Boron Deficiency:

A global Micronutrient Deficiency in Soils

Globally, B deficiency is one of the most prevalent
micronutrient deficiencies. Even some reports indicate that
B deficiency is the most widespread micronutrient
deficiency around the world (Shorrocks, 1997, Plant and
Soil. 193, 121–148)

Boron should be included in the fertilizer formulations

and fertilizer programs to ensure balanced fertilization and
balanced crop nutrition.
 Three major mechanisms leading to boron
deficiency in plants
I. Boron Adsorption
(high pH soils, low organic matter, high clay)

II. Boron Leaching

(usually in soils with low pH,
low organic matter and high sand)

III. Immobility of Boron in Plants

(poor transport of B from vegetative to generative
organs/youngest shoot parts)
 Soil B adsorption increased with increasing
solution pH, reached a maximum around pH 9
(soil from a salinity lab in Riverside)

Soil pH

Goldberg et al., 2004, Vadose Zone Journal 3:676–680

Soil Acidity and Alkalinity
(Regions with red color : acidic soil)

Acidity pH Alkalinity
 Soil Acidity and Alkalinity
(Regions with red color : acidic soil)

Boron deficiency commonly occurs in low pH soils

due to leaching and lime-induced B deficieny

Acidity pH Alkalinity
 Root growth is highly sensitive to Al
Wheat seedlings grown in soil with
decreasing aluminum concentrations

High Al No or Low Aluminum

https://www.agric.wa.gov.au/soil-acidity/effects-soil-acidity Photo: S Carr

Liming of Low pH Soils and Root Growth of Barley

Slide: Dr. L. Kochian

Liming in Low pH Soils

https://www2.ctahr.hawaii.edu/ctahr2001/Soil/
 Effect of liming
2.5 t lime/ha in 1984, barley photographed in 2000
Wongan Hills, W. Australia

Limed Unlimed
Liming is effective to correct pH for a certain
period.

However, liming may induce boron deficiency:

The lime-induced B deficiency is closely related to
adsorption of B onto AI(OH)3 which forms following
liming practice

Gupta et al, 1985; Can. J. Soil Sci., 65: 381-409; Goldberg, 1997, Plant and Soil
193: 35–48, 1997; Hajduk et al., 2017, J. Elementology, 22: 411-426
Liming is a common agronomic practice against Al toxicity

Acid Soils Lime

Al3+ Ca2+
Clay + 3 CaCO3 + 3 H2O Clay Ca2+ + 2 Al(OH)3 +3 CO2
Al3+ Ca2+
pH >6.0

Precipitation of Al
AI(OH)3

Redrawn from Rengel Z (2002)

Encyclopedia of Soil Science
Boron content of newly developed needles in
Norway spruce seedlings grown in a growth
medium with CaCO3 and CaSO4 addition

Lehto and Malkönen, 1994, Plant Soil 163: 55-64

Why plants need B?
Major Physiological Changes
under B Deficieny
• Cell wall structure and function
• Structural stability and functional integrity of
membranes
• Root nutrient uptake
• Photosynthesis and photoxidation
• Nitrogen Fixation
• Generation of ROS (reactive O2 species)
• Phenol metabolism
• ….
 Boron is essential for the Structure and
Function of Cell Walls (Structural Element)
• Cell wall contains a unique B binding sugar “Apiose”
that is likely the site of B attachment and function in
cell walls
• >90% B is associated with Cell Walls
• B deficiency reduces cell wall extensibility: resulting in
shortened, thickened cell walls

Brown et al 1994; 2002).

 Boron and
Cell Wall Stability and Function
• Boron is known to be primarily
involved in establishment of
networks of pectic polysaccharides
in cell walls, particularly with
rhamnogalacturonan-II (RG-II).

• Maintenance of mechanical stability

and proper function of cell walls is
closely related to the formation of
RG-II-B complexes

Funakawa and Miwa, 2015, Front. Plant Sci.

About 90% of the RG-II pectin
complex is associated with boron.

RG-II pectin complex is primarily

required for pollen germination,
pollen tube elongation and pollen
function

(Dumont et al., 2014; Ann. Bot; Funakawa

and Miwa, 2015, Front. Plant Sci).
 Boron and Root Growth
Rapid and substantial physiological changes occur in
plants when exposed to low B conditions. Probably,
the most common and immediate reaction of plants
to low B supply is the impairment in root growth.

