, and chlorine, were essential for the groirth of plants* He further remarked that iron, bromine, iodine, lithium, and copper, though occurring in minute amounts, might also be necessary to the plants* However, not much attention vas paid to his statement in the nineteenth century* Seventy years later, Maze, using water culture experiments with carefully prepared culture vessels and nutrient salts, shoved that manganese, zinc, boron, aluminum, and silicon were all essential in small amounts for the healthy growth of the plant (33, 34, 35). His work stimulated maqy other plant physiologists and aroused their interest in studying minor elements with critical water cultures* The findings of Hase were later confirmed ty the work of McHargue (36), Barnette and Warner (5), Brenchley et al* (8), and Sommer (57). Sommer (58) also found that addition of copper in low concentrations increased the growth of sunflower, flax, and tomatoes, and demonstrated that copper was essential for plant growth. The last element added to the group of essential minor elements was molybdenum. Araon and Stout (1) obtained evidence that molybdenum was necessary for the growth of green plants. Although cobalt has not been proven to be essential for plant growth, it is required in the growth of animals, because cobalt is a constituent of vitamin (46). Underwood anil Filner (61) found that 4 when sheep or cattle grazed on a pasture low in cobalt content, a mineral deficiency disease occurred which could be cured ty feeding several doses of cobalt nitrate* Strontium is another element that has not been recognized as essential* However, McHargue (37) found significant yield increases of several grain crops when using a small amount of strontium carbonate in the presence of calcium carbonate, During recent years, strontium was found to be important in the bones of animals, because it stimulates deposition of calcium (49)* The Accumulation of Some Minor Elements in Plants Aluminum Latshaw and Miller (31) reported that the amount of aluminum in different parts of a fully developed corn plant, expressed as percent of dry weight, was 0*07# in leaves, 0*013# in stems, 0*023# in grains, 0,052% in cobs, and 0*98# in roots, McLean and Gilbert (38) found in several species of plants the greatest accumulation of aluadnxn was in the cortex of the roots. It accumulated in the protoplasm rather than in the cell walls or vacuoles* Hoffer et al. (23, 24) reported that aluminum accumulated in the vascular region of the corn nodes, and related this accumulation to the root-rot disease. This was also suggested as one of the means of liagnosing phosphorus deficiency. Boron Boron has been reported ty several investigators to be more concentrated in the leaves than in the other parts of the plant (11, 41, 3). Boron toxicity is generally characterized by leaf injury which is More pronounced in the older leaves (64). The symptoms are yellowing of tips and Margins, often followed ty marginal or spotted turning. Milikan (39) investigated the toxic concentration of boron, and found that 2.5-12.5 p.p.e. excess of boron caused the death of older flax leaves but no chlorosis. Shive et al. (9, 45) reported that boron toxicity was progressively accentuated ty increasing the potassium concentration in nutrient substrates, but vas decreased ty increasing the calcium concentration. Jones and Scarseth (28) found that a high calcium concentration resulted in boron deficiency unless the boron concentration was also high. Gauch and Dugger (15) suggested that the role of boron in the plant might be the formation of an ionizable sugar-borate complex which would facilitate sugar translocation. Calcium and Magnesium Latshav and Miller (31) found that about 60% of the calcium in a fully developed c o m plant vas in the leaves, and only 4% in the grain and cob. Magnesium vas present in somewhat smaller amounts than calcium and distributed differently, the grains and the leaves each containing one third of the total magnesium.