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Boron nutrition and mobility,

and its relation to the
elemental composition of
greenhouse grown root crops.
II radish
a a a
B.J. Shelp , V.I. Shattuck & J.T.A. Proctor
a
Department of Horticultural Science , University
of Guelph , Guelph, Ontario, N1G 2W1, Canada
Published online: 11 Nov 2008.

To cite this article: B.J. Shelp , V.I. Shattuck & J.T.A. Proctor (1987) Boron nutrition
and mobility, and its relation to the elemental composition of greenhouse grown root
crops. II radish, Communications in Soil Science and Plant Analysis, 18:2, 203-219,
DOI: 10.1080/00103628709367811

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 COMMUN. IN SOIL SCI. PLANT ANAL., 18(2), 203-219 (1987)

BORON NUTRITION AND MOBILITY, AND ITS RELATION To

THE ELEMENTAL COMPOSITION OF GREENHOUSE GROWN ROOT CROPS. II RADISH

KEYWORDS: Raphanus sativus, brown heart, nutrient mobility, B

retranslocation, Ca mobility, tissue contents
Downloaded by [University of Cambridge] at 17:30 31 August 2015

B.J. Shelp, V.I. Shattuck and J.T.A. Proctor

Department of Horticultural Science
University of Guelph
Guelph, Ontario Canada N1G 2W1

ABSTRACT
The elemental distribution between the leaves and roots of
mature radish (Raphanus sativus cv Cherry Belle) plants grown in
the greenhouse with various concentrations of nutrient solution B
or Ca was determined to assess the role of phloem in the provision
of nutrients to the root, and the retranslocation of B under
deficient conditions. The relative composition and accumulation
of elements in different parts, and the ratio of their
concentrations in leaves:roots were used as a measure of their
uptake, relative mobility and retranslocation. The data indicate
that B, but not Ca is retranslocated in the phloem to the roots
when that particular element was in short supply in the nutrient
solution. B deficiency induced brown heart disorder in radish
roots but the severity was dependent on the degree of deficiency
below 28 µg g - l DM in the root. These symptoms were alleviated
when the root B concentrations were enhanced by foliar applications
of B. It is concluded that radish responded to B deficiency in a
fashion similar to that reported previously for rutabaga and that

203

Copyright © 1987 by Marcel Dekker, Inc.

 204 SHELP, SHATTUCK, AND PROCTOR

it might serve as a time-saving model system for examining the

mechanisms responsible for brown heart in rutabaga.

INTRODUCTION
Aspects of nutrition and mobility of B and its influence on
the elemental composition of rutabaga plants were reported in a
companion paper22. Continuing study of this question as well as
other nutritional requirements of rutabagas is hindered by its
lengthy growing season of about 14 weeks 2 2 under optimal greenhouse
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conditions of lighting and nutrients. To partly overcome this

problem, we are examining the possibility that summer radishes
could serve as a model system for rutabagas. They are ready for
use 3-6 weeks after sowing. Both radish and rutabaga are members
of the same family, Cruciferae, and although they are not in the
same genus, the roots develop similarly and consist of the upper
part of the true root, the hypocotyl, and the lower part of the
true stem. Also, both are known to exhibit signs of brown heart
and skin cracking or splitting as a result of B deficiency.20
In this paper, we report studies on B nutrition and mobility
in radishes grown in the greenhouse during the fall and winter
months under suboptimal lighting. Ca nutrition and mobility are
used for comparison because Ca is the classic example of a phloem
immobile element in herb
herbaceous plants ' and a Ca/B interaction
see 22
is reported often

MATERIALS AND METHODS

Radish (Raphanus sativus L. cv. Cherry Belle) plants were
grown in the greenhouse without supplemental lighting from the
early fall to late spring in 9L pots containing vermiculite. Night
and day temperatures varied but were maintained in the 15-25°C
range. Plants were fertilized every third day with 1L of full
strength Hoagland's solution as described by Lorenz and Maynard^^
with the exception that the sulphate salts of the micronutrients
were replaced by chloride salts. The standard solution was
prepared with distilled water and contained 0.05 mgB L"* and 200 mg

