Plant and Soil 150: 289-294, 1993.

© 1993 KluwerAcademic Publishers. Printedin the Netherlands. PLSO 9824

Boron deficiency in soybean [Glycine max (L.) Merr.] peanut (Arachis

hypogaea L.) and black gram [Vigna mungo (L.) Hepper]: Symptoms in
seeds and differences among soybean cultivars in susceptibility to boron
deficiency

B. R E R K A S E M 1, R.W. BELL,2 S. L O D K A E W a and J.F. L O N E R A G A N 2

1Faculty of Agriculture, Chiang Mai University, Chiang Mai, Thailand and 2School of Biological and
Environmental Science, Murdoch University, Murdoch, Western Australia

Received 19 October 1992. Accepted in revised form 27 January 1993

Key words: black gram, boron deficiency, hollow heart, peanut, soybean cultivars

Abstract

The susceptibility of soybean cultivars (cvs) to boron (B) deficiency was examined in three experiments
on a Typic Tropaqualf in Northern Thailand: one experiment also included peanut cv Tainan 9 and
black gram cv Regur.
Without added B (B0), B deficiency depressed seed yield by 60% in soybean cv NW1 compared with
30% in cv SJ5, 40% in cv 7016, 45% in peanut, and 93% in black gram. B deficiency also induced a
localised depression on the internal surface of one or both cotyledons of some soybean seeds resembling
the symptom of 'hollow heart' in peanut seeds. It induced 50% hollow heart in peanut, 17% in soybean
cv NWl, 5% in SJ5, and 1% in 7016: black gram seeds had no symptoms. Addition of B decreased or
eliminated the symptoms.
In a comparison of 19 soybean cvs, the incidence of hollow heart symptoms in seeds at B0 varied
widely from none in two cvs to 75% in cv Buchanan: cv NWl showed intermediate susceptibility to B
deficiency with 36% hollow heart in its seed while SJ5 was insensitive with 1%. Application of B
eliminated hollow heart except from one seed in one cultivar. The results suggest that susceptibility to B
deficiency is sufficiently important and variable among soybean genotypes to warrant its inclusion as a
selection criterion when breeding cultivars for areas with low soil B.

Introduction widely used throughout the country. However,

Soybean is generally regarded as having a low
susceptibility to B deficiency (Martens and Wes-
termann, 1991). Recent experiments on a low B
soil in Northern Thailand supported this opinion
when B deficiency had no effect on soybean but
halved the seed yield of sunflower and black
gram and induced severe hollow heart symptoms
in peanut seed (Rerkasem et al., 1988). The
soybean cultivar used in the Thai experiments
was SJ5, an established cultivar which has been
when a new cultivar (NW1) was released and
grown in the area, it became B deficient sug-
gesting that the reputation of soybean for low
susceptibility to B deficiency may be more an
accidental property of the cultivars in common
use than an inherent property of the specie in
general. The present experiments were under-
taken to explore differences among soybean
cultivars in their susceptibility to B deficiency
and to compare them with one common cultivar
each of peanut and black gram.
290 Rerkasem et al.
Materials and m e t h o d s
Soybean (Glycine max cv. N W l ) seeds were
sown in three experiments in the field at Chiang
Mai, Thailand on a sandy loam textured Typic
Tropaqualf of the San Sai series.
In Experiments 1 and 2, seeds were sown on
the site of an earlier experiment with four
replications of four B treatments as main plots
and four cropping sequences as subplots (Experi-
ment 3, Rerkasem et al., 1988). In the original
experiment, B was applied as borax
(Na2B4OT.10H20) to supply 0, 1, 2, and 4 k g B
ha -1 (referred to as B0, B1, B2, B4) immedi-
ately prior to the sowing of the first crop in
D e c e m b e r 1983. In January 1987, soybean was
sown as the 9th crop in the fourth cropping
sequence subplots and its response to the origi-
nal B treatments is reported as Experiment 1 of
the present paper. On the same subplots rice was
sown as crop 10 and, in January 1988, the
subplots were split for planting five genotypes as
the l l t h crop since applying the original borax
treatments. Prior to sowing the genotypes of
crop 11, 13kg P h a -1 as calcium aerophos
(Ca(H2PO4)2) and 25 kg K ha -1 as K2SO 4 were
applied to each subplot. The hot water soluble B
levels in the 0 to 10 cm layer of soil in B0, B1,
B2 and B4 treatments after crop 11 were 0.05,
0.08, 0.10 and 0.12mg B kg -1 soil respectively,
when determined as described previously (Rer-
kasem et al., 1988). Crop 11 is reported as
Experiment 2 in the present paper.
Experiment i
N W l soybean seeds were harvested at maturity,
examined for symptoms, dried at 75°C, dry
ashed at 500°C for 10 h, and analysed for B by an
azomethine-H procedure as described previously
( R e r k a s e m et al., 1988). The cotyledons of 100
seeds from each B treatment were separated and
examined for indentations of their internal sur-
faces; seeds with abnormal indentations were
classified as 'hollow heart' because of the re-
semblance of the disorder to the symptom of
'hollow heart' in B deficient peanut seed (Harris
and Brolman, 1966).
Experiment 2
In January 1988, soybean cv. NW1, soybean cv.
SJ5, soybean cv 7016 peanut (Arachis hypogaea
cv. Tainan 9), and black gram (Vigna mungo cv
Regur) were sown as subplots in each of the
residual B treatment main plots of Experiment 1.
Seeds were harvested, examined and analysed as
described for Experiment 1.
Experiment 3
In January 1989, 19 soybean genotypes from
diverse sources were sown as subplots within
duplicate main plots of two B treatments, no B
(B0) and B applied as borax at 1 kg B ha -1 (B1).
The seeds of each genotype were sown at a
spacing of 0.20 m in rows 0.25 m apart on raised
beds 1 × 2 m 2 in size. At maturity, 100 seeds
from each subplot were assessed for hollow
heart.
Results and discussion
Hollow heart and DM of seed
In the absence of applied B in Experiment 1, B
deficiency depressed by 33% the seed dry matter
(DM) of the recently released soybean cv N W l
(Table 1). It also induced a localized depression
on the internal surface of one or both cotyledons
of 22% of the seeds. In severe cases, the internal
depression was very large and the seeds were
shrivelled and deformed. Some seeds also had a
small external 'dimple' on the testa surface
Table 1. Effects of soil B treatments on dry matter, B
concentration and incidence of hollow heart in seeds of
soybean cv. NWl. (Experiment 1)
Soil B treatment
B0 B1 B2 B4
B concentration
(mg B kg- 1) 10a 14bc 16c 20d
Dry matter (kg ha -1) 1219a 1893b 1833b 1646b
Hollow heart (%) 22a 2b lb 0b
Values within a row are significantly different (p <0.05)
unless followed by the same letter; Duncan's Multiple Range
Test.
 Boron deficiency in soybean 291

