Pearl millet as an alternative crop for boosting productivity of salt

affected soils
Ravikiran KT1, Jogendra Singh2, Vijayata Singh2, Sanjay Arora1 and Anoop Kumar
Dixit1
1
ICAR-Central Soil Salinity Research Institute, Regional Research Station, Lucknow –
226 0002, Uttar Pradesh, India
2
ICAR-Central Soil Salinity Research Institute, Karnal – 132 001, Haryana, India
Corresponding email: Ravi.KT@icar.gov.in
Introduction
Soil salinity is a major issue that affects agricultural productivity worldwide. The
total area of salt-affected lands globally is 932.2 million hectares - 351.2 million hectares
are saline and 581 million hectares are sodic. In India, around 6.74 million hectares of
land are salt-affected, which is 2.1% of the country's geographical area. Out of this, 2.96
million hectares are saline and the rest (3.77 million hectares) are sodic (Mandal et al.
2018). Around 2.35 million hectares of salt-affected soil can be found in the Indo-
Gangetic plains of India, out of which 0.56 million hectares are saline and 1.787 million
hectares are sodic. Uttar Pradesh alone has 1.35 million hectares of sodic soil, which is
35.6% of the total sodic soil in the country (Arora and Sharma 2017). Salinity increases
the accumulation of ions like Na+ and Cl- in the plant root zone, which results in
decreased water potential in the rhizosphere and leads to osmotic stress. Unless these
salts are leached to lower soil horizons, either naturally or through human intervention,
plants will be forced to take up harmful levels of Na+ and Cl-. This impacts plant
growth and development, as Na+ in particular competes with K+ at various cation
transport channels. On the other hand, soil sodicity/alkalinity disrupts the physical
structure of soil by weakening its stability and lowering hydraulic conductivity,
aeration, and infiltration rates. At higher pH levels, some nutrients become deficient
(nitrogen, calcium, magnesium, iron, manganese, copper, and zinc), while others rise to
toxic concentrations (boron and molybdenum). As a result, plant growth under sodicity
is constrained by both physical and chemical stresses. Calcium deficiency severely
affects root growth, hindering nutrient and water uptake, and exacerbating nutrient
deficiencies already experienced by plants under alkalinity. Poor hydraulic conductivity
and disturbed capillary movement of water lead to either waterlogging or drought.

Pearl millet (Pennisetum glaucum (L.) R. Br.) is a small-grained cereal crop that is
diploid (2n = 2x = 14), cross-pollinated (protogynous), and belongs to the C4 family. It
has a large genome size of approximately 2.4 Gb and is known for its high tillering. It is
typically grown in marginal lands of arid and semi-arid regions of Sub-Saharan Africa
and India. Pearl millet is a hardy crop that can tolerate low moisture, high temperature,
and nutrient deficiency, making it suitable for challenging environments. It is the sixth
most important cereal crop, providing calories for approximately 90 million people in
Africa and Asia. Due to its nutrient density, particularly in iron and zinc, pearl millet is
considered a valuable candidate for combating micronutrient malnutrition. For this
reason, it has been declared the International Year of Millets by the FAO for 2023 upon
the recommendation of India, the world's largest producer of millets. In India during
2021-22, pearl millet production was 9.62 million tonnes with a productivity of 1239 kg
ha-1, which is higher than the world average of 1229 kg ha-1. Despite being categorized
as a moderately salinity tolerant crop (Mass, 1993), previous studies have shown a
significant reduction in seed germination, plumule length, root and shoot lengths, fresh
weights, leaf number, and seedling vigor due to salt stress. The symptoms of salt injury
include chlorosis and necrosis resulting from decreased chlorophyll and carotenoid
content, leading to decreased photosynthesis. Salt stress during the vegetative stage
results in a reduction in shoot length, biomass, test weight, and ultimately grain yield
(up to ~60%). However, most of these studies are based on the seedling or early
vegetative stages, particularly under saline stress. There is very limited literature
available on the effect of sodicity on pearl millet growth and development, with only
one report published. To date, no salt-tolerant pearl millet varieties have been released
in India.
Effect of salt stress on pearl millet
In contrast to other crops, such as rice and maize, where the effect of salinity on
plant growth and development is limited to certain phenological stages (seedling and
reproductive stages in rice and vegetative stage in maize), pearl millet seems to suffer
from salt stress at all growth stages starting from seed germination to maturity. The
effects of salinity on pearl millet growth and development are summarized in Table 1.
At germination, there will be a significant reduction in germination followed by the
death of germinated seedlings. Seedlings show reduced vigour, root and shoot lengths,
root and shoot fresh weights and dry weights. At biochemical level decrease in
chlorophyll content was also observed in the seedlings. Sneha et al (2013) observed a
decrease in chlorophyll content and an increase in Catalase activity under salt stress
with no change in Ascorbate peroxidase activity. At the vegetative stage, there will be a
reduction in leaf number and area of emerged leaves. Reduced plant height and stem
girth were also observed under salinity. At the tillering stage, the number of tillers will
be reduced. All of these will ultimately result in reduced biomass/straw yield and grain
yield. For instance, a mean yield reduction of 16.0%, 37.1% and 64.4% were registered
when saline irrigation water of 3, 6 and 9 dS/m was applied in comparison to the best
available water (Sheoran et al., 2016).

