Received: 13 September 2023 Revised: 29 December 2023 Accepted: 16 February 2024

DOI: 10.1002/fft2.381

REVIEW ARTICLE

Strategies to promote the dietary use of pigeon pea (Cajanus

cajan L.) for human nutrition and health

Jun Wu Qian Zhou Chenhaojin Zhou Ka-Wing Cheng Mingfu Wang

Shenzhen Key Laboratory of Food Nutrition

and Health, Institute for Advanced Study,
Shenzhen University, Shenzhen, P. R. China
Correspondence
Qian Zhou, Shenzhen Key Laboratory of Food
Nutrition and Health, Institute for Advanced
Study, Shenzhen University, Shenzhen, P. R.
China. Email: qzhou@szu.edu.cn
Mingfu Wang, Shenzhen Key Laboratory of
Food Nutrition and Health, Institute for
Advanced Study, Shenzhen University,
Shenzhen, P. R. China. Email:
mfwang@szu.edu.cn
Funding information
National Natural Science Foundation of China,
Grant/Award Number: 32101935; Shenzhen
Science and Technology Program,
Grant/Award Number:
ZDSYS20220117155800001; Guangdong
Basic and Applied Basic Research Foundation
of China, Grant/Award Number:
2022A1515012098
Abstract
Pigeon pea (Cajanus cajan L.) is one of the most important legume crops grown in
subtropical and tropical regions with high nutritional quality. However, the consump-
tion of pigeon pea is very limited worldwide owing to its high level of anti-nutrients,
hard texture, and unpleasant characteristic taste. The present review summarized the
nutritional profile and health benefit of pigeon pea with special attention to its bioac-
tive phytochemicals (e.g., cajanin, cajanol, and cajaninstilbene acid). Furthermore, some
aspects that can help promote the application of pigeon pea in foods were highlighted
from the perspectives of physical (i.e., thermal, extrusion, and ultrasound-assisted),
chemical (i.e., chemical modification and enzymatic hydrolysis), and biological treat-
ment (i.e., sprouting and fermentation). The primary impact of physical methods was
confined to the mitigation of anti-nutrients. Chemical approaches were less explored
and mainly targeted on the modification of starch and protein in pigeon pea. In contrast,
the biological methods were found to decrease the level of anti-nutritional factors,
increase the production of bioactive phytochemicals, and improve the sensory proper-
ties of pigeon pea. Therefore, biological methods, especially sprouting techniques, are
considered to be more promising.

KEYWORDS
anti-nutrients, cajanin, fermentation, pigeon pea, sprout

1 INTRODUCTION
Pigeon pea (Cajanus cajan L.), also known as red gram and Arhar, is
one of the oldest crops and ranked as the sixth most important edible
legume in the world (Figure 1). It is drought-tolerant and nitrogen-
fixing, growing in tropical and subtropical regions. The global area of
pigeon pea cultivation is around 5 million hectares, with South Asia,
the Caribbean, and Africa being the major planting regions (Varsh-
ney et al., 2012; Zhao et al., 2020). Noteworthy, pigeon pea is largely
considered an orphan crop, which has been domesticated by human
beings but not fully utilized so far. Orphan crops are usually fit to low-
input agricultural conditions. They not only play a critical role in local
agriculture and ecology but also possess high potential to meet the
requirements of food security and nutritional health worldwide (Ye &
Fan, 2021). Thus, growing interests have been aroused in the study
of pigeon pea in recent years. Unlike other legume crops, pigeon pea
is tolerant to environmental stimuli, unique in nutritional profile, high
in biomass productivity, and friendly in the retention of soil nutrients
(Odeny, 2007). In addition, current literature data have exemplified the
health benefits of pigeon pea, as well as pointing out that pigeon pea