Published reports using model plants show very quick

stop in root cell elongation upon transfer of plants to B-
deficient growth conditions (within hours)

Marschner, 2012; Oiwa et al., Soil Sci. Plant Nutr. 59: 621–62
 Effect of Boron Nutrition on
Root Growth of Squash Plants
50
+Boron
Root Elongation (mm)

40

30 -Boron +B Resupply

20

10 +B - Boron

0
0 10 20 30 40
Boron treatment, h
Marschner, 2012
Growth of plants with low (-B) and
adequate (+B) boron

Canola

-B

+B
Growth of plants with low (-B) and
adequate (+B) boron

Soybean

-B
+B
Growth of plants with low (-B) and
adequate (+B) boron

Corn

-B +B
Growth of plants with low (-B)
Corn
and adequate (+B) boron
Soybean
Canola

-B
+B

-B
+B
-B +B
 Boron Content of Plant Species Grown
Under Identical Environmental Conditions.

Plant Species Boron Content

(µg.g dwt)
Wheat (Nable, 1990)
Cv ‘Sterling’ 6
CV ‘Sahara’ 23
Timothy 15
Tobacco 30
Brussels Sprout 50
Carrots 75
Sugar Beet 102
From Gupta, 1979
Shoot growth of soybean plants
depending on B supply

Low B Medium B Adequate B

Shoot and root growth of soybean plants
depending on B supply

Low B Medium B Adequate B

 Sunflower roots Adequate boron
under low B

Low boron

Very low boron

Low boron Adequate boron
 Effect of Boron Nutrition on
Root Growth of Squash Plants
50
Adequate Boron
Root Elongation (mm)

40

30
Low Boron +B Resupply
20

10 +B Low Boron

0
0 10 20 30 40
Boron treatment, h

Marschner, 2012
Alfalfa root penetration in Al-toxic soil with (+B) and
without B (-B)
Root Penetration

+B -B alfalfa roots penetrated

approx. 38 cm into the high-
Al toxic subsoil when soil
treated with B (left), while the
roots without B penetrated
only 15 cm
15 cm

A good boron nutrition

38 cm is protective against Al toxicity

Modified from Lenoble et al 1996, Plant Cell Environ.

Effect of different
Reduced B treatments
ascorbiconacid
concentration
of reduced ascorbic acid in sunflower young
in young leaves
leaves
mol Ascorbate · g-1FW

6 -B
+B

0
0 4 6 8 10

Plant Age, days

Cakmak and Romheld, 1997, Plant and Soil
Ascorbic acid (AA helps to mitigate
root damage under low B
Root Growth inhibition
in Boron-Deficient or
Aluminum-
Stressed Squash May
Be a Result of
Impaired Ascorbate
Metabolism

Lukaszewski and Blewins,

1996, Plant Physiol
 Al binding within the cell wall is
the first step leading to Al toxicity

Published reports show that adequate B supply

to plants is required for minimizing root Al
uptake and binding of Al to root cell walls
 Histochemical determination of Al by
hematoxylin staining of the rapeseed roots.

Stronger blue color (more Al) in the roots under B

deficiency in rapeseed/canola

Riaz et al., 2018, J. Plant Interactions, 13: 270-276

Shoot and root growth of trifoliate orange seedlings
under different B and Al treatments

Low B - Al Low B +Al +B - Al +B +Al

Riaz et al., 2018, Ecotox. Environ. 153: 107-115

 Negativity of the apoplast (Al-binding sites) in
root cell walls is higher in B-deficient plants

B deficiency increases high amount of negative charges in

cell walls for Al binding (due to high level of un-methyl
esterified pectin) (Horst et al., 1999; Li et al., 2017; Franco et
al., 2002; Stass et al., 2007)

A further reason might be that B is competitive with Al for

binding sites in cell walls. B supply is shielding Al binding
sites.
 Boron and Nitrogen Fixation

Boron is also required for better nitrogenase

activity (N-fixing enzyme), nodule weight and
nodule formation in legume plants
 BORON and N-FIXATION

B deficiency-induced impairments in N fixation

Adequate Boron Low Boron

Bolanus et al., 1994, Plant Physiol.

 BORON and N-FIXATION

Nodule Weight Nitrogenase

Nodule Weight and Nitrogenase Activity

in Pea Roots with and without B supply
Bolanus et al., 1994, Plant Physiol.
Boron affects nitrogenase activity
in roots of soybean plants

W: irrigated

WB: irrigated,+B

WS: Water Stress

WSB: Water Stress, +B

Bellaloui, 2011, Amer. J. Plant Sci.

Boron also protects plants from
high light/ radiation damage
Growth of sunflower plants with sufficient and deficient
boron supply under low and high light conditions.
Plants under low boron supply are quickly
damaged when exposed to high light intensity

Cakmak and Romheld, 1997, Physiol. Plant.