The response of pearl millet under sodicity is poorly understood. There is only
one report which demonstrated the reduction of grain yield of an average of 37% at pH
9.0 and 77% at pH 9.4 relative to pH 8.5 (Sharma et al., 2011). However, the genotypes
used in the study were very few, i.e., eight, of which six are near isogenic lines (NILs),
making the net number of genotypes only three. A preliminary experiment at our
institute revealed that pearl millet is sensitive to soil sodicity (pH ~ 9.6). At the
vegetative stage, a reduction in plant height, stem diameter, leaf width and root
biomass was observed under sodicity. Furthermore, roots are more affected by sodicity
than the shoots.
Genetic variability for salt tolerance in pearl millet

Appreciable variability was reported for response of pearl millet under salt
stresses. For instance, among the wild relatives of pearl millet, Panicum clandestinum,
commonly called Kikuyu grass was studied by Muscolo et al (2003) who observed a
decrease in growth and leaf length with an increase in salinity. However, the genotype
showed tolerance to salinity up to 100 mM resulting of unaffected carbohydrate
utilization. Pearl millet varieties Babapuri, WCC-75, and ST-II were found more tolerant
than other varieties studied and could survive at 12 dS/m (Sharma and Gill, 1992). This
tolerance was attributed to restriction in the uptake of Na to the stem, particularly
nodes and internodes, increased diffusion resistance, decreased transpiration, increased
proline content and lesser reduction in Nitrate reductase activity. Pearl millet hybrids
HHB 223, I-IHB 272, HJ-IB 146, ICMA 004444 × IP 13150 and ICMA 03222 × ICMV
05777 were found to the promising showing lesser yield reduction and lower Na/K
ratio in the shoot with the increasing irrigation water salinity. Markhana et al (2019)
proposed the hybrid ICFH -15 and ICFH -16 as suitable for dry matter yield (dry fodder
purpose) while accessions ICFH-5, ICFH-2 and 7 for higher grain yield under saline
irrigation conditions. Several other tolerant sources identified by various studies are
presented in Table 7.4.2.