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2 WU ET AL .
F I G U R E 1 The appearance of pigeon pea in different forms. Source: Adapted from
https://specialtyproduce.com/produce/Pigeon_Peas_11653.php and https://www.academia.edu/31170922/Indian_Food_Composition_Tables.
is an important income source for smallholders in some undeveloped
countries (Sharma et al., 2019). As a perennial pulse from Fabaceae
family, pigeon pea works as a perfect source of protein supplement for
traditional cereal-based diets to avoid protein deficiency. Owing to the
expensive cost of animal-derived proteins and rapid growing popula-
tions, protein deficiency might be a common and serious problem in
some pigeon pea–planting regions (Odeny, 2007). Pigeon pea has one
of the highest protein contents (average 25 g/100 g) in the legume fam-
ily. In addition, it is abundant in calcium, phosphorus, potassium, and a
variety of essential amino acids, which determines that it has a good
nutritional effect. It is also rich in polyphenols, which can contribute
to its health benefits. Hence, from this perspective of view, the study
of pigeon pea to promote its application in food products is of great
importance.
Pigeon pea has been used in tropical and subtropical regions as food
and animal feeds for centuries. It is directly eaten after cooking in
South Asia and can also be applied as a functional ingredient in biscuits
(Adeola & Ohizua, 2018; Okpala et al., 2013; Tiwari et al., 2011), fla-
vored snacks (Ohizua et al., 2017; Rampersad et al., 2003), and noodles
(Yadav et al., 2011). As an example, it is reported that the fortification
of 5% pigeon pea flour into cassava flour can produce a satisfier texture
with higher protein content and better flavor (Rampersad et al., 2003).
Moreover, pigeon pea has been extruded into a mixture with maize to
make technologically low-cost grain–legume snack, a product that can
both alleviate the problem of malnutrition (especially for children) and
potentially bring benefits to local families (Fasoyiro et al., 2017). More-
over, they were processed and consumed as traditional foods (e.g., bean
cake and pudding) in some developing countries (Fasoyiro et al., 2010).
However, the dietary consumption of pigeon pea is very limited across
the world, and its application in food industry is also rare by far. At least
three reasons may account for this issue. The first and foremost rea-
son is that pigeon pea contains a high level of various anti-nutrients
(Singh & Kumar, 2022), such as enzyme inhibitors, phytic acid, and
lectins (Singh, 1988; Soetan & Oyewole, 2009). As the name implies,
anti-nutrients are the compounds/components that block the absorp-
tion of nutrients in foods. Even more, some anti-nutrients are toxic
to human body to some extent, including inducing malnutrition and
micronutrient deficiencies, and impairing the integrity of the intestinal
mucosa (Soetan & Oyewole, 2009). Among enzyme inhibitors, trypsin
inhibitors and amylase inhibitors are the leading problems. Trypsin
inhibitors can reduce the digestibility of proteins by forming complex
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with trypsin and chymotrypsin. Similarly, amylase inhibitors can inhibit
the activity of amylase, thereby reducing the digestion rate of carbohy-
drates (Avilés-Gaxiola et al., 2018; Khattab & Arntfield, 2009; Vagadia
et al., 2017). Phytic acid is able to trap minerals (e.g., iron, calcium,
and zinc), inhibiting their intracellular uptake. Lectins can also interfere
with the absorption of minerals, bind to the lining of the gut, and conse-
quently lead to gut inflammation (Coulibaly et al., 2011; Feizollahi et al.,
2021; Thakur et al., 2019). Moreover, pigeon pea is extremely hard,
which makes cooking it a tough job. In particular, it takes up to 24 h
to prepare a pigeon pea meal from its dried seeds when applying tra-
ditional thermal processing methods (Samtiya et al., 2020). In addition,
pigeon pea has a characteristic taste that distinguishes it from other
legumes. This special taste from our perspective certainly restrains its
application in our daily diets. Considering the economic and nutritional
potential of pigeon pea, it is meaningful to seek practical strategies
to enhance its edibility and fully use the beneficial functions of its
bioactive components.
In the past years, various efforts have been made to pave the way for
the application of pigeon pea in food products, primarily by targeting
on anti-nutrients. Such efforts include physical (i.e., thermal, extru-
sion, and ultrasound-assisted), chemical (i.e., chemical modification and
enzymatic hydrolysis), and biological (i.e., sprouting and fermentation)
treatments (Figure 2). By using these methods alone or in combina-
tion, it is effective in decreasing the level of anti-nutritional factors, as
well as ameliorating undesirable flavor and hard texture. Furthermore,
previous research also helped to figure out the changes of bioactive
phytochemicals (e.g., chemical structure and concentration) in pigeon
pea under different treatments, except affecting its basic physicochem-
ical properties (Rizvi et al., 2022). Therefore, the nutritional value and
health benefits of pigeon pea might be improved after processing,
which probably added up its potentials in food application. The present
review aims to address these issues and summarize the advances in this
field.
2 NUTRITIONAL PROFILE OF PIGEON PEA
Pigeon pea is highly recommended ascribing to its abundance of
nutrients. As summarized in Table 1, pigeon pea is rich in carbohy-
drates (19.46–62.68 g/100 g), proteins (16.76–29.26 g/100 g), and
fat (1.08–5.31 g/100 g), as well as crude fibers (1.20–8.63 g/100 g)
 26438429, 0, Downloaded from https://onlinelibrary.wiley.com/doi/10.1002/fft2.381 by Cochrane Philippines, Wiley Online Library on [19/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
WU ET AL . 3