apparently due to a localised depression on the
external surface of the cotyledon.
The internal symptoms on the cotyledons of
the affected soybean seeds resembled the symp-
toms of 'hollow heart' in peanuts (Harris and
Brolman, 1966) so closely that the same term
'hollow heart' will be used to describe the
symptom in both species. Application of B at all
levels enhanced seed DM and decreased the
incidence of hollow heart to 2% or less.
When compared with the established soybean
cv SJ5 and the promising new cv 7016 in Experi-
ment 2, NWl was more susceptible to B de-
ficiency (Table 2). At B0, B deficiency induced
17% hollow heart in NW1 but only 5% in SJ5
and 0.5% in 7016: it also depressed seed DM by
60% in NW1, 30% in SJ5 and 40% in 7016.
In the development of hollow heart symptoms,
soybean cv NW1 was appreciably less susceptible
to B deficiency than peanut cv Tainan 9 but in
the depression of seed DM it was more suscep-
tible (Table 2). At B0, B deficiency induced 50%
hollow heart in peanut cv Tainan 9 and de-
pressed seed DM by 45%. Black gram cv Regur
showed no symptoms of hollow heart but, at B0,
many of its seeds were shrivelled and B de-
ficiency depressed its seed DM by over 90%.
The results indicate that seed yield of soybean
cv NWl was more susceptible to B deficiency
than were seed yields of peanut cv Tainan 9 and
soybean cvs SJ5 and 7016 but much less sucept-
ible than seed yield of black gram cv Regur. In
the expression of hollow heart symptoms, soy-
bean cv NW1 was also more susceptible to B
deficiency than soybean cvs SJ5 and 7016 al-
though less susceptible than peanut.
That soybean cv NW1 is more susceptible to B
deficiency than cv SJ5 was confirmed when 19
soybean cvs were compared in Experiment 3: at
B0, NWl had 36% hollow heart in its seed while
SJ5 had 1%. Only Buchanan with 75% hollow
heart was more susceptible than NWl. Two
cultivars (8219-59-4 and 8037-42-1) were as sus-
ceptible as NW1 and five others (B107, 82A-
1842, CM60, Wilis, SJ1) were as insensitive as
SJ5. Nine cultivars (UFV80-84, Farrer, 7016,