Table 1. Effect of salinity on various growth stages of pearl millet

Effect Reference

Reduced Germination Gundalia et al., 1992; Krishnamurthy et al.,

Seedling death
Reduced seedling vigour
Reduced root length, fresh
weight and dry weight
Reduced chlorophyll content
Reduction in leaf number
2007; Yakubu et al 2010; Sharma et al., 2011
Saxena and Kolarkar, 1981
Ali and Irdis et al, 2015
Muscolo et al., 2003; Ali et al., 2004; Yakubu et
al., 2010; Bukhari et al., 2012
Sharma et al., 2011; Sharma et al., 2014
Muscolo et al., 2003; Rahim et al., 2020
 Reduction in flag leaf area/leaf
area/length
Reduction in shoot fresh & dry
weight
Reduction in plant height/ shoot
length
Reduction in grain yield
Reduction in biomass/straw
yield
Reduction in stem girth and
number of tillers
Table 2. Salinity tolerant sources of pearl millet
Tolerant Genotypes
A1/3
Babapuri, WCC-75, and ST-II
93613, KAT/PM-2 Kitui, and Kitui
ICMV-95151, ICMV-95490 and Gana
10876, 10878, 18406, 18570, ICMV-93753, ICMV-94474
HTP 94/54, CZI 9621, IP 3757
Maiwa
HHB 223, I-IHB 272, HJ-IB 146, ICMA 004444 x IP 13150
and ICMA 03222 x ICMV 05777
Bukhari et al., 2012; Radhouane, 2013; Rahim et
al., 2020
Bhakt et al., 2000; Yakubu et al., 2010; Ali and
Irdis, 2015
Alam and Naqvi, 1991; Gundalia et al., 1992;
Yakubu et al., 2010; Hussain et al., 2010;
Bukhari et al., 2012; Radhouane, 2013; Rahim et
al., 2020; Hajlaoui et al., 2021
Gundalia et al., 1992; Radhouane, 2013; Sharma
et al., 2014; Toderich et al., 2018; Sheoron et al.,
2016
Alam and Naqvi, 1991; Gundalia et al., 1992;
Sharma et al., 2011; Rahim et al., 2020; Hajlaoui
et al., 2021
Rahim et al., 2020
Reference
Ashraf and McNeilly,
1987
Sharma and Gill, 1992
Ashraf and McNeilly,
1992
Bhakt et al., 2000
Ali et al., 2004
Krishnamurthy et al.,
2007
Yakubu et al., 2010
Sheoron et al., 2016
 HHVBC Tall and IP 19586 Toderich et al., 2018
ICFH -15 and ICFH -16 (fodder) Makarana et a., 2019
ICFH-5, ICFH-2 and 7 (grain yield)
IP22269 Hajlaoui et al., 2021

Mechanism of salt tolerance in pearl millet known so far

Three studies have elucidated the salinity tolerance mechanism in pearl millet
(Sharma and Gill, 1992; Sharma et al., 2011; Sharma et al., 2014). Under salt stress with
high Na+ ions in the root zone, plants try to
restrict the uptake of Na+ by reducing
transpiration or increasing the diffusion
resistance. The Na+ ions that escape into the
plant system will be further curtailed in the root
system. Once this gate is breached, the plant
tries to localize the Na+ in the stem, particularly
in the nodes and internodes. After this, Na+ will
be transferred to older leaves to be shed at a
later stage of plant life. Simultaneously, plants
accumulate proline in younger leaves as an
osmotic adjustment mechanism. However, this
mechanism is applicable to salinity, whereas no
clear mechanism was proposed for sodicity.

Conclusion

There are very few studies available on the effects of salt stress on pearl
millet. The majority of these studies focus on the seedling stage, which may not be an
accurate representation of the plant's overall tolerance to salt stress. Although some
studies have been done on pearl millet genotypes at the reproductive stage, these
studies have evaluated only a small number of entries, which does not adequately
reflect the plant's genetic variability. Moreover, researchers have given limited attention
to how pearl millet responds to sodicity/alkalinity, which requires further
investigation.

References

Alam, S.M. and Naqvi, S.S.M., 1991. Effect of salinity on growth and nutrient content of
Pearl millet grown in desert sand and gravel. Pakistan Journal of Botany, 23(4), pp.223-
226.

Albassam BA (2001) Effect of nitrate nutrition on growth and nitrogen assimilation of

pearl millet exposed to sodium chloride stress. Journal of Plant Nutrition, 24, pp.1325–
1335

Ali, G.M., Khan, N.M., Hazara, R. and McNeilly, T., 2004. Variability in the response of
pearl millet [Pennisetum americanum (L.) Leeke] accessions to salinity. Acta Agronomica
Hungarica, 52(3), pp.277-286.

Ashraf M (1994) Breeding for salinity tolerance in plant. Crit Reviews in Plant Science, 13,
pp.17–42

Ashraf M, McNeilly T (1987) Salinity effects on five cultivars/ lines of pearl millet
(Pennisetum americanum [L.] Leeke). Plant Soil 103, pp.13–19

Ashraf M, McNeilly TM (1992) The potential for exploiting variation in salinity

tolerance in pearl millet (Pennisetum americanum [L.] Leeke). Plant Breeding 104, pp.234–
240

Bukhari, I.A., Khalid, S., Waheed, T., Ahmed, S., Jabeen, F. and Riaz, M.U., 2012. Effect
of NaCl on the morphological attributes of the pearl millet (Pennisetum
glaucum). International Journal of Water Resources and Environmental Sciences, 1(4), pp.98-
101.
Cuin, T.A., Miller, A.J., Laurie, S.A. and Leigh, R.A., 2003. Potassium activities in cell
compartments of salt‐grown barley leaves. Journal of Experimental Botany, 54(383),
pp.657-661.

