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<strong>Boron</strong> <strong>Tolerance</strong> <strong>of</strong> <strong>Sugar</strong> <strong>Beets</strong> <strong>in</strong> <strong>Relation</strong> <strong>to</strong><br />

<strong>the</strong> <strong>Growth</strong> <strong>and</strong> <strong>Boron</strong> Content <strong>of</strong> Tissues<br />

JAMES VLAMIS AND ALBERT CLRICH i<br />

Receivedlor publication March 2,1973<br />

The extensive work <strong>of</strong> Ea<strong>to</strong>n (4)2 established sugar beet plants as<br />

hav<strong>in</strong>g a high requirement for boron <strong>and</strong> a high <strong>to</strong>lerance for boron<br />

excess. His work <strong>and</strong> that <strong>of</strong> o<strong>the</strong>rs <strong>in</strong>dicates that plant tissues <strong>in</strong> <strong>the</strong><br />

deficiency range tend <strong>to</strong> run about 10 <strong>to</strong> 35 parts per million boron (1,<br />

7, 12).<br />

In <strong>the</strong> <strong>to</strong>xic range Ea<strong>to</strong>n showed that <strong>the</strong> <strong>to</strong>ps <strong>and</strong> lam<strong>in</strong>ae <strong>of</strong>sugar<br />

beets could conta<strong>in</strong> hundreds <strong>of</strong> ppm B without affect<strong>in</strong>g yield very<br />

markedly.<br />

This work presents <strong>the</strong> results <strong>of</strong> experiments designed <strong>to</strong> study<br />

<strong>the</strong> growth o[ sugar beets <strong>in</strong> solutions o[ high concentrations <strong>of</strong> B<strong>and</strong><br />

<strong>to</strong> correlate growth with <strong>the</strong> B content <strong>of</strong> selected tissues. A previous<br />

report covered <strong>the</strong> B nutrition <strong>of</strong> sugar beets <strong>in</strong> <strong>the</strong> deficiency range<br />

(12).<br />

Plant culture<br />

Materials <strong>and</strong> Methods<br />

<strong>Sugar</strong> beet seeds were planted <strong>in</strong> vermiculite <strong>and</strong> irrigated with a<br />

nutrient solution conta<strong>in</strong><strong>in</strong>g 0.25 ppm B (Table 1). The seeds used<br />

(F58-554H I-MS <strong>of</strong> NB 1 X NB4) were previously treated with a fungicide,<br />

Phygon XL, at a rate <strong>of</strong> 1 gram per 100 grams <strong>of</strong> seed.<br />

Eighteen days later <strong>the</strong> seedl<strong>in</strong>gs were each wrapped with Dacron<br />

fiber <strong>and</strong> transferred <strong>to</strong> cork r<strong>in</strong>gs. Three seedl<strong>in</strong>gs were placed <strong>in</strong><br />

Table l.-Chemical composition <strong>of</strong> nutrient solution added at two <strong>in</strong>tervals.<br />

Salts added, meq/l Microelements, mg/l<br />

Ca(N03), 5.0 Mn , as MnSO,· 4 H,O 0.25<br />

KN03 2.5 Ln, as ZnSO, . 7 H,O 0025<br />

MgSO, 2.0 C u, as CuSO, · 5 H2O 0010<br />

KH,PO, 1.0 Mo, as Mo03 0.005<br />

K,SO, 0.5 Fe,asEDTA 2.5<br />

NaCI 0.5 <strong>Boron</strong> (see Tabk 2)<br />

1 Plant Physiologists, Department <strong>of</strong>Soils <strong>and</strong> Plant Nutrition, University <strong>of</strong>CaJifornia, Berkeley.<br />

'Numbers <strong>in</strong> paren<strong>the</strong>ses refer <strong>to</strong> lit erature cited.