Table 2. Effects of soil B treatments on dry matter, B concentration and incidence of hollow heart in seeds of soybean cultivars
NW1, SJ5 and 7016, peanut cv. Tainan 9, and black gram cv. Regur. (Experiment 2)
Cultivar Soil B treatment
B0 B1 B2 B4
Seed dry matter (kg ha l)
NW1 668a 1176b 1400bc 1697c
SJ5 1333a 1809b 1755b 1901b
7016 1457a 1964ab 2019b 2315b
Tainan 9 2170a 3503b 3749b 3973b
Regur 84a 51 lb 720c 1193d

B concentration (mg B kg 1)o

NWl 7cd llefg llefg 19ij
SJ5 8cd 10def 14ghi 16hi
7016 8cd 12efgh 13fgh 18ij
Tainan 9 4b llefg 1lefg 14ghi
Regur 3a 5b 7cd 9de

Hollow heart ( % )b
NWl 16.8b 3.3c 4.8c 0.0d
SJ5 4.8c 0.0d 0.0d 0.0d
7016 0.5d 0.0d 0.0d 0.0d
Tainan 9 49.8a 10.0b 8.0bc 0.0d
Values within rows for seed dry matter and values within rows and columns for B concentration and hollow heart are significantly
different (p < 0.05) unless followed by the same letter; Duncan's Multiple Range Test.
aAnalysis of variance after log transformation.
bAnalysis of variance after arcsin transformation. No deformed seeds were observed in black gram.
292 R e r k a s e m et al.

V a l d e r , 8207-24-3, C a n a p o l i s , SK1, Shi Shi, h o l l o w h e a r t while S J5 s e e d w i t h 10 m g B kg -1

B R 1 ) w e r e i n t e r m e d i a t e b e t w e e n N W l a n d SJ5
in t h e i r s u s c e p t i b i l i t y to B deficiency. A p p l i c a -
t i o n o f B e l i m i n a t e d h o l l o w h e a r t in all cultivars
e x c e p t B R 1 w h i c h h a d s y m p t o m s in o n l y o n e
seed.
B o r o n concentrations in seeds
I n E x p e r i m e n t 1, s e e d s f r o m t h e B0 t r e a t m e n t
w i t h a high i n c i d e n c e o f h o l l o w h e a r t h a d an
a v e r a g e B c o n c e n t r a t i o n o f 1 0 m g B kg -~ D M :
s e e d s f r o m t r e a t m e n t s w i t h 2 % o r less h o l l o w
h e a r t h a d 14 o r m o r e m g B kg -~ D M ( T a b l e 1).
I n e a c h B t r e a t m e n t o f E x p e r i m e n t 2, all
s o y b e a n cvs h a d s i m i l a r B c o n c e n t r a t i o n s in t h e i r
s e e d : t h e y i n c r e a s e d f r o m a b o u t 8 m g kg -1 D M
at B 0 t o 16 o r m o r e at B4 ( T a b l e 2). S o y b e a n
s e e d w i t h 12 o r m o r e m g B kg -~ D M h a d no
h o l l o w h e a r t in a n y cv o r t r e a t m e n t . S o y b e a n cv
N W 1 s e e d with l l m g B kg -1 h a d 3 to 5 %
h a d n o n e a n d 7016 w i t h 8 m g B kg -1 h a d o n l y
0.5% hollow heart. While not conclusive, the
results suggest t h a t t h e g r e a t e r s u s c e p t i b i l i t y o f
cv N W l to B d e f i c i e n c y results f r o m a p o o r e r
u t i l i z a t i o n o f B in t h e s e e d r a t h e r t h a n in t h e
s u p p l y o f B to t h e s e e d . W h e n t a k e n t o g e t h e r
with t h e results o f E x p e r i m e n t 1, t h e r e s u l t s also
suggest t h a t s o y b e a n p l a n t s with s e e d B c o n c e n -
t r a t i o n s o f 8 o r less m g B k g -1 D M a r e B
deficient, o f 9 to 13 m a y b e deficient, a n d o f 14
o r m o r e a r e n o t deficient.
I n e a c h B t r e a t m e n t o f E x p e r i m e n t 2, b l a c k
g r a m cv R e g u r h a d B c o n c e n t r a t i o n s in its s e e d
o f a p p r o x i m a t e l y half o f t h o s e in t h e s o y b e a n
s e e d . Since in e a c h t r e a t m e n t its s e e d y i e l d was
also l o w e r t h a n t h a t o f t h e s o y b e a n cvs, t h e
h i g h e r s u s c e p t i b i l i t y o f b l a c k g r a m cv R e g u r to B
deficiency a p p e a r s to r e s u l t f r o m a p o o r e r a b i l i t y
to s u p p l y its s e e d with B. B l a c k g r a m cv R e g u r
p l a n t s with s e e d B c o n c e n t r a t i o n s o f 7 o r less m g

