Plant Physiol.

(1973) 52, 183-185

Short Communication

Vanadium and Plant Nutrition

THE GROWTH OF LETTUCE (LACTUCA SATIVA L.) AND TOMATO (LYCOPERSICON ESCULENTUM

MILL.) PLANTS IN NUTRIENT SOLUTIONS LOW IN VANADIUM'

Received for publication March 12,1973

Ross M. WELCH AND EDWARD W. D. HUFFMAN, JR.2 United States Plant, Soil and Nutrition Laboratory, Agricultural Research Service, United States Department of Agriculture, Ithaca, New York 14850 and Department of Agronomy, Cornell University, Ithaca, New York 14850
ABSTRACT Lettuce (Lactuca sativa L.) and tomato (Lycopersicon esculentum Mill.) plants were grown in purified nutrient solutions with and without the addition of 50 nanograms per milliliter V. These experiments showed that lettuce and tomato plants can be grown to maturity on nutrient solutions containing less than 0.04 nanogram per milliliter V with tissue concentrations of less than 2 to 18 nanograms per gram V. Growth and dry matter yield were comparable to those of plants grown on nutrient solutions containing 50 nanograms per milliliter with tissue levels of V from 117 to 418 nanograms per gram. Thus if V is an essential element for lettuce and tomato plants, the adequate tissue level would be less than 2 nanograms per gram V derivable from a growth medium containing less than 0.04 nanogram per milliliter V.

The evidence that V is essential for the growth of higher plants is not conclusive (3-6, 11, 12). Its essentiality for microorganisms also appears to be questionable (4, 5), with a possible exception being the growth of the green alga, Scenedesmus obliquus (1, 2). However, growth rates of various microorganisms, higher plants, and animals reportedly have been increased by small additions of V to the growth media (1, 2, 7, 8, 11, 12, 14). The experiments reported here were carried out to test the hypothesis that V is an essential element for the normal growth of higher plants.

MATERIALS AND METHODS Double distilled deionized water and C.P. grade reagents were used throughout the experiment. All HNO. was glassdistilled before use. Glassware and plastic materials were washed in hot soapy water, rinsed, dipped in an acid bath (10% HCI), and finally rinsed several times with distilled water. Cheesecloth and glass wool were rinsed repeatedly in dilute HNO3, then thoroughly in distilled water.
1 Cornell University Department of Agronomy Paper No. 1022. - Present address: Huffman Laboratories, Inc., 3830 High Court, P. 0. Box 350, Wheat Ridge, Colo. 80033.

stock solutions were further purified by extracting (2-liter volumes, pH 4) twice with 100 ml of 8-quinolinol in chloroform (0.5% w/v) followed by an extraction with 2.5 liters of chloroform. The residual chloroform in the solutions was removed by extracting twice with ethyl ether. The ethyl ether residue was then removed by boiling the solutions. Tomato (Lycopersicon esculentum Mill. cv. "Glamar") and romaine lettuce (Lactuca sativa subsp. longifolia L., experimental line C898C, Department of Vegetable Crops, Cornell University) seeds were germinated, and the seedlings were grown as described previously (9). Briefly, the seeds were soaked in aerated water overnight and then germinated in the dark on cheesecloth in purified nutrient solution one-fifth the strength of that described previously (9). After 3 to 5 days, the seeds were transferred to a growth chamber with a growth regime of 24 C/ 18 C day-night temperatures, 16-hr days, and 2300 ft-c light intensity. The plants were grown for 1 to 2 weeks before transfer to 3.2-liter polyethylene containers, containing the nutrient solution described previously (9). The nutrient solutions were changed approximately every 10 days. Four harvests of lettuce and two of tomato were obtained. The lettuce plants were harvested after 26, 35, 43, or 56 days (4, 2, 3, and 2 plants per treatment, respectively), and the tomato plants after 26 or 35 days (4 and 6 plants per treatment, respectively). The tops and roots were harvested separately and freeze-dried. The dried plant tissues were pulverized by hand in plastic bags and then analyzed for V and, in some cases, Mo. The plants were grown in the purified nutrient solution with or without added NH4VO3 to give 50 ng/ml V in solution. Plants were also grown in unpurified nutrient solutions to test for possible effects of chemical residues left in the nutrient solutions from the purification procedures. No differences in yields were found between plants grown in unpurified and purified nutrient solutions. In one experiment, lettuce and tomato plants were grown on purified nutrient solutions as described above, but the Mo concentration in the nutrient solutions was reduced to onefifth its normal level. As before, the V treatments received 50 ng/ml V. The V concentrations in salts, nutrient solutions, and experimental plant tissues were determined by a catalytic method

Of the salts used to prepare the purified nutrient solutions, only KNO3, CaCI,,, MgSO,, and FeCi, contained significant concentrations of V. All nutrient salt solutions were purified as described previously (9). The 2 M KNO3, CaCI2, and MgSO,

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Plant Physiol. Vol.