Growth of sunflower plants with sufficient and deficient
boron supply under low and high light conditions.
Plants under low boron supply are quickly
damaged when exposed to high light intensity
Cakmak and Romheld, 1997, Physiol. Plant.
B-Deficient plants are highly sensitive to high light

Growth of sunflower plants with sufficient and deficient

boron supply under low and high light conditions.
Plants under low boron supply are quickly
damaged when exposed to high light intensity
Cakmak and Romheld, 1997, Physiol. Plant.
 Superoxide Generation and
Photooxidative Damage

Photooxidative damage to
membrane and chlorophyll can
be expected in B-deficient
leaves as a result of enhanced
photogeneration of toxic oxygen
Shaded Unshaded
free radicals caused by impaired
utilization of light energy in
photosynthesis

Cakmak and Romheld,

1997, Physiol. Plant.
Flower organs require more B than
vegetative organs
 Pollen formation
is probably the most critical step in plant
development that determines the size of
yield formation,

and pollen formation stage is the most B

sensitive phase of life cycle of plants
Wheat anthers with low and adequate B

Adequate B B Deficiency

Photo courtesy R. Bell and P. Brown

B Deficiency: Flower Abortion in Walnut
Flower Death

1 inch

Adequate B Low B

Photo courtesy R. Bell and P. Brown

Boron concentrations of flowers in canola , mg kg-1

38 (control) 19 15
Low B
increased
abnormal
flowers and
dropped floral
buds

Boron concentration in these

lines were reduced by
eliminating B-transporter gene
Zhang et al., 2017, Plant, Cell and Environ. 40: 1819–1833
 High B demand for pollination
Effect of increasing B supply on vegetative
and reproductive growth of red clover
B supply Shoot DW Flowers Seed yield
(mg/kg (g/pot) (No/pot) (mg/pot)
soil)
0 12.8 0 0
0.25 13.0 6 0
0.5 12.6 13 0
1.0 12.3 37 430
2.0 12.3 37 1190
(Sherell, 1983, New Zeal. J. Agric. Res., 26, 197-203)
 Fertilization and Boron

Pollen Tube

Function of pollen
tubes is significaly
impaired in B-deficient
plants

http://lcbiology.webnode.com/
 Growth and length of pollen in lilly plants
at different B concentrations

RG-II Pectin
Complex

Boron Concentration
mg/l
https://mammothmemory.net/biology/
From Marschner, 2012 plants/sexual-reproduction-in-plants
Effect of Boron on Seed Formation in Pea

-BORON

+BORON

Bonillo et al. 2009, in Plant Phys. Dev.

 Sunflower head
with good B
nutrition

Sunflower head
with low B
nutrition

https://www.flickr.com/photos/
25531432@N02/2807733867/
Flower organs require more B than vegetative organs
Critical Value B for different organs (mg/kg)
More
16 38 sensitive

1 16
Less
sensitive
www.dreamstime.com

Dell and Huang, 1997, Plant Soil, 193:103–120.

Boron Nutrition and
Nutrient Uptake
Boron Nutrition and Nutrient Uptake
Adequate boron nutrition is required for maintenance
of nutrient uptake by roots, probably due to its direct
contribution to the structural and functional integrity of
root cell membranes
.
Plant roots secrete protons through a plasma-
membrane bound H+-ATPase system, and thus,
generate a membrane potential gradient across the
membranes. The generated membrane potential is
used as a driving force for active uptake and
transport of mineral nutrients
(Marschner, 2012, Marschners' Mineral Nutrition of Higher Plants, 3rd Ed9
The plasma membrane H+-ATPase generates the gradient of
electrochemical potential of H+ across the plasma membrane

Soil Particle

H+ Outside Cell
K+

Inside Cell
K+
H+
Membrane Potential: Driving force for
-120 mV K Uptake

Modified from plantcellbiology.masters

Effect of Boron on Em of Sunflower Root Cells

H3BO3 caused a gradual hyperpolarization of the root

cell membranes. The mean Em value of -174.9 mV
became nearly 5 mV more negative within the first 3 min of
exposure to B
Schon et al 1990, Plant Physiol
 Driving Force for K uptake
Adequate B

Low B

Changes in membrane potential (driving force for K

uptake) of root cell membranes (within minutes)

Schon et al. (1990, Plant Physiol. 93: 566-571).

Changes of the plasma
membrane ATP hydrolysis
activity (a) and H+ transport
(b) in the roots of coffee
seedlings treated with
increasing B supply .

De Souza et al., 2022, J. Plant Nutr.

45: 2123-2134
Effect of increasing B
supply on plasma
membrane H+-ATPase
activity in different young
Citrus trees

Ferreira et al., 2021, Ann Appl Biol.

178:293–303.
 Shoot concentrations of K and Ca in tomato
plants grown in nutrient solution upon root and
foliar applied boron

Davis et al., 2003, J. Amer. Soc Hort. Sci

Relationships between leaf B and leaf K in sweet orange
grafted onto rootstocks of rangpur lime, sunki mandarin, shingle citrumelo
grown on an oxisol (pH:5.7) in Brazil
Leaf K, g kg-1

Leaf B, mg kg-1

Mattos et al., 2017, J. Integ. Agricult.