_
 BORON NUTRITION AND MOBILITY. II. 205

Ca L" 1 ; extra precautions, such as the use of recrystallized

chemicals and B-free glassware were not taken to remove
contaminating B. In the first experiment, B or Ca deficiencies
were induced by removing B from the standard solution or
substituting Na salts where appropriate. In the second experiment
B in the nutrient solution was varied from 0 to 2.5 mg L" 1 . In the
third experiment 3 B treatments were used: (1) B was not added to
the nutrient solution, (2) B at 0.05 mg L" 1 was added to the
nutrient solution, and (3) B was not added to the nutrient solution
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but was foliar applied twice weekly by painting the upper surface
of the leaves with a solution containing 300 mg B L" 1 as
recommended by Ontario Ministry of Agriculture and Food Publication
363. Precautions were taken to prevent B contamination of the soil
surface during painting, which began 21 or 28 days after sowing and
continued the 2 or 3 weeks until harvest. At 21 days the roots
were about 0.5 cm in diameter. Experiments were of a completely
randomized block design of 4 replicates with 8 plants/pot. Plants
were harvested after 6-8 weeks of growth when the control roots
reached the size desired for commercial harvest.
In preliminary experiments radish plants grown in the presence
(0.5 mg L" 1 ) and absence of B respectively contained 28.0 + 0.6
(n=3 pots of 8 plants each) and 10.6 ± 0.3 vg g-1 DM B (n=8 pots of
8 plants each) in their roots. Since the variation in B
concentrations was small between pots, the plants were usually
first assessed for deficiency symptoms (eg. root cracking,
splitting, blackening and elongation 17 ) and the fresh weight of the
plant parts (leaves and roots) determined separately before the
pots were bulked for dry weight and nutrient analysis as described
elsewhere 22 . Roots were rated as 0 for no splitting, and 1,2 and 3
respectively for mild, moderate, and severe splitting.

RESULTS
In the absence of added B the dry weights of the leaves and
roots of radish plants were unaffected, but the % DM was increased
(Table 1 ) . With the exception of B and perhaps Cu, the composition
Downloaded by [University of Cambridge] at 17:30 31 August 2015

TABLE 1

Effect of B and Ca Deficiencies on Yield Characteristics and Elemental Composition of Raphanus sativus cv.
Cherry Belle.

Plant part/ Element

treatment Fresh wt. Dry wt. DM Ca Mg Mn Cu Zn B
g/pota % % DM vg/g D M
Leaves
Control 32.6 ± 2.2b 2.51 ± 0.19 7.7 ± 0.1 4.90 0.56 5.60 4.95 1.15 161 4 15 160

-B 27.3 ± 5.0 2.56 ± 0.23 9.8 ± 0.8 4.35 0.53 6.16 5.32 1.28 154 2 14 43

-Ca 13.4 ± 0.7 1.36 ± 0.10 10.2 ± 0.5 4.60 0.74 3.80 2.41 1.84 157 2 12 190

Roots
Control 54.3 ± 6.9 2.85 ± 0.35 5.3 ± 0.1 2.75 0.46 7.80 0.72 0.42 20 1 15 60

-B 39.4 ± 4.0 2.78 ± 0.30 7.0 ± 0.2 3.45 0.55 7.33 0.89 0.42 22 3 17 25

-Ca 38.0 ± 5.0 2.46 ± 0.36 6.4 ± 0.2 2.80 0.46 6.21 0.35 0.32 19 1 10 45

a
each pot contains 8 plants
b
mean ± SE (n=4)
 BORON NUTRITION AND MOBILITY. II. 207