Hajlaoui, H., Akrimi, R. and Hajlaoui, F., 2021. Screening field grown pearl millet
(Pennisetum glaucum L.) genotypes for salinity tolerance in the North of Tunisia. IOP
Conference Series: Earth and Environmental Science, 904, p.012046

Hussain, K., Ashraf, M. and Ashraf, M.Y., 2008. Relationship between growth and ion
relation in pearl millet (Pennisetum glaucum (L.) R. Br.) at different growth stages under
salt stress. African Journal of Plant Science, 2(3), pp.023-027.

Jehan-Bakht, B., Banarus Khan, M. and Shafi, M., 2000. Studies of pearl millet under
salinity stress at early growth stage. Pakistan Journal of Biological Science, 3(10), pp.1577-
1579.

Krishnamurthy, L., Serraj, R., Rai, K.N., Hash, C.T. and Dakheel, A.J., 2007.
Identification of pearl millet [Pennisetum glaucum (L.) R. Br.] lines tolerant to soil
salinity. Euphytica, 158, pp.179-188.

Maas, E.V., 1993, August. Testing crops for salinity tolerance. In Proc. workshop on
adaptation of plants to soil stresses (Vol. 94, pp. 234-247). Lincoln, NE: University of
Nebraska.

Makarana, G., Yadav, R.K., Kumar, A., Kumar, R., Sheoran, P., Kushwaha, M. and
Yadav, T., 2019. Physiological, Biochemical and Yield Traits of Pearl-millet (Pennisetum
glaucum L.) Accessions under Saline Irrigation. Journal of Soil Salinity and Water
Quality, 11(1), pp.98-107.

Muscolo, A., Panuccio, M.R. and Sidari, M., 2003. Effects of salinity on growth,
carbohydrate metabolism and nutritive properties of kikuyu grass (Pennisetum
clandestinum Hochst). Plant science, 164(6), pp.1103-1110.
P. C. Sharma & K. S. Gill (1992) Effect of salinity on yield and ion distribution in pearl
millet genotypes. Arid Soil Research and Rehabilitation, 6(3), pp.253-260, DOI:
10.1080/15324989209381319

Radhouane, L., 2013. Agronomic and physiological responses of pearl millet ecotype
(Pennisetum glaucum (L.) R. Br.) to saline irrigation. Emirates Journal of Food and
Agriculture, pp.109-116.

Rahim, Z., Parveen, G., Gul, S. and Rehman, K., 2020. Ameliorating Effects of Salt Stress
(KCl, NaCl) on Growth and Germination Parameters of Pearl Millet (Pennisetum
americanum). Pakistan Journal of Agricultural Research, 33(4), pp.951-956.

Sharma, P.C., Sehgal, D., Singh, D., Singh, G. and Yadav, R.S., 2011. A major terminal
drought tolerance QTL of pearl millet is also associated with reduced salt uptake and
enhanced growth under salt stress. Molecular Breeding, 27, pp.207-222.

Sharma, P.C., Singh, D., Sehgal, D., Singh, G., Hash, C.T. and Yadav, R.S., 2014. Further
evidence that a terminal drought tolerance QTL of pearl millet is associated with
reduced salt uptake. Environmental and experimental botany, 102, pp.48-57.

Sheoran, R., Kumar, A., Sharma, P.C., Lata, C., Mann, A., Gupta, S.K., Yadav, D. and
Sharma, D.K., 2016. Impact of salinity on physiological and biochemical traits in pearl
millet. In: 4th International Agronomy Congress, Nov 22- 26, 2016, New Delhi, India.

Toderich, K., Shuyskaya, E., Rakhmankulova, Z., Bukarev, R., Khujanazarov, T.,
Zhapaev, R., Ismail, S., Gupta, S.K., Yamanaka, N. and Boboev, F., 2018. Threshold
tolerance of new genotypes of Pennisetum glaucum (L.) R. Br. to salinity and
drought. Agronomy, 8(10), p.230.

Yakubu, H., Ngala, A.L. and Dugje, I.Y., 2010. Screening of millet (Pennisetum glaucum
L.) varieties for salt tolerance in semi-arid soil of Northern Nigeria. World Journal of
Agricultural Sciences, 6(4), pp.374-380.