TA B L E 1 Nutritional profile of pigeon pea in different forms.

Green seed
Category (unmatured) Dry seed (matured) Dal (peeled off) Ref.
(1) Basic elements (g/100 g)
Carbohydrates 19.5–23.9 62.7–62.8 55.23–60.4 Ade-Omowaye et al. (2015); Adenekan
et al. (2018); Akubor (2017);
Protein 7.2–21.0 16.8–29.2 19.7–24.6
Athanas (2022); Choi et al. (2020);
Fat 0.9–2.3 1.1–2.4 1.4–5.3 Faris (1987); Longvah et al. (2017);
Crude fiber 2.3–8.2 3.2–8.6 1.2–3.2 Rizvi et al. (2022); Torres et al.
(2006); Usda (2019)
Ash 1.4–1.6 3.2–4.0 2.0–3.5
Moisture 64.0–65.9 8.2–9.3 6.7–10.5
(2) Amino acids (g/100 g)
Leucine 0.2 1.4–1.6 0.3–0.5 Ade-Omowaye et al. (2015); Adenekan
et al. (2018); Athanas (2022);
Lysine 0.3 0.3–1.5 0.3
Kunyanga et al. (2013); Longvah
Valine 0.2 0.9–1.1 0.2–0.6 et al. (2017); Usda (2019)
Isoleucine 0.2 0.6–0.8 0.2–0.3
Histidine 0.2 0.7–0.8 0.1–0.3
Threonine 0.2 0.7–0.8 0.2–0.4
Methionine 0.02 0.2–0.3 0.04–0.1
Tryptophan 0.05 0.2–0.6 0.04–0.1
Phenylalanine 0.4 1.7–8.7 0.4–0.5
Arginine 0.3 1.0–1.4 0.2–0.3
Cystine 0.04 0.3–0.5 0.03–0.2
Alanine 0.3 0.9–1.0 0.1–0.2
Tyrosine 0.1 0.5 0.1–0.4
Glycine 0.2 0.7–0.8 0.2–0.5
Serine 0.3 1.0–1.1 0.2–0.3
Aspartic acid 0.5 2.0–2.3 0.2–0.5
Glutamic acid 1.0 3.1–5.0 0.1–1.1
Asparagine / 0.2
Proline 0.2 0.9–1.1 0.1–0.2
(3) Vitamins (mg/100 g)
Folic acid (VB9 ) 0–0.4 100 / Athanas (2022); Ellong et al. (2015);
Kunyanga et al. (2013); Longvah
Ascorbic acid 15.1–569.2 0–4.8 /
et al. (2017); Torres et al. (2006);
Niacin (VB3 ) 2.1–2.2 2.4–3.0 2.1 Usda (2019)
Thiamin (VB1 ) 0.2–0.4 0.3–0.7 0.5
Pyridoxine (VB6 ) 0.2–0.7 0.3–0.4 0.2
Riboflavin (VB2 ) 0.1–0.2 0.1–0.4 0.1
Vitamin E (VE) 0.4–1.8 2.3 0.8
Pantothenic acid (VB5 ) 0.7 1.56 1.3
Biotin (VB7 ) 2.9 0.7 0.3
Total folates (VB9 ) 94.2 229 108
(4) Fatty acids (mg/100 g)
Palmitic acid (C16:0) 181 236–262 222 Kunyanga et al. (2013); Longvah et al.
(2017); Usda (2019)
Oleic acid (C18:1) 13.0–15.1 78.6–92 89.9
Stearic acid (C18:0) 30.0 40.0 35.2
Linoleic acid (C18:2) 427–835 537–778 600
Linolenic acid (C18:3) / 35 /

(Continues)
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4 WU ET AL .