(Table 1 ) , accumulation and partitioning (Table 2) of the elements

in the plant parts also were similar. B, on the other hand,
declined from 160 and 60 u9 g~* DM respectively in the leaves and
roots of healthy plants to 43 and 25 in B-deficient plants (Table
1) resulting in a 30% increase in the B harvest index (Table 2 ) . B
and Cu showed the greatest percentage increase in the ratio of the
element concentration in the root:leaves (retranslocation index),
with smaller changes found for N, P and Zn. The roots of B-
deficient plants were either elongated and misshapen with patchy
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torn surface and dull skin, or severely split. Where the roots
were misshapen the internal tissue was also very glassy or watery
in appearance with one or two brown necrotic areas; the foliage did
not show any discernible symptoms which might be linked to B
deficiency.
When Ca was not added to the nutrient solution, the fresh and
dry weights of the leaves decreased and the % DM increased (Table
1). The dry weight of the roots was not significantly reduced, but
there was an increase in % DM. Thus the DM harvest index increased
to 0.64 from 0.53 in the control (Table 2 ) . The Ca concentration
of the leaves and roots was reduced by about 50% (Table 1 ) . K was
the only element which showed a decrease (21-33%) in both plant
parts; P, Mg and B concentration increased in leaves whereas in
roots Mg and B decreased (Table 1 ) . The total uptake of Ca
declined by 71%, accompanied by reductions of all other elements,
ranging from 18-50%, corresponding to a plant weight decrease of
28% (Table 2 ) . The harvest index however, was increased for a
number of nutrients (N,K,Mn,Zn,Cu) and in particular for Ca (by
50%). The retranslocation index (element concentration in
root:leaves) was decreased by about 50% for P and Mg, 40% for B,
and 17% for Zn; Ca and Cu were unaffected. In Ca-starved plants
the roots were slightly deformed and more pointed at the base than
the control; in some cases the internal tissue exhibited mild
necrosis with red discoloration. The leaves were generally smaller
than the control treatment with the leaf margins of young leaves
exhibiting forward rolling and eventually browning.
 TABLE 2 O
00
Effect of B and Ca Deficiencies on the Accumulation of Elements and the Harvest Indices for
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Growth and Nutrients in Raphanus sativus cv. Cherry Belle. Data are calculated from Table 1.

Element
Treatment Dry wt. N Ca Mg Mn Zn Cu
g/plant accumulation (mg/plant) accumulation (pg/plant)
Control 0.67 25.2 2.7 45.4 18.1 5.1 58 10 13 71
-B 0.67 25.9 2.9 45.1 20.1 5.6 57 11 13 23
-Ca 0.48 16.5 2.7 25.6 5.2 4.2 33 47

harvest index(accumulation in root:plant)

Control 0.53 0.39 0.48 0.61 0.14 0.29 0.12 0.50 0.2 0.30
en
-B 0.52 0.46 0.52 0.56 0.15 0.26 0.14 0.55 0.6 0.39
-Ca 0.64 0.52 0.51 0.74 0.21 0.23 0.18 0.60 0.5 0.30
retranslocation index (concentration in root:leaves)
G
n
Control 0.56 0.82 1.39 0.15 0.37 0.12 1.00 0.3 0.38 73

0.79 1.04 1.19 0.17 0.33 0.14 1.21 1.5 0.58

-Ca 0.61 0.62 1.63 0.15 0.17 0.12 0.83 0.5 0.23 o
3
3
o
pa
 BORON NUTRITION AND MOBILITY. II. 209

Figure 1 shows that the maximum DM accumulation occurred at

1.0 mg B L" 1 , although the root yield was relatively unaffected up
to at least 1.0 mg B L~l. The lowest yield for leaves was found at
0.0 mg B L" 1 ; 1.0 mg B L' 1 gave a slightly higher yield. With
increasing B in the nutrient solution the B concentration increased
drastically in the leaves but not in the roots. In both plant
parts, the B concentration reached a temporary plateau at 0.25 -
1.0 mg B L" 1 . Above this level, the B concentration increased to a
greater extent than the total B in that part. In fact, the 40 ug B
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g~l DM found in the root at 2.5 mg B L~l in the nutrient solution

was apparently toxic because yield and total accumulation of B
declined in comparison to the 1.0 mg B L~l treatment where the root
concentration was 30 ug B g"1 DM. Both the DM and B harvest
indices exhibited higher values at low B levels in the nutrient
level, but levelled off at 0.25 mg B L" 1 . Also the index of
retranslocation for B increased from 0.19 at 0.25 mg B L" 1 to 0.36
and 0.40 respectively at 0.125 and 0.0 mg B L" 1 .
In the third experiment, the absence of added B decreased the
radish yields, particularly the roots, which also exhibited a
greater DM content (Table 3 ) . The B accumulation was considerably
less than the control tissue yielding a much higher B harvest index
and increasing the incidence of severe root splitting. The yields
also were reduced when B was foliar-applied beginning 21 and 28
days after sowing; however, the B concentration and accumulation
increased by 129-285% to levels comparable to the 0.5 mg B L" 1
treatment. Possibly the most important observation was the
dramatic reduction in the incidence of severe root splitting.

DISCUSSION
In a companion paper dealing with rutabaga 22 , we assessed the
retranslocation of several elements, in particular boron after
transfer from sufficient B to the absence of added B by comparing
the element's concentration in roots or young leaves to that in
mature leaves. The magnitude of the ratio should give the relative
phloem mobility for each element, and an increased ratio under
 210 SHELP, SHATTUCK, AND PROCTOR
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'• 125

3.