Root nutrient uptake of 47 days old alfalfa (Medicago sativa
cv. Diane) plants grown under varied B concentrations
under hydroponic conditions.

B Treatment Nutrient Uptake mg /g root

uM K Mg P S Ca

B 0 9.0 ±2.9 1.52 ±0.26 2.49 ±0.65 3.35 ±0.36 8.4 ± 1.5
0.25 9.8 ±0.9 1.48 ±0.22 2.51 ±0.36 3.56 ±1.00 9.2 ± 1.8
1 24.7 ±2.3 2.02 ±0.24 3.79 ±0.23 3.12 ±0.83 11.7 ± 1.1
25 53.5 ±3.3 1.55 ±0.06 4.85 ±0.92 2.55 ±1.13 13.5 ± 1.2

Ceylan et al, 2022

 Boron Deficiency-Induced
Membrane Leakage (K+-Efflux)

800 low B
K+ Efflux, µg g-1 FW

+B Potassium absorbed
600 by the roots is
released (efflux) again
400 into soil if roots are
exposed to low B
200 nutrition

0
0 4 6 8 10

Plant Age, days

Cakmak et al 1996, Physiol Plant
Phosphorus (P) uptake of root tips of faba
bean and maize plants grown with
adequate and deficient supply of boron

Marschner 2012
Boron Fertilization
How we should fertilize crop plants with B
to achieve highest agronomic impact,
foliar or soil ?
Main issue to consider:
phloem mobility of boron
In most of the plant species B
is NOT phloem mobile
 Boron Mobility

It is generally believed from older literature that B

is phloem immobile in plants (Oertli and
Richardson, 1970), and this is indeed true for most
plant species.
However, it has been recently shown that in plants
with sugar alcohols (sorbitol, mannitol etc.) in
phloem, B is freely phloem mobile (mainly plant
species in the order of Rosales) (see Brown and
Shelp, 1997)

Practical importance?
B Spray to low-B plants

B Spray to
low B plants
 Limited or No
B Spray to B-Transport to
Low-B Plants New Growth

New
Growth
B

B
2 weeks after

In most of the
plant species
 .
Only 3.2 % of the B taken up by the leaves was
translocated within the plants 240 days after spray

Boron transport REDISTRIBUTION (10B) DAYS AFTER SPRAYING

in citrus (%) 30 120 240

NEW PARTS
(grown after spraying)
--- 0.6 2.5

Sprayed Part PARTS SPRAYED 100 99.4 96.8

OLD PARTS
(protected during spraying)
0 0.1 0.7

% Total Translocated 0 0.5 3.2

Boaretto, et al. 2007, in Advances in Plant and Animal

Boron Nutrition, Pages: 125-131
It is obvious that agronomic effectiveness of foliar B
spray on shoots is minimal (but NO effect on roots)

Foliar B-fertilizers are effective only on the tissues

sprayed, and the B absorbed by the leaves cannot be
translocated into new growthimost of the crop plants

Soil Fertilization…..
 Soil B-Fertilization:
Soluble vs less soluble B fertilizers

Almost all commonly available soil-applied B fertilizers

are highly soluble, leading high risk of leaching and
seedling damage. Probably, major reason for high
prevalence of B deficiency in humid regions and acidic
soils is related to leaching of B from soil profile.

Leaching losses may result in high risk of B deficiency

later in the season. The fact, that reproductive organs
have higher demand for B than the vegetative organs,
indicates high importance of slow-release B-fertilizers.
Solubility of B-Fertilizers
(highly soluble)

50 % Col+50 % Borax

(less soluble)

Da Siva et 2018, Soil Sci. Soc. Am. J.

Higher solubility= higher leaching risk
Borax

Colemanite

Da Silva et 2018, Soil Sci. Soc. Am. J.

Conclusion
Some concerns about foliar spray of B for improving B
demand of plants. B existing in leaf tissue cannot be
translocated in the new growth to sustain/maintain and
development (and pollination). Timing of foliar B spray !
Providing both early- and late-season (less soluble) boron
products seems to be of great importance in soils,
especially in low pH soils
It is important to maintain a sufficient B supply to
plants in order ensure better K uptake and achieve
better response to K fertilization.

Even in high K-soils, a low B nutrition of plants may

reduce K uptake from soils
Since boron is not
mobile in most plant
species, continuous
availability of boron
around roots is
required during the B B
entire growth stage
B B
 Boron in the leaves
B or boron in the roots
Shoot
Root cannot be transported
B to the newly growing
root tips.

continuous availability of boron around

roots is required during the entire growth
stage….slow-B-releasing fertilizers