VOL. 17, No.3, APRIL 1973<br />

281<br />

perforated masonite covers mounted on 5-gallon cans pa<strong>in</strong>ted with<br />

alum<strong>in</strong>um pa<strong>in</strong>t on <strong>the</strong> outside <strong>and</strong> Amercoat No. 33 on <strong>the</strong> <strong>in</strong>side.<br />

The masonite lids were pa<strong>in</strong>ted with alum<strong>in</strong>um on <strong>the</strong> upper surface<br />

<strong>and</strong> Valspar underneath. The conta<strong>in</strong>ers were filled with nutrient<br />

solutions with compositions as given <strong>in</strong> Table 1.<br />

<strong>Boron</strong> was added <strong>to</strong> <strong>the</strong> solutions so as <strong>to</strong> give <strong>the</strong> follow<strong>in</strong>g range<br />

<strong>of</strong>B <strong>in</strong> ppm: 0.5, 2,4,8,16,32,64, <strong>and</strong> l28 (Table 2). In two <strong>of</strong> <strong>the</strong><br />

experiments a 1 ppm B solution was <strong>in</strong>cluded. Three replicates <strong>of</strong>each<br />

rate were set up. The experimental period for <strong>the</strong> ma<strong>in</strong> experiment<br />

extended from April 26 <strong>to</strong> June 8. Midway <strong>in</strong> this period ano<strong>the</strong>r dose<br />

<strong>of</strong>nutrients was added <strong>to</strong> each pot, but without a fur<strong>the</strong>r addition <strong>of</strong>B.<br />

The test was conducted <strong>in</strong> a smog-free clear glass greenhouse. The<br />

nutrient solutions were kept at a pH <strong>of</strong> 5.0 <strong>to</strong> 6.5, <strong>and</strong> distilled water<br />

was added as needed.<br />

Harvest<strong>in</strong>g<br />

The plants were harvested 6 weeks after transplant<strong>in</strong>g. Fresh<br />

weights were taken <strong>of</strong> <strong>the</strong> shoots <strong>and</strong> roots separately. The leaves were<br />

divided <strong>in</strong><strong>to</strong> three age groups-young, mature, <strong>and</strong> old. Leaves which<br />

had not reached full development were classified young, <strong>and</strong> those<br />

show<strong>in</strong>g signs <strong>of</strong> senescence were placed <strong>in</strong> <strong>the</strong> old category. The<br />

<strong>in</strong>termediate leaves were called mature. The leaves were <strong>the</strong>n separated<br />

<strong>in</strong><strong>to</strong> blades <strong>and</strong> petioles for boron analysis. The residue consisted<br />

<strong>of</strong> dead or wilted leaves. The fibrous roots were removed from <strong>the</strong><br />

s<strong>to</strong>rage root, centrifuged <strong>to</strong> remove free water, <strong>and</strong> fresh weights<br />

taken.<br />

Preparation <strong>of</strong> samples <strong>and</strong> chemical analysis<br />

The plant parts were placed <strong>in</strong> paper bags <strong>and</strong> dried at 70°C <strong>in</strong> a<br />

forced-draft oven for 72 hours. The dried material was weighed,<br />

ground <strong>in</strong> a Wiley mill with a 40-mesh screen, <strong>and</strong> s<strong>to</strong>red <strong>in</strong> plastic vials.<br />

The samples were analyzed for boron at room temperature by <strong>the</strong><br />

curcum<strong>in</strong> method, us<strong>in</strong>g an acetate buffer (6). All o<strong>the</strong>r analyses were<br />

made by established methods (2, 8). .<br />

<strong>Boron</strong> excess <strong>and</strong> yield<br />

Results<br />

The data obta<strong>in</strong>ed are shown <strong>in</strong> Table 2 <strong>and</strong> figure 1. The growth<br />

curve for <strong>the</strong> <strong>to</strong>ps shows that maximum yields were obta<strong>in</strong>ed at 0.5, 2,<br />

4, <strong>and</strong> 8 ppm B <strong>in</strong> <strong>the</strong> external solution . This is followed by a steep drop<br />

at 16 ppm <strong>and</strong> <strong>the</strong>n a more gradual decl<strong>in</strong>e <strong>in</strong> yield for <strong>the</strong> rema<strong>in</strong>der<br />

<strong>of</strong> <strong>the</strong> curve. The beet curve follows <strong>the</strong> one for <strong>the</strong> <strong>to</strong>ps except for a<br />

small shift <strong>to</strong> <strong>the</strong> left. Thus, <strong>the</strong> first significant drop <strong>in</strong> beet yield occu rs<br />

at 8 ppm as opposed <strong>to</strong> 16 ppm for <strong>the</strong> <strong>to</strong>ps. The fibrous root response<br />

resembles very closely that <strong>of</strong> <strong>the</strong> <strong>to</strong>ps.