E
3
U
o

0 1 2
SOLUTION B ( mg L"1 )

Fig. 1 . Effect of boron concentration in the nutrient solution on

dry matter accumulation ( o « )» B content ( A A ) and accumulation
( D M ) in leaves (open symbols) and roots (closed symbols), and
the partitioning of dry matter ( , no symbols) and B ( , no
symbols) between leaves and roots of Raphanus sativus cv. Cherry
Belle.
 BORON NUTRITION AND MOBILITY. II. 211

TABLE 3
Effect of B Application on Characteristics of Yield and B
Accumulation in Leaves (L) and Roots (R) of Raphanus sativus cv.
Cherry Belle.
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Roots
exhibiting
Harvest moderate to
Fresh Dry B B i ndex severe
wt DM cone accum DM B splitting
g/plant % ug/g DM
nutr. soln 0.5 mg B/L
L 4.83 0.87 9.8 170 80
R 12.94 0.72 5.5 34 25 0.61 0.24 22
nutr. soln 0.0 mg B/L
L 3.45 0.38 11.1 19 7
R 6.62 0.48 7.2 15 7 0.56 0.50 82
nutr. soln 0.0 mg B/L; foliar applied 300 mg B/L - 21d
L 3.06 0.34 11.0 ND a - - -
R 6.67 0.41 6.1 40 16 0.57 - 28
nutr. soln 0.0 mg B/L; foliar applied 300 mg B/L - 28d
L 3.63 0.41 11.2 ND - - -
R 9.55 0.54 5.7 49 27 0.57 - 34

a
ND, Not determined
b
4 replicate pots were bulked for weight and nutrient analyses
 212 SHELP, SHATTUCK, AND PROCTOR

deficiency conditions should indicate retranslocation of that

element 25 . In the present study, this approach as well as the
determination of plant elemental uptake and partitioning between
the roots and leaves (harvest index) has been used to investigate B
nutrition and mobility in radish. B mobility and retranslocation
were compared to that of calcium which is a classic example of a
phloem immobile element 1 5 ' 1 8 ' s e e refs * c i t e d in 2 2
In the control plants, the relative concentration of B or Ca,
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and N,K or P in the roots compared to the leaves (Tables 1 and 2)

indicates that the roots are predominantly fed with nutrients
through the phloem1^. N,P and K which are present at adequate
levels are known to be phloem mobile and readily
1 *•> 1 8
retranslocated ' . Mn, Cu and to a lesser extent Mg was also low
in the root suggesting that they are also relatively phloem •
immobile. On the other hand, Zn appeared to be readily mobile in
the phloem of radish. Loneragan1^ suggested that Mn, Cu and Zn
have a variable and limited mobility from old leaves with the
retranslocation depending on the nutrient status of the plant. The
Mn level reported here for leaves was in the sufficient supply
(100-600 pg g"1 DM), perhaps approaching toxic levels seen in some
other plants 13 (eg. 160 yg g"1 DM in soybean1-6). Both the Cu and
Zn content in the leaves are in the range of low to sufficient
values seen for most plants " ' . Although further work is
required to establish the optimal tissue range for these elements
in root crops the data suggest that Zn was mobile even when
concentrations are not at luxury levels, contrary to Loneragan's
proposal1^.
When B was omitted from the nutrient solution, the tissue
content (Table 1) and plant accumulation of the element decreased,
accompanied by an increased partitioning of B to the roots (Table
2 ) . The net result is that retranslocation of this element from
the leaves to the roots occurred under conditions of B starvation
(Table 2 ) . With the exception of Cu evidence was lacking for an
effect of B nutrition on the elemental composition of leaves as
suggested by Francois for Mg, K, P and Ca in tomato^, and broccoli,
 BORON NUTRITION AND MOBILITY. II. 213

cauliflower and radish (cv Crimson Giant)^. In those studies

however, the changes were found at much higher solution
concentrations of B. For Cu, the plant uptake was unaffected but
its partitioning to the root was markedly increased resulting in a
4-fold increase in copper mobility due to B deficiency. This might
be attributed to induction of leaf senescence which is known to
promote Cu retranslocation^, or to stimulation in the roots of
phenol metabolism 9 '23 , a process involving enzymes with bound
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Cu3,23 B deficiency in roots of radishes and rutabaga is often