TA B L E 1 (Continued)

Green seed
Category (unmatured) Dry seed (matured) Dal (peeled off) Ref.
(5) Minerals (mg/100 g)
Potassium 552–616 1290–1390 1395 Akubor (2017); Athanas (2022); Faris
(1987); Kunyanga et al. (2013);
Phosphorus 127–141 312–367 328
Kuraz Abebe (2022); Longvah et al.
Calcium 42.0–94.6 80.5–200.9 16.3–110.0 (2017); Rizvi et al. (2022); Torres
Magnesium 57.0–113.7 20.4–183.0 78.9–146.0 et al. (2006); Usda (2019)

Sodium 2.6–5.0 0.3–89.7 18.0

Iron 1.2–4.6 5.2–33.5 2.9–5.0
Zinc 1.0–2.5 2.3–58.5 2.6–3.0
Copper 0.1–1.4 1.0–17.0 11.1–1.3

FIGURE 2 Strategies to pave up the application of pigeon pea in food products.

(Adamu & Oyetunde, 2013; Ade-Omowaye et al., 2015; Adenekan
et al., 2018; Akubor, 2017; Amarteifio et al., 2002; Athanas, 2022; Choi
et al., 2020; Rizvi et al., 2022). Moreover, pigeon pea is sufficient in
essential amino acids, including leucine (0.2–0.6 g/100 g), lysine (0.3–
1.5 g/100 g), valine (0.2–1.1 g/100 g), isoleucine (0.2–0.8 g/100 g),
histidine (0.1–0.8 g/100 g), threonine (0.2–0.8 g/100 g), methionine
(0.02–0.3 g/100 g), tryptophan (0.04–0.6 g/100 g), and phenylalanine
(0.4–8.7 g/100 g) (Adenekan et al., 2018; Athanas, 2022; Kuraz Abebe,
2022). The high level of leucine and lysine makes pigeon pea espe-
cially suitable for people with obesity, diabetes mellitus (Pedroso et al.,
2015), and people who need more calcium such as athletes (Chromiak
& Antonio, 2002). For fatty acids, palmitic acid (181–262 mg/100 g)
is the main saturated fatty acid in pigeon pea, whereas oleic acid
(13–92 mg/100 g) is the most abundant unsaturated fatty acid (Ade-
Omowaye et al., 2015; Longvah et al., 2017). In the process of analyzing
published data, we noticed the concentration of macronutrients in

WU ET AL .
pigeon pea varied significantly among different sources. This might
be ascribed to their variation in genotypes and maturities (Table 1).
For instance, Choi et al. evaluated the nutritional value of 82 differ-
ent pigeon peas cultivated in Korea and found that they obviously
diversed in the content of crude protein (16.76–22.61 g/100 g), crude
fiber (4.70–8.63 g/100 g), and dietary fiber (12.98–33.19 g/100 g)
(Choi et al., 2020). In addition, the interesting thing is that green
seed (unmatured), dry seed (matured), and dal (peeled off) (Figure 1)
are all editable in different countries. Consequently, researchers from
different countries might use different forms of pigeon pea as their
tested materials. Moreover, a pigeon pea can be divided into embryo,
cotyledon, and seed coat. Its proteins, carbohydrates, lipids, and ash
are mainly distributed in the cotyledon, whereas the dietary fiber is
mainly presented in the seed coat (Salunkhe et al., 1986). This also
probably contributes to its nutrient variances. Given that the pro-
tein content in pigeon pea is particularly higher than other legume
crops (Ade-Omowaye et al., 2015; Saxena et al., 2002), pigeon pea
is a potential plant protein supplement for human and animal diets.
This is very important and meaningful, considering that pigeon pea is
mostly planted in developing countries located in tropical and subtrop-
ical regions (Bekele, 2007). With a fast-growing population, people may
have increased demands for proteins. Pigeon pea offers sufficient pro-
teins at a lower price compared to meat. In addition, its proteins can be
easily digested in the body (Akporhonor et al., 2006). Therefore, pigeon
pea can offer the body with enough nutrients (especially proteins and
amino acids) and act as a well-balanced diet, either as a staple food or
in combination with other grains and beans.
Except macronutrients, pigeon pea is also a good source of micronu-
trients. For vitamins, pigeon is abundant in vitamin B1 (VB1 , thiamin;
0.23–0.74 mg/100 g), vitamin B2 (VB2 , riboflavin; 0.11–0.39 mg/100 g),
vitamin B3 (VB3 , niacin; 2.09–2.97 mg/100 g), vitamin B6 (VB6 ,
pyridoxine; 0.24–0.68 mg/100 g), vitamin B9 (VB9 , folic acid; 0.39–
100 mg/100 g), vitamin C (VC, ascorbic acid; 4.80–569.2 mg/100 g),
and vitamin E (VE; 0.39–2.29 mg/100 g), and vitamin A precur-
sor (β-carotene; 0.05–0.36 mg/100 g). Thereinto, Ellong et al. found
that pigeon pea was surprisingly high in VC (569.2 mg/100 g) and
β-carotene (0.364 mg/100 g) contents amongst other 27 crops,
which certainly contributed to the high antioxidant ability of pigeon
pea (Ellong et al., 2015). For minerals, except phosphorus (127–
367 mg/100 g) and potassium (552–1392 mg/100 g), pigeon pea is
characterized by its richness in calcium (Ca; 16.3–200.9 mg/100 g) and
magnesium (Mg; 68–183 mg/100 g) (Athanas, 2022; Kunyanga et al.,
2013; Kuraz Abebe, 2022; Olalekan & Bosede, 2010; Rizvi et al., 2022).
3 HEALTH BENEFITS OF PIGEON PEA AND ITS
BIOACTIVE PHYTOCHEMICALS
3.1 Beneficial effects of pigeon pea
The consumption of pigeon pea is believed to be beneficial and has been
applied in the prevention/treatment of chronic diseases in Asia for a
long history (Hassan et al., 2016). For instance, pigeon pea is widely
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5
used to treat hepatitis, superficial infections, and even measles and
chickenpox in Oman (Al-Saeedi & Hossain, 2015); pigeon pea exerts
potentials in the relief of pains and is considered a traditional medicine
by Indians to treat stomatitis and gingivitis (Ganesan, 2008). Except
that, recent research has also demonstrated the health benefits of
pigeon pea with its advances, summarized in Table 2. Its functional
activities can be basically categorized as (1) antioxidants, which were
considered the major benefits of pigeon pea. Several in vitro DPPH, fer-
ric reducing antioxidant potential (FRAP), and Trolox equivalent antiox-
idant capacity experiments have confirmed the appreciable antioxidant
property of pigeon pea (Cheboi et al., 2019; Choi et al., 2020; Sekhon
et al., 2017). Meanwhile, pigeon pea was also demonstrated to increase
the activity of antioxidant enzymes in cells, including superoxide dis-
mutase, catalase, glutathione peroxidase, and glutathione reductase
(Lai et al., 2012). Intriguingly, the seed coat of pigeon pea was believed
to possess higher antioxidant potential, given that it is hard in texture
and digestion (Rani et al., 2014). In addition, the phenotype with brown
seed coat had higher antioxidant activity than its white coat companion
(Ukom et al., 2023). In addition, the solvents applied in the extrac-
tion of pigeon pea are also important, and hexane extracts showed
the highest antioxidant capability by using the DPPH method (Al-
Saeedi & Hossain, 2015). (2) Anti-inflammatory: Lai et al. (2012) found
pigeon inhibited the expression of pro-inflammatory cytokines, such
as tumor necrosis factor-a (TNF-α), interleukin-1β, and interleukin-
6 (IL-6), in lipopolysaccharide (LPS)-induced macrophage (Lai et al.,
2012), and Hassan (2016) reported the administration of pigeon pea
extracts effectively ameliorated carrageenan-induced inflammation in
Adult Swiss albino mice via reducing TNF-α, IL-6, and IgG serum levels
(Hassan et al., 2016). (3) Anti-hyperglycemic and prevent lipid peroxi-
dation: Uchegbu et al. proved that slightly sprouted pigeon pea extract
obviously lowered the fasting blood glucose levels and decreased lipid
peroxidation in diabetic rats. On this basis, pigeon pea was recognized
as a good dietary supplement for controlling hyperglycemia (Uchegbu
& Ishiwu, 2016); (4) anti-hypertensive: Olagunju et al. (2022) used
pepsin and pancreatic enzymes to digest pigeon pea, and the pep-
tides obtained both showed better inhibitory activity on renin and
angiotensin-I-converting enzyme (ACE).
3.2 Bioactive ingredients in pigeon pea
The health benefits of pigeon pea might be closely associated with
its high content of polyphenols, which exhibit favorable bioactivities.
As reported by Ellong et al. (2015), the total phenolic content in
pigeon pea was 978.6 mg/100 g, which was significantly higher than
that of other crops. Ade-Omowaye et al. (2015) also observed sim-
ilar results and found pigeon pea had high antioxidant activity by
using the FRAP method. Noteworthy, the content of polyphenols might
vary in different genotypes of pigeon peas. Agricultural characteris-
tics of 82 genotypes cultivated in South Korea have been studied for
their phenolic content by DPPH and Trolox assays (Choi et al., 2020),
and positive correlation was found between high polyphenol level
and high DPPH activity. Similar conclusion was achieved by Cheboi

6 WU ET AL .
TA B L E 2 The health benefits of pigeon pea extract and its characteristic phytochemicals.
Extracts/phytochemicals
(1) Extracts of pigeon pea
Water extracts of pigeon pea sprout flour Chemical models
Methanol, hexane, chloroform, ethyl
acetate, and butanol extracts of pigeon
pea
Butanol extracts of pigeon pea
50% ethanol extract pigeon pea
200 and 400 mg/kg hexane extracts of
pigeon pea
(2) Characteristic phytochemicals of pigeon pea
Cajanin
Cajanol
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Models Bioactivities and effects Ref.
- ↑ Total flavonoid compounds and antioxidant activity Ariviani et al.
(2022)
- ↑ α-Amylase and α-glucosidase inhibitory activities
- ↑ Potential to antidiabetic
Chemical models - Total phenol content: Al-Saeedi and
hexane > chloroform > methanol > ethyl Hossain
acetate > butanal > water (2015)
- Total flavonoid content: chloroform > methanol > ethyl
acetate > butanol > water > hexane
- Antioxidant activity: hexane > butanol > methanol > ethyl
acetate > chloroform > water
Chemical models - ↑ Antioxidant activity and total reduction capability Hassan et al.
(2016)
- ↓ Lipid peroxidation
In vitro, 50 µM - ↑ Antioxidant enzyme activity (SOD, CAT, GPx, and GR) Lai et al. (2012)
H2 O2 -treated RAW
264.7 cells - ↓ GSH and MDA production
- ↓ DNA damages
In vitro, 1 µg/mL - ↓ NO and PGE2 level
LPS-treated RAW 264.7
cells - ↓ Inflammatory cytokines, including TNF-α, IL-1β, and IL-6
- Protect against oxidative stress and inflammation
In vivo, - ↓ Swelling of rat paw edema after carrageenan injection Hassan et al.
carrageenan-induced (2016)
rats, and their paw - ↓ Inflammatory cytokines, including TNF-α and IL-6
edema
- ↓ Carrageenan-mediated increase in IgG level
In vivo, adult Swiss albino - ↓ The number of writhing in 20 min observation period
mice with 2% in saline
acetic acid injected - ↑ The antinociceptive effect
In vitro, 0.5 µg/mL - ↓ LPS-activated NO, IL-6, TNF-α, and iNOS Hong et al.
LPS-treated RAW 264.7 (2022)
cells - ↓ The phosphorylation and translocation of NF-κB
- Blockade of NF-κB transcription factor activity
In vitro, bone marrow - ↑ Differentiation and proliferation of BMSCs Zhao et al.
mesenchymal stem cells (2019)
(BMSCs) - Osteogenesis markers were upregulated by cajanin
treatment at both protein and mRNA levels
- The cell cycle regulator proteins were phosphorylated or
upregulated
In vitro, MCF-7 human - ↓ The growth of MCF-7 cells Luo et al. (2010)
breast cancer cells
- Arrest the cell cycle in the G2/M phase and induced
apoptosis via a ROS-mediated mitochondria-dependent
pathway
- Inhibit Bcl-2 expression and induce Bax expression to
disintegrate the outer mitochondrial membrane, causing
cytochrome c release
- Prevent apoptosis
(Continues)
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WU ET AL . 7

TA B L E 2 (Continued)

Extracts/phytochemicals Models Bioactivities and effects Ref.

In vitro, human lung - Fungal cajanol possessed stronger cytotoxicity toward A549 Zhao et al.
carcinoma cells (A549), cells (2013)
noncancer cells
(MC3T3-E1 and RAW
246.7 cells)
In vitro, 6 kinds of bacteria - Strong antibacterial activity towards all bacteria Liu et al. (2011)
(e.g., Escherichia coli and
Staphylococcus aureus) - Inhibit E. coli only by DNA damage

- Inhibit S. aureus by affecting both the lecithin and phosphate

groups on the cellular membrane and DNA
Cajaninstilbene acid Chemical models - ↑ Antioxidant activity Wu et al. (2011)

- ↓ Radical and peroxide generation

- ↓ XOD inhibition and formation of nicked DNA

- ↑ Formation of supercoiled DNA

- ↓ DNA damage
In vitro, 0.1 µg/mL - ↓ Release of inflammatory mediator and cytokines (NO, Huang et al.
LPS-treated RAW 264.7 TNF-α, and IL-6) (2016)
cells
In vitro/vivo, OGD/R and - ↓ Cell death in OGD/R and t-BHP injured SH-SY5Y cells Xu et al. (2021)
t-BHP injured SH-SY5Y
cells, MCAO/R rats - ↓ Infarct volume and neurological deficits in MCAO/R rats

- Neuroprotective activity via AMPK/Nrf2 signaling pathway

In vivo, behavioral despair - ↓ The immobility time of mice in tail suspension test and Zhang et al.
mice models forced swimming test (2020)

- ↑ Antidepressant effects
In vivo, chronic - Reversed the depressive-like behavioral changes
unpredictable mild
stress model - ↑ Antidepressant effects and the effects might be attributed
to regulating TRP metabolism, promoting BDNF
expression and activating Akt/mTOR pathway
In vivo, injury transgenic - ↓ The production and migration of neutrophils and primitive Huang et al.
zebrafish larvae macrophages (2016)

Abbreviations: Akt, Ak strain transforming; AMPK, AMP-activated protein kinase; Bax, BCL2 associated X; Bcl-2, B-cell lymphoma 2; BDNF, brain-derived
neurotrophic factor; CAT, catalase; GPx, glutathione peroxidase; GR, glutathione reductase; GSH, glutathione; IgG, immunoglobulin G; IL, interleukin; iNOS,
inducible nitric oxide synthase; LPS, lipopolysaccharide; MCAO/R, middle cerebral artery occlusion/reperfusion; MCF-7, a breast cancer cell line; MDA,
malondialdehyde; mTOR, mechanistic target of rapamycin; NF-κB, nuclear factor kappa-light-chain-enhancer of activated B cells; Nrf2, nuclear factor ery-
throid 2-related factor 2; OGD/R, oxygen-glucose deprivation/reoxygenation; PGE2 , prostaglandin E2 ; ROS, reactive oxygen species; SH-SY5Y cells, a human
neuroblastoma cell line; SOD, superoxide dismutase; t-BHP, tert-butyl hydroperoxide; TNF-α, tumor necrosis factor-α; TRP, tryptophan; XOD, