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Role of boron in plant growth: a review

Article  in  Journal of Agricultural Research · January 2009

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 Role of boron in plant growth 329

ROLE OF BORON IN PLANT GROWTH: A REVIEW

Waqar Ahmad*, A. Niaz**, S. Kanwal***, Rahmatullah***
and M. Khalid Rasheed**

ABSTRACT

Boron (B) is considered as an essential element for plant growth and

development. Sexual reproduction in plant is more sensitive to low B, than
vegetative growth. Considerable research activities have been directed at
accentuating the physiological and biochemical role of B in plant growth and
development. This paper reviews the literature (upto the year 2006) focusing on
the role of boron in cell wall integrity, cell division, plasma membranes, phenol
metabolism, and its requirement for the nitrogen fixation and in the
reproductive growth of plants.

KEYWORDS: Boron; plant growth; cell division; cell wall; Pakistan.

INTRODUCTION

Mineral nutrition of plants is important for controlling physiological and

biochemical processes of plants. Its deficiency may lead to changes in these
processes and disturbed plant growth and yield. Boron is one of mineral
nutrients that are required for normal plant growth. The essentiality of B for
growth and development of higher plants has been earlier demonstrated (27,
43, 46). The main functions of boron relate to cell wall strength and
development, cell division, fruit and seed development, sugar transport and
hormone development. Some functions of boron interrelate with those of
nitrogen, phosphorus, potassium and calcium in plant. The most important
functions of boron in plants are thought to be its structural role in cell wall
development; and stimulation or inhibition of specific metabolism pathways.

Boron and plant cell wall

*Sustainable Land Management Project, Ministry of Environment, Islamabad Email:

[email protected], **Soil Chemistry Section, AARI, Faisalabad, ***Plant Nutrition
Lab., Institute of Soil and Environmental Sciences, University of Agriculture, Faisalabad,
Pakistan.

J. Agric. Res., 2009, 47(3)

330 W. Ahmad et al.

Plant cell wall is composed of three main layers i.e. primary wall, secondary
wall and middle lamella. Primary wall is set down by cells before and during
active growth and comprises pectic polysaccharides (Ca. 30%), cross-linking
glycans-hemicellulose (Ca. 25%), cellulose (15-30%) and protein (Ca. 20%)
(13). Secondary wall in some cells deposits additional layers inside the
primary wall. This occurs after growth stops or when the cell begins to
differentiate. The secondary wall is mainly for support and comprises
primarily cellulose and lignin. Middle lamella that binds adjacent cells is
composed of pectic polysaccharides. The actual content of wall components
varies with species and age. Some researchers estimate that over 90
++
percent of total B is localized in cell walls (26, 29). Boron along with Ca is
able to form complexes with several cell wall components such as pectins
(21), polyhydroxyl polymers, polyols and Ca++ (31, 44). This is the reason that
B is implicated in synthesis and stability of cell wall (16) by forming esters
with cis diol groups present in cell wall (26). This provides rigidity, strength
and shape to the cell.

Cell division

Boron is considered to be essential for actively growing regions of plants,

such as root tips, new leaf and bud development. According to Rerkasem
(37) boron is especially required more in meristematic cells than in mature
tissues. That is why; first effect of boron deficiency usually appears in
meristems, as described by Warington (46). Higher meristematic B
requirement may rise because of low phloem transport from shoots to other
parts of plant, leading to higher accumulation of B in leaves (37).

Ion fluxes

Boron plays an important role in both structural and functional integrity of

plasma membranes (28, 31). In B-deficient plants, plasma membranes are
highly leaky and lose their functional integrity (11). In many instances, it is
+ + + -3 +2
proved that fleet alterations in ion fluxes i.e. H , K , Rb , PO4 , and Ca is
associated with B deficiency (38, 39).

The proton efflux by membrane-bound ATPases is a main driving force for

ion uptake and responsible for a gradient in electrical potential across the
membranes (36, 41). The stimulated activity of plasma membrane NADH
+
oxidase and H secretion in cultured carrot cells with boron has also been
reported by Barr and Crane (3). Apparently, auxin affects B-induced H+
extrusion. According to Hu and Brown (20) the stimulatory effect of B on H+-

J. Agric. Res., 2009, 47(3)

 Role of boron in plant growth 331

ATPase activity requires the presence of auxin, or enhancement in proton

release by auxin requires the presence of B. In the studies with sunflower
root cells and dense leaved elodea (Elodea densa) leaf cells, a significant
depolarisation of membranes was found after transfer of cells from B-
containing to B-free solution. Hence, boron nutrition has marked effects on
proton secretion and creation of an electrical potential gradient across the
membranes (5).