284 J OC R:-;AL OF TH E A. S. S. B. T .<br />

Figure 2.-0n <strong>the</strong> left, leaf series <strong>of</strong> one plant grown <strong>in</strong> 64 ppm B<br />

show<strong>in</strong>g severe symp<strong>to</strong>ms on old leaves. On <strong>the</strong> right, closeup <strong>of</strong> second<br />

oldest leaf with severe symp<strong>to</strong>ms <strong>of</strong> B <strong>to</strong>xicity.<br />

>-<br />

0::<br />

o<br />

10 -<br />

- 0 YOUNG BLADES<br />

-0 MATURE BLADES<br />

--x OLD<br />

BLADES<br />

L L L L<br />

600 1200 1800 2400<br />

PPM BORON IN BLADES<br />

Figure 3.-<strong>Relation</strong> <strong>of</strong> <strong>to</strong>p dry weights <strong>to</strong> boron content <strong>of</strong> young, mature,<br />

<strong>and</strong> old blades.


VOL. 17, No.3, APRIL 1973 287<br />

closer <strong>to</strong> 16 ppm. This puts <strong>the</strong> sugar beet plant <strong>in</strong> <strong>the</strong> category <strong>of</strong><br />

plants <strong>to</strong>lerant <strong>of</strong> excess B <strong>in</strong> comparison with many cultivated plants<br />

(11). In <strong>the</strong> extensive work by Ea<strong>to</strong>n (4) several varieties <strong>of</strong> sugar beet<br />

were all found <strong>to</strong> be fairly <strong>to</strong>lerant <strong>of</strong> high B <strong>in</strong> <strong>the</strong> external solution. A<br />

recent study by EI-Sheikh et al., (5) showed that cucumber, squash,<br />

melon, <strong>and</strong> corn had a 50% reduction <strong>in</strong> growth with 6,12,12, <strong>and</strong> 16<br />

ppm B <strong>in</strong> solution, respectively.<br />

In <strong>the</strong> case <strong>of</strong>sugar beets <strong>the</strong> focus has <strong>to</strong> be on <strong>the</strong> beet root more<br />

than on <strong>the</strong> shoot. The first suggestion <strong>of</strong> a significant decrease <strong>in</strong> beet<br />

yield is at 8 ppm B. The tissue analysis, however, shows that <strong>the</strong>re was<br />

very little B accumulation <strong>in</strong> ei<strong>the</strong>r <strong>the</strong> beet or <strong>the</strong> fibrous roots.<br />

Therefore, <strong>the</strong> reason <strong>the</strong> beet yield dropped at a lower B concentration<br />

than <strong>the</strong> <strong>to</strong>ps must be an <strong>in</strong>direct one. The pho<strong>to</strong>syn<strong>the</strong>tic rate<br />

must be adversely affected before <strong>the</strong> yield <strong>of</strong><strong>the</strong> shoots is decreased so<br />

that <strong>the</strong>re is enough sugar manufactured for <strong>the</strong> growth <strong>of</strong> <strong>the</strong> <strong>to</strong>ps<br />

<strong>and</strong> fibrous roots but not enough for <strong>the</strong> s<strong>to</strong>rage root.<br />

The boron analysis <strong>of</strong> <strong>the</strong> tissues <strong>in</strong>dicates that <strong>the</strong> yield <strong>and</strong> <strong>the</strong><br />

appearance <strong>of</strong> <strong>the</strong> leaves are not affected until <strong>the</strong> B content <strong>of</strong> <strong>the</strong> old<br />

leaves reaches 600 ppm. At <strong>the</strong> same po<strong>in</strong>t <strong>the</strong> mature leaves have 450<br />

<strong>and</strong> <strong>the</strong> young leaves 300 ppm, both without symp<strong>to</strong>ms. Br<strong>and</strong>enburg<br />

showed that B content <strong>in</strong>creases with <strong>the</strong> physiological age <strong>of</strong> <strong>the</strong><br />

tissues (1) . In this <strong>in</strong>stance, s<strong>in</strong>ce <strong>the</strong> <strong>in</strong>jury first affects <strong>the</strong> old leaves, it<br />

is preferable <strong>to</strong> use <strong>the</strong> B content <strong>of</strong> <strong>the</strong>se tissues as an <strong>in</strong>dication <strong>of</strong> <strong>the</strong><br />

status <strong>of</strong> sugar beet plants from <strong>the</strong> <strong>to</strong>xicity po<strong>in</strong>t <strong>of</strong> view.<br />