20 22
associated with browning of the internal tissue ' , an effect
commonly attributed to changes in phenol production^.
These studies demonstrate that B, although relatively
immobile, was retranslocated in the phloem. As discussed in our
companion paper22, it is uncertain as to how foliar-applied B
reached the root (Table 3). Nevertheless, radish are similar to
rutabagas in remobilizing the B to the root and this is effective
in alleviating the symptoms of brown heart and its associated
splitting (Table 3 ) .
The severity of B deficiency varied between experiments
(Tables 1 and 3; Fig 1 ) . This may be due to the changing daylength
encountered throughout the fall and winter months in the greenhouse
without supplemental lighting2!. For example, plants growing in
decreasing daylength require less boron see 19. Moreover, with our
experimental procedure, there was unexplained B contamination (See
Materials and Methods) since the levels in the tissue under
deficiency conditions were not as low as observed by others using
1 21 24
sand culture systems ' ' or as might be predicted based on the
low B content of young radish seedlings^.
In experiments where the yield was slightly reduced (Fig 1)
or not at all (Table 2) the partitioning of dry matter to the root
was also only slightly affected. This corresponded to leaf and
root B levels of 43-74 and 25-27 ug g"1 DM respectively (Table 1,
Fig 1 ) . In contrast, when yield was markedly decreased (Table 3)
the DM harvest index was decreased and the corresponding B levels
were 19 and 15 yg g"l DM respectively in the leaves and roots.
 214 SHELP, SHATTUCK, AND PROCTOR

Despite this variation in the response of dry matter to B

starvation, there was always an increase in % DM and the harvest
and retranslocation indices for B. Good growth was found at B
levels of 150-170 ug g"1 DM in the leaves and 28-60 \tq g-1 DM in
the roots. Toxic levels were possibly attained in one experiment
(Fig 1) when leaves and roots contained 260 and 40 yg B g"1 DM
respectively, but a greater range of solution B is required to
carefully evaluate this proposal.
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In a study of R_. sativus cv Cherry Belle, conducted on sandy

loam in the greenhouse, Gupta? listed the deficient, sufficient,
and toxic levels for plant tops when the roots began to swell as
<9, 96-217, and >217 u9 g"1 DM; however, as the author pointed out
only a small range of B application rates, and none between 0 to 1
jjg L"1 were used. In our study, B levels sufficient for growth
ranged from (>27) 28-60 (<40) in the roots and (>74) 150-170 in
the leaves at harvest. Below these levels, the exact effect on
growth will likely depend on the B concentration. Our results are
in agreement with those for another summer radish cultivar, Scarlet
Globe, which showed that B deficiency enhanced the % DM, and
decreased the yield of both plant parts causing a decrease in the
DM harvest index2^. Furthermore, maximum and relatively consistent
fresh weight yields have been observed from 0.25-5.0 mg B L" 1 in
the nutrient solution by Scripture and McHargue 21 . With Raphanus
sativus L., Agarwala et al. 1 found that root yield was more
dramatically reduced than leaves by B deficiency at leaf
concentrations ranging from 10.5 to 18.5 ug B g"1 DM. Levels of
68.5 (0.37 mg L" 1 in the nutrient solution) and 850 (18.5 mg L"1 in
the nutrient solution) yg g"1 DM produced good and poor growth
respectively.
Together with the B deficiency-induced decline in the harvest
index, Scripture and McHargue 21 , and Bible et a!.2 also found that
the reducing sugars decreased and increased respectively in the
roots and leaves; sugars released by invertase treatment
accumulated in both parts 21 . It is well established in radish 24
and other plants1? that drastic anatomical changes in the vascular
 BORON NUTRITION AND MOBILITY. II. 215

tissue may occur with B-deficiency; however, little is known about

how such changes influence the physiological function of such
tissues. Considering that in the present study, B retranslocation
in the phloem was apparently not hindered, it is suggested that B-
deficiency restricts the movement of sugars by preventing the
loading of sugars onto the phloem for translocation to the root.
However Shkolnik^^, on the basis of extensive literature review,
has argued that the effects of B-deficiency on sugar translocation
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are indirect, reflecting growth processes rather than transport.