52, 1973

(detection limit of 5 ng V per sample) previously reported (15). lower than that reported as a result of Mo interference with V The Mo content of plant tissues was determined by a modifica- determinations). tion of the thiocyanate colorimetric method of Purvis and The lettuce and tomato plants grown in nutrient solutions Peterson (13). with one-fifth the normal level of Mo developed severe symptoms characteristic of Mo deficiency (i.e. interveinal chlorosis, pale green veins, mottled leaves, curling of leaf margins, and RESULTS AND DISCUSSION stunted growth) after about 4 weeks of growth. These obThe average yields of both lettuce and tomato tops grown servations were independent of V additions and suggest that V in purified nutrient solution with and without 50 ng/ml V addi- cannot substitute for Mo in the nutrition of higher plants. tions are shown in Table I. Vanadium analyses of the purified The seeds of lettuce and tomato plants were analyzed for V nutrient solutions showed them to contain less than 0.04 ng/ml content. Tomato seeds contained 0.181 isg/g V (or approxiV. A growth response to added V was not evident. All plants mately 0.556 ng V/seed), and lettuce seeds contained 0.041 looked healthy with no observable differences between treat- ,ug/g V (or approximately 0.064 ng V/seed). Thus, the lettuce ments in over-all growth and development; no visual deficiency and tomato seeds contributed an insignificant amount of V to symptoms attributable to V developed in any of the plants. the plants grown on purified nutrient solutions. The mean, median, and range of V concentrations in the In one experiment, tomato plants were grown to maturity on tops and roots of the lettuce and tomato plants are shown in purified nutrient solutions with or without added V. Both treatTable II. The V concentration in plants receiving V additions ments produced healthy looking fruit and no differences in was in the normal range reported for various plant species (15). growth were observable between treatments. Because of possible Mo interference with V determination The results (Tables I and II) show that lettuce and tomato via the catalytic method used for V analysis (i.e. when Mo plants can be grown successfully to maturity on nutrient soluconcentrations exceed 10 ,ug/sample), the Mo concentration tions containing less than 0.04 ng/ml V with tissue concentrawas determined in several representative samples of lettuce tions of less than 2 to 18 ng/g V. Growth and dry matter and tomato tops and roots. The average Mo concentrations yield were comparable to those of plants grown on nutrient were 0.55 ,Ag/g and 52 jAg/g for lettuce tops and roots, and solutions containing 50 ng/ml V with tissue levels of V from 3.96 ,Ag/g and 8.4 ,g/g for tomato tops and roots. Thus Mo 117 to 418 ng/g. Thus, if V is essential for lettuce and tomato may have interfered with V determinations in the roots but not plant growth, the adequate tissue level must be less than 2 in the tops (i.e. the V concentration of the roots may have been ng/g obtainable from a growth medium containing less than 0.04 ng/ml. Table I. Yields of Plant Tops Grown on Of the 16 elements established as essential for the growth of Purified Nutrient Solutions with and higher plants, Mo is required in the least amount (4, 5). Plants without V Added as NH4V0O have been reported to show visual symptoms of severe Mo deficiency when grown in nutrient solutions containing less than V in Nutrient 10 ng/ml Mo (6, 10). Deficiencies of Mo have been reported Plant Plant Top Yield Replications for plant tissues (grown in solution cultures) containing from 0.01 to 0.5 ,ug Mo/g dry weight (10). The data reported in No. g dry wt ng/mi Table II show that most of the plant tops analyzed had V Lettuce <0.04 9 13.871 concentrations '5 to 250 of the Mo tissue concentrations at 50 Lettuce 9 16.00 which Mo deficiency symptoms occur. Therefore, if V is an essential element, it is required at a level much lower than the 8 <0.04 16.88 Tomato required level of Mo. 50 8 17.01 Tomato Hewitt (6) grew sugar beets, tomato, lettuce, alfalfa, white a Differences in means between treatments are not significant clover, and alsike clover in purified nutrient solutions contain(P > 0.05). ing from less than 2 to 3 ng/ml V. He was unable to show any response of these plants to V additions. However, levels of V ranged from 0.10 to 0.97 jtg/g in the tops and from 4 to 6.3 Table II. Vanadium Concentration in Plant Tops and ,ug/g V in the roots of the plants grown in solutions without Roots Grown on Purified Nutrient Solutions added V. He attributed these high tissue concentrations of V with and without V Added as NH4VO3 to greenhouse dust, which contained 150 ug/g V. Arnon and Wessel (2) have reported that V is an essential V in Nutrient Mean' Median' Plant Range element for the growth of the green alga, Scenedesmus obliquus, which required 0.1 ,tg/ml V in the nutrient medium. No. ng/ml ng/g dry wt Schwarz and Milne (14) reported that the physiologically es11 <0.04 6 5 <2-182 Lettuce, tops sential levels of V for the rat lie at or below 0.1 s.g/g dry matter 73 3 <0.04 45-119 Lettuce, roots in their diet. Diets containing 10 ng/g V decreased growth rates in rats and reduced wing and tail feather growth in chicks (7, 11 13 10 <0.04 <5-162 Tomato, tops 3 <0.04 61 <5-126 14). Thus, if V is essential to higher plants, it is required at a Tomato, roots level of 1s25o to less than 500 that required for the normal growth of Scenedesmus obliquus, rats, or chicks. 11 50 283 308 128-419 Lettuce, tops 10 50 278 234 117-418 Tomato, tops Acknowledgments-We thank Dr. W. H. Allaway for his many helpful suggestions and discussions, and V. A. Lazar for his Mo analyses of the plant
ma-

l In computing the mean and median values, samples containing less than 2 ng/g V were assumed to have 2 ng/g, and those containing less than 5 ng/g V were assumed to have 5 ng/g. 2of the 21 lettuce and tomato plants grown in nutrient solutions containing no added V, 13 contained no detectable levels of V.

terials.
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VANADIUM AND PLANT NUTRITION

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