It is <strong>of</strong> economic <strong>in</strong>terest that sugar beets <strong>to</strong>lerate high B s<strong>in</strong>ce<br />

<strong>the</strong>re are many areas where high boron occurs <strong>in</strong> soils or irrigation<br />

waters. These areas occur primarily <strong>in</strong> <strong>the</strong> western parts <strong>of</strong>this country<br />

(3,9, 10) where sugar beets should be given preference over crops with<br />

a greater sensitivity <strong>to</strong> high B situations.<br />

Summary<br />

<strong>Sugar</strong> beet plants were grown for 6 weeks <strong>in</strong> 20-liter pots conta<strong>in</strong><strong>in</strong>g<br />

nutrient solutions with B rang<strong>in</strong>g <strong>in</strong> amount from adequate <strong>to</strong><br />

extremely <strong>to</strong>xic. The lowest B concentration <strong>in</strong> solution was 0.5 ppm<br />

<strong>and</strong> <strong>the</strong> highest was 128 ppm.<br />

No significant decrease <strong>in</strong> growth <strong>of</strong><strong>to</strong>ps or fibrous roots occurred<br />

<strong>in</strong> plants grow<strong>in</strong>g <strong>in</strong> 0.5, 1,2,4, or 8 ppm B. The first drop <strong>in</strong> yield for<br />

<strong>the</strong>se plant parts came at 16 ppm <strong>and</strong> <strong>the</strong>n cont<strong>in</strong>ued <strong>to</strong> drop at a fairly<br />

uniform rate at 32, 64, <strong>and</strong> 128 ppm. A 50% reduction <strong>in</strong> growth for<br />

<strong>to</strong>ps <strong>and</strong> fibrous roots came at 28 ppm, <strong>and</strong> for beet roots at 16 ppm B.<br />

Only a trace <strong>of</strong> symp<strong>to</strong>ms appeared at 8 ppm <strong>and</strong> showed as a<br />

mosaic chlorosis <strong>and</strong> cupp<strong>in</strong>g <strong>of</strong> tne old leaves. 'These symp<strong>to</strong>ms became<br />

worse with time <strong>and</strong> were very severe at <strong>the</strong> highest B concentrations.<br />

A f<strong>in</strong>al symp<strong>to</strong>m apeared as a spott<strong>in</strong>g at <strong>the</strong> tips <strong>and</strong> edges <strong>of</strong> <strong>the</strong><br />

old leaves that moved downward with time as a cont<strong>in</strong>uous mass <strong>of</strong>


288 JOURNAL OF THE A. S. S. B. T.<br />

necrotic tissues <strong>to</strong>ward <strong>the</strong> bases <strong>of</strong> <strong>the</strong> leaves. The roots appeared<br />

healthy at all B levels <strong>and</strong> a decrease <strong>in</strong> yield was <strong>the</strong> only evidence <strong>of</strong><br />

<strong>to</strong>xicity, even at 128 ppm B.<br />

The B content <strong>of</strong> <strong>the</strong> leaves <strong>in</strong>creased with <strong>the</strong>ir physiological age.<br />

Blades were higher <strong>in</strong> B than <strong>the</strong> correspond<strong>in</strong>g petioles by a fac<strong>to</strong>r <strong>of</strong><br />

as much as twelve. The B content <strong>of</strong> <strong>the</strong> old blades <strong>of</strong> <strong>the</strong> plants where<br />

<strong>the</strong> first drop <strong>in</strong> <strong>to</strong>p yield occurred was above 600 ppm. At this po<strong>in</strong>t<br />

<strong>the</strong> mature blades had 450 <strong>and</strong> <strong>the</strong> young blades 300 ppm. It was<br />

suggested that <strong>the</strong> old blades could be used as diagnostic material <strong>in</strong><br />

determ<strong>in</strong><strong>in</strong>g <strong>the</strong> B status <strong>of</strong> <strong>the</strong> nutrient medium <strong>in</strong> <strong>the</strong> <strong>to</strong>xic range.<br />