Boron supply enhances the activity of membrane bound ATPase and

subsequently causes hyperpolarization of plasma membrane by stimulating
+
K ion uptake The pumping activity of the membranes with subsequent
membrane hyperpolarization, results in an increased driving force for K+
influx (42). As far as K+ is concerned, it is involved in the opening and closing
+
of stomata. By affecting K influx, B was also found involving in enhancement
of stomatal opening in epidermal strips of dayflower (Commelina communis)
+
(41). A number of researchers have observed the excessive leakage of K
with B deficiency that leads to a decrease in membrane integrity. Some
researchers consider this effect of B as a primary effect while others attribute
it to secondary effect of boron deficiency.

Uptake rate of phosphate is decreased by B deficiency and is rapidly restored

by resupply of B to deficient plants for one hour (40). More rapid effects of B
on ion influx or efflux were shown by Poole (32) in Zea mays (maize). He
found that resupply of B to deficient maize for one hour caused a significant
restoration of phosphate uptake (Table 1).

The data (Table 1) show the variation in phosphorus uptake (nmol/g/h) in

faba bean and maize under the pretreatment of root tips with B for one hour
and without pretreatment (no B) of root tips. The data markedly indicate that
pretreatment of root tips increased the uptake of inorganic phosphorus both
in faba bean and maize as compared to without pretreatment.
Table 1. Effect of treatment on subsequent uptake of inorganic phosphate
by root tip zones of faba bean (Vicia faba) and maize (Zea mays)
grown in (+ B) or (- B)

Pretreatment of root tips Phosphate uptake (nmol/g/h)

Faba bean Maize
+B -B +B -B
No boron 112 52 116 66
-5
10 M B 152 108 190 171
Source: Poole (32)

J. Agric. Res., 2009, 47(3)

332 W. Ahmad et al.

Phenol metabolism

Phenol metabolism also plays a very important role in plant growth which
seems to be affected by B nutrition. Kamali and Childers (24) studied the
effects of B deficiency on several phenolics and enzyme activities involved in
the biosynthesis of these compounds in tobacco plant. In B-deficient plants a
higher amount of phenolic compounds is accumulated (10). This higher
amount of phenolic is very hazardous for plants. With increase in the amount
of phenolic compounds, enzymatic or non-enzymatic oxidation takes place, in
which phenolic compounds act as substrate. Thus, toxic quinones and
destructive O2 species are generated (1, 23). The phenolics after oxidation
corroborate changes in ion fluxes accompanied by changes in membrane
potential (19). It has been suggested that phenolics result in reversible
alterations in membrane permeability, and this effect of phenolics occurs
during their passage through the membranes (18, 19).

In B-deficient plants, not only the phenolic compounds increase but also
defence capacity of cells against toxic O2 species is weakened due to
reduced levels of ascorbic acid, SH-compounds and H2O2 scavenging
enzymes (9). So, it can be suggested that B deficiency renders membrane
leakiness and alteration in ion flux characterized by peroxidative damage and
structural alteration in plasma membranes.

Boron and nitrogen fixation

Boron is also involved in nitrogen fixation. Loomis and Durst (26) reported
that boron is an essential micronutrient required for growth and development
of vascular plants, diatoms and species of marine algal flagellates, while
bacteria, fungi, green algae and animals apparently do not require B. Not
only the leguminous plants but also cyanobacteria require B when dependent
on N2 fixation. Under B-deficient conditions, nodule weight and N2 fixation
capacity of legumes is usually decreased. Bolanos et al. (6) investigated the
effect of boron on symbiotic nitrogen fixation in pea (Pisum sativum). The
absence of boron in culture medium resulted in lower number of nodules and
alterations in nodule development. Nodules were not only found less in
number but also nodule structure was disorganized and not easily
distinguishable in B deficient media. Examination of boron deficient nodules
showed dramatic changes in cell wall and in both peribacteriod and infection
thread membranes, suggesting a role of boron in the stability of these
structures (7). The formations of ineffective nodules have also been reported
earlier (8, 42). This alteration of nodule development led to an inhibition of

J. Agric. Res., 2009, 47(3)

 Role of boron in plant growth 333

nitrogenase activity. These results indicated that boron is a requirement for

normal nodule development and functionality.