Ca starvation decreased yield as well as tissue Ca by about
50% (Table 1) indicating that the sufficient Ca concentration was
likely between 24.1 and 49.5, and 3.5 and 7.2 mg g"1 DM
respectively in leaves and roots. Higher plants often contain Ca
in the order of 5-30 mg g~l DVfl-5 but there is considerable range
between plants in the Ca concentration required for optimal yields.
For example in ryegrass, this occurs at 0.7 mg g"l DM and in tomato
at 12.9 mg g-1 DM • . Thus radish would appear to have a
relatively high Ca requirement. Since Ca accumulation was
depressed more extensively than tissue Ca and yields (Tables 1 and
2) it is suggested that root growth and function may be impaired.
Edwards e_t al_4. have shown that Ca starvation in peach seedling
depresses the uptake rates of P,K, and Mg. That this proposal
applies to the present study seems likely since the reduction in
the uptake of all other elements was consistent with the reduction
in growth (Table 2 ) . Contrary to the previous suggestion by
Tanaka^S, we found no evidence that Ca may inhibit B uptake apart
from the reduction in growth, although that author analyzed the B
concentration of the leaves only. With the exception of B, Mg and
P, the partitioning of DM and other elements to the roots was
enhanced by a similar amount. However, the other three elements,
particularly Mg and B, showed marked reductions in retranslocation,
perhaps contributing to symptoms of calcium deficiency by altering
the balance of elements in the roots. Interestingly, Ca mobility
was unaffected suggesting that either Ca is not retranslocated
 216 SHELP, SHATTUCK, AND PROCTOR

and/or the root obtains a major portion of its Ca supply by

absorption through its surface^.
In conclusion, this study provides additional evidence that B
movement and retranslocation, although limited, occurred in the
phloem at least under conditions of B starvation in root crops, and
that Ca retranslocation and mobility the in phloem is extremely
limited. Considering that the mature leaves of both radish and
rutabaga plant are fed predominantly by the xylem stream, B levels
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in these tissues may vary considerably depending on the temperature

and relative humidity of the environment. On the other hand the B
concentrations of the roots should be less dependent on the
environmental conditions and vary less because they are fed with
nutrients predominatly via the phloem. A comparison of radish and
rutabaga22 B contents under the slightly differential environmental
conditions of our studies shows that B deficiency is associated
with 15-27 and 14-20 yg B g~l DM respectively in roots of radish
and rutabaga. Adequate root B levels were 28-60 yg g~l DM for
radish and 27 ug g~* DM for rutabaga. It is suggested that the
optimal B levels are similar for both crops but a larger range of
solution and tissue B levels in rutabaga are required to complete
the comparison. More importantly, radish responded similarly to B-
deficiency and foliar application of B as rutabaga suggesting that
it might be an appropriate time-saving model system for
investigating the mechanism(s) responsible for the brown heart
disorder in rutabaga.

ACKNOWLEDGEMENTS
This research was supported by financial assistance from the
Natural Sciences and Engineering Research Council of Canada
(B.J.S.) and the Department of Horticultural Science. The authors
gratefully acknowledge Earl Gagnon for performing the elemental
analysis, M. Vindum, M. da Silva and D. Louttit for assisting with
the care of plants, and J. Beswick for typing the manuscript.
 BORON NUTRITION AND MOBILITY. II. 217

REFERENCES
1. Agarwala, S.C., S. Farooq and C.P. Sharma. 1978.
Interaction of boron supply and nitrogen source in growth,
boron uptake, nitrogen fractions and sugars in radish. In:
Nitrogen Assimilation and Crop Productivity, eds. S.P.
Sen, Y.P. Abrol and S.K. Sinha. Assoc. Publ. Co., New
Delhi, p 185-194.
2. Bible, B.B., H-Y. Ju, and C. Chong. 1981. Boron
Downloaded by [University of Cambridge] at 17:30 31 August 2015