Table 3. Effect of soil B treatments on incidence of hollow heart in seeds of nineteen soybean cultivars. (Experiment 3)
Cultivar Source Soil B treatment a Significance b
B0 B1
Hollow heart (%)
B107 Australia 0a 0 NS
82A- 1842 Thailand 0a 0 NS

CM60 Thailand lab 0 NS

SJ5 Thailand lab 0 NS

Wilis Indonesia lab 0 NS

SJ1 Thailand 4ab 0 NS

UFV 80-84 Australia 8bc 0 *

Farrer Australia 9bc 0 *
7016 Thailand 9bc 0 *
Valder Australia 1lbc 0 *
8207-24-3 Thailand 1lbc 0 *
Canapolis Australia 13bc 0 *
SK1 Thailand 17c 0 *
Shi Shi Taiwan 19c 0 *
BR1 Australia 19c 1 *

NW1 Thailand 36de 0 **

8219-59-4 Thailand 41de 0 **
8037-42-1 Thailand 54ef 0 **

Buchanan Australia 75f 0 ***

aStatistical analysis after arcsin transformation of square root of proportion of seeds with hollow heart; for comparisons within the
B0 column, values are significantly different (p < 0.05) unless followed by the same letter; Duncan's Multiple Range Test.
bFor comparisons within a row: NS = not significant at p <0.05; *significant at p <0.05, **p <0.01; ***p < 0.001.

B k g - I were B deficient and of 9 or m o r e were
not.
A t B0, the B concentration in the seed of
p e a n u t cv Tainan 9 was only half that of the
soybean cvs: in all other B treatments, it was
similar. Peanut seed with 11 or less mg B kg -1
had 8% or m o r e hollow heart symptoms and
seed with 14 or m o r e mg B kg -1 had none.
Discussion
T h e present study has shown that B deficiency
produces internal d a m a g e of cotyledons in soy-
b e a n seeds similar to the hollow heart s y m p t o m
in p e a n u t seeds. In this paper, the s y m p t o m in
soybean has b e e n called hollow heart and it is
p r o p o s e d that this n a m e be adopted for general
use.
In peanuts, hollow heart is specific to B
deficiency and has been used for diagnosis of B
deficiency in crops (Harris and Gilman, 1957;
Netsangtip et al., 1985). It is suggested that,
since there are no reports of other factors caus-
ing hollow heart in soybean seed, the symptom
m a y also be specific to B deficiency in soybean
and might therefore provide a simple criterion
for diagnosis of B deficiency.
In the present experiments, hollow heart in
soybean a p p e a r e d no m o r e sensitive to B de-
ficiency than seed D M , since appreciable levels
of hollow heart were always associated with a
significant depression in seed D M . In this respect
it contrasts with hollow heart in peanut which is
generally a m o r e sensitive indicator of B de-
ficiency than is seed D M (Cox et al., 1982; Hill
and Morrill, 1974; Morrill et al., 1977; R e r k a s e m
et al., 1988).
In several studies, soybean crops did not
respond to B when grown on low B soils on
which other crop species were severely B de-
ficient (Oertli and Roth, 1969; R e r k a s e m et al.,
1988), leading to the generalisation that soybean
crops are insensitive to B deficiency (Martens
and Westermann, 1991). The present experi-
ments show that this generalisation is not valid
for the specie as a whole, since soybean cultivars
varied widely in their susceptibility to B de-
ficiency on a low B soil. The high susceptibility
to B deficiency of peanut and black gram shown
Boron deficiency in soybean 293
in the present and other experiments ( R e r k a s e m
et al., 1988) should also not be taken as an
indication of the behaviour of these species in
general since, in other experiments a wide vari-
ation in the susceptibility of their cultivars to B
deficiency has been found (Keerati-Kasikorn
personal communication; R e r k a s e m , 1990).
The results suggest that the reputation of
soybean for low susceptibility to B deficiency
may be more an accidental property of the
cultivars in c o m m o n use than an inherent prop-
erty of the specie in general; they also suggest
that susceptibility to B deficiency is sufficiently
important and variable a m o n g soybean geno-
types to warrant its inclusion as a selection
criterion when breeding cultivars for areas with
low soil B. It may be noted that of the Thai
genotypes in the present study, SJ1, SJ5 and
CM60 which were relatively tolerant of low B
were selected in Chiang Mai on soil types similar
to that of the present study: N W l and SK1 which
were appreciably m o r e susceptible to B de-
ficiency were selected in areas where B de-
ficiency is not common.
Acknowledgements
The authors are grateful to the University of
Chiang Mai for the provision of facilities and to
the Australian Centre for International Agricul-
tural Research, C a n b e r r a (Project 8603) for
financial support.
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Section editor: A C Borstlap