The B content <strong>of</strong> <strong>the</strong> roots was low <strong>and</strong> <strong>of</strong> <strong>the</strong> order <strong>of</strong> magnitude<br />

found <strong>in</strong> <strong>the</strong> petioles.<br />

The sugar beet plant was found <strong>to</strong> be <strong>to</strong>lerant <strong>of</strong> B relative <strong>to</strong> some<br />

crop'; studied by o<strong>the</strong>r <strong>in</strong>vestiga<strong>to</strong>rs.<br />

Literature Cited<br />

(I) BRANDENB URG , E. 1940. Ueber die Grundlagen del' Boranwendung <strong>in</strong><br />

der L<strong>and</strong>wirtschaft. Phy<strong>to</strong>path. Zeitschr. 12: 1-112.<br />

(2) CHAPMAN, HOMER D. <strong>and</strong> PARKER F. PRATT. 1961. Methods <strong>of</strong> Analysis<br />

for Soils, Plants <strong>and</strong> Waters. University <strong>of</strong> Califormia, Division <strong>of</strong><br />

Agricultural Sciences, Berkeley, Ca. 309 p.<br />

(3) EATON, F. M. 1935. <strong>Boron</strong> <strong>in</strong> soil <strong>and</strong> irrigation water <strong>and</strong> its effects<br />

on plants with particula r references <strong>to</strong> <strong>the</strong> San Joaqu<strong>in</strong> Valley<br />

<strong>of</strong> California. USDA Tech. Bul. 448.<br />

(4) EATON, F. M. 1944. Deficiency, <strong>to</strong>xicity, <strong>and</strong> accumulation <strong>of</strong> boron <strong>in</strong><br />

plants. J. Agr. Res. 69:237-277.<br />

(5) EL-SHEIKH, ADELM., A ULRICH, S. K. AWAD <strong>and</strong> A. E. MAWARDY. 1971.<br />

<strong>Boron</strong> <strong>to</strong>lerance <strong>of</strong> squash, melon, cucumber <strong>and</strong> corn. J. Am . Soc.<br />

Hort. Sci. 96: 536-537.<br />

(6) GRINSTEAD, R. R. <strong>and</strong> SI GR ID SNIDER. 1967. Modification <strong>of</strong> <strong>the</strong><br />

method for 10" level boron determ<strong>in</strong>ation. Analyst 92: 532-533.<br />

(7) IlCGHES, J. D. 1959. A nOlt' on foliar diagnosis <strong>of</strong> boron deficiency <strong>in</strong><br />

beetroot. Queensl<strong>and</strong> J. Agr. Sci. 16: 85-86.<br />

(8) JOHNSO N, C. M. <strong>and</strong> A. ULRICH. 1959. Analytical methods for use <strong>in</strong><br />

plant analysis. Cal. Agr. Exp. Sta. Bu!. 766.<br />

(9) KELLEY, W. P. <strong>and</strong> S. M. BROWN. 1928. <strong>Boron</strong> <strong>in</strong> <strong>the</strong> soils <strong>and</strong> irrigation<br />

waters <strong>of</strong> sou<strong>the</strong>rn California <strong>and</strong> its relation <strong>to</strong> citrus <strong>and</strong> wa<strong>in</strong> ut<br />

culture. Hilgardia 3: 445-458.<br />

(10) REEVE , R. C., A. F. PILLSBURY <strong>and</strong> L. V. WILCOX. 1955. Reclamation<br />

<strong>of</strong> a sal<strong>in</strong>e <strong>and</strong> high boron soil <strong>in</strong> <strong>the</strong> Coachella Valley <strong>of</strong> California.<br />

Hilgardia 24:69-91.<br />

(II) UNITED STATES SALINITY LABORATORY STAFF, 1954. Diagnosis<br />

<strong>and</strong> improvement <strong>of</strong> sal<strong>in</strong>e <strong>and</strong> alkali soils. USDA H<strong>and</strong>book 60,<br />

U. S. Govt. Pr<strong>in</strong>t<strong>in</strong>g Office, Wash<strong>in</strong>g<strong>to</strong>n, D. C. 160 p.<br />

(12) VLAMIS,JAMES <strong>and</strong> ALBERT ULRICH. 1971 . bOl'On nutrition <strong>in</strong> <strong>the</strong> growth<br />

<strong>and</strong> sugar content <strong>of</strong> sugarbeets. J. Am. Soc. <strong>Sugar</strong> Beet Technol.<br />

16: 428-439.

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