Boron and plant reproductive growth

It has been reported that boron deficiency limits reproductive growth. In

wheat, B deficiency causes poor anther and pollen development, low grain
set and stunted growth (12, 33). In vitro germination tests also showed that B
was required for pollen germination and tube growth in wheat (12). Seed
setting is important for improving yield from any soil particularly salt affected
soils. Aslam et al. (2) reported better pollination, seed setting, low spike
sterility and more grain formation in different cultivars of rice as an effect of
boron nutrition. Rashid et al. (33) observed a substantial increase in grain
yield of rice cultivars, due to reduced panicle sterility after B application. Jiang
and Miles (22) identified critical stages of anther development of wheat
during which B deficiency caused a significant and irreversible decrease of
floret fertility. According to Cakmak (9), B has limited phloem mobility in
crops like wheat. Hence, B supply is required for healthy reproductive growth
(anther, pollen and ovule development). Loomis and Durst (25) reported the
abortion of apical meristems leading to the lack of flowering and development
in peach (Prunus persica) in boron free medium. Some physiologists (4, 35,
36, 47) have observed the effect of boron nutrition on flowering, stamens,
pistil and young reproductive leaves (Table 2 and 3). Generally, the
concentration of boron in flower, stamen, and pistil is increased with increase
in boron supply from low to adequate level. This is the indicator of response
of young reproductive plant parts and leaves to supply of boron.

Table 2. Boron concentration (mg/g dry weight) in young reproductive parts and
leaves in oilseed rape (Brassica napus), wheat (Triticum aestivum) and green
gram (Vigna radiata) in relation to B supply.

Crops B supply Flower Stamen Pistil Leaf

1
Oilseed rape Low - 19 19 10
Adequate - 38 40 17
2 3
Green gram Low 30 - 19 11
Adequate 61 - - 34
4 5
Wheat Adequate - 16 8 3
1 2
Adapted from Dell B. L. Huang (14), Pot study in soil by Zhang et al. (47) Pot
study in solution culture by Bell et al. (4). The data for the low boron plants is
3 4
after transfer to solution without B for 4 days. Data for young pods. Field
5
study by Rerkasem (37). Data for anthers.

J. Agric. Res., 2009, 47(3)

334 W. Ahmad et al.

Table 3. Mean B concentration (mg/kg) in young whole leaves as affected by B

application to a B-deficient media in wheat (Triticum aestivum L.) and rice
(Oryza sativa L.) cultivars.

B concentration (mg/kg) in leaves

Wheat cultivars Control +B Rice cultivars Control +B
Rohtas-90 5.2 11.0 Super Basmati 7.58 11.24
Sindh-81 9.0 17.0 Basmati-6129 9.36 16.19
Faisalabad-85 8.0 10.3 DR-83 9.07 14.70
Rawal-87 9.3 18.0 KS-282 9.86 16.67
Pak-81 8.7 15.8 Basmati-385 7.14 8.42
Sariab-92 10.0 19.5 Pakhal 9.29 11.56
Inqalab-91 7.2 20.7 Basmati-370 7.43 9.79
Bakhtawar 11.0 21.0 IR-6 8.62 11.31
Adapted from Rashid et al. (35)

SUMMARY

Above review indicates that boron performs many functions in cell walls (30)
and cellular activities (11). Boron deficiency renders decrease in cell wall
plasticity (21) leading to failure of newly divided cells to enlarge (15). As far
as plasma membrane is concerned, adequate level of boron stops the
accumulation of phenolics and ceases the oxidation of components of
+
plasma membranes. Further it is also involved in the generation of H
ATPase, which is a driving force for ion uptake. Hence, integrity and
functionality of plasma membranes is ensured with adequate supply of boron.
The nitrogen fixation ultimately correlates with nitrogenase activity. Oxygen
sensitivity of nitrogenase is well known and relates to toxicity of O2 species
such as O2- and H2O2 (13). Therefore, activity of O2- and H2O2 scavenging
enzymes may play an essential role in protecting nitrogenase against toxic O2
species. Since nitrogenase activity is dependent on the development of
effective nodules, any alteration in nodule development inhibits the
nitrogenase activity. Boron accelerates nitrogenase activity through effective
nodule development for nitrogen fixation. Plants reproductive growth is
ceased with the deficiency of boron. This retarding growth is considered due
to the low phloem mobility of boron. In brief, the formation of B complexes
with the constituents of cell wall and plasma membrane as well as with the
phenolic compounds is a major reason to affect the physiological functions of
boron.

Pollard et al. (31) reported several impairments as a result of B deficiency,

such as sugar transport, cell wall synthesis, signification, cell wall structure,
carbohydrate metabolism, RNA metabolism; respiration, indole acetic acid
(IAA) metabolism, phenol metabolism, and membrane integrity.

J. Agric. Res., 2009, 47(3)

 Role of boron in plant growth 335

The above discussion reveals unique physiological role played by boron in

plants. However, despite such numerous effects of B, no evidence has yet
been presented that B is an enzyme constituent or it has a direct role in
enzyme activities. It is also not clear whether above mentioned changes are
precursor of direct functions of B or the changes are of an indirect nature.
Therefore, a lot of research activities are required to have an insight whether
role played by boron nutrition is primary or secondary.

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