deficiency in relation to growth and thiocyanate toxin

content of radish. Scientia Hort. 15:201-205.
3. Bould, C., E.J. Hewitt and P. Needham. 1983. Diagnosis of
Mineral Disorders in Plants. Vol. 1, Principles. H.M.
Stationery Office, London. 174pp.
4. Edwards, J.H., D.W. Reeves, B.D. Horton and J.M. Thompson.
1985. Influence of calcium concentration on growth, tissue
concentration, and nutrient uptake of in. vitro propagated
plums and Lovell seedlings. J. Plant Nutrit. 8:327-344.
5. Francois, L.E. 1984. The effect of excess boron on tomato
yield, fruit size, and vegetative growth. J. Amer. Soc.
Hort. Sci. 109:322-324.
6. Francois, L.E. 1986. Effect of excess boron on broccoli,
cauliflower, and radish. J. Amer. Soc. Hort. Sci. 111:494-
498.
7. Gupta, U.C. 1983. Boron deficiency and toxicity symptoms
for several crops as related to tissue boron levels. J.
Plant Nutrit. 6:387-395.
8. Kahata-Pendias, A. and H. Pendias. 1984. Trace Elements
in Soils and Plants. CRC Press, Inc., Boca Raton, Florida.
315pp.
9. Lewis, D.H. 1980. Boron, lignification and the origin of
vascular plants - a unified hypothesis. New Phytol.
84:209-229.
10. Loneragan, J.F. 1975. The availability and absorption of
trace elements in soil-plant systems and their relation to
movement and concentrations of trace elements in plants.
 218 SHELP, SHATTUCK, AND PROCTOR

In: Trace Elements in Soil-Plant-Animal Systems, eds.

D.J.D. Nicholas and A.R. Egan. Academic Press, N.Y.
p 109-134.
11. Loneragan, J.F. and K. Snowball. 1969. Calcium
requirements of plants. Aust. J. Agric. Res. 20:465-478.
12. Loneragan, J.F., J.S. Gladestones and W.J. Simmons. 1968.
Mineral contents in temperate crop and pasture plants II.
Calcium. Aust. J. Agric. Res. 19:353-64.
Downloaded by [University of Cambridge] at 17:30 31 August 2015

13. Lorenz, O.A. and D.N. Maynard. 1980. Knott's Handbook For
Vegetable Growers. Wiley-Interscience, New York. 390pp.
14. Melati, M.R. and E. Bellini. 1981. Boron distribution in
barley and radish seedlings. Plant Sci. Lett. 22:205-209.
15. Mengel, K. and E.A. Kirby. 1982. Principles of Plant
Nutrition. Intern. Potash Inst., Switzerland. 654pp.
16. Ohki, K. 1977. Manganese and zinc status related to
maximum growth for selection agronomic crops. Proc.
Intern. Symp. Soil Environm. Fert. Mgmt. Intensive Agric.
Tokyo, p.659-668.
17. Palser, B.F. and W.J. Mcllrath. 1956. Responses of
tomato, turnips, and cotton to variations in boron
nutrition. II. Anatomical responses. Bot. Gaz. 118:58-
71.
18. Pate, J.S. 1975. Exchange of solutes between phloem and
xylem and circulation in the whole plant. In: Encyc.
Plant Physiology Vol. 1, Transport In Plants I Phloem
Transport, eds., M.H. Zimmerman and J.A. Milburn.
Springer-Verlag, Berlin, p 451-473.
19. Pi1beam, D.J. and E.A. Kirby. 1983. The physiological
role of boron in plants. J. Plant Nutrit. 6:563-582.
20. Scaife, A. and M. Turner. 1986. Diagnosis of Mineral
Disorders in Plants. Vol 2, Vegetables. H.M. Stationery
Office, London. 96pp.
21. Scripture, P.N. and J.S. McHargue. 1945. Boron supply in
relation to carbohydrate metabolism and distribution in the
radish. J. Amer. Soc. Agron. 37:360-364.
 BORON NUTRITION AND MOBILITY. II. 219

22. Shelp, B.J. and V.I. Shattuck. 1986. Boron nutrition and
mobility, and its relation to the elemental composition of
greenhouse grown root crops I. Rutabaga. Commun. Soil Sci.
Plant Anal. in press.
23. Shkolnik, M.Y. 1984. Trace Elements In Plants.
Developments in Crop Science, Vol 6. Elsevier Scientific
Publ. Co., New York. 463 pp.
24. Skok, J. 1941. Effect of boron on growth and development
Downloaded by [University of Cambridge] at 17:30 31 August 2015

of the radish. Bot. Gaz. 103:280-294.

25. Tanaka, H. 1967. Boron absorption by crop plants as
affected by other nutrients of the medium. Soil Sci. Plant
Nutrit. 13:41-44.
26. Van Goor, B.J. and P. Van Lune. 1980. Redistribution of
potassium, boron, iron, magnesium and calcium in apple
trees determined by an indirect method. Physiol. Plant.
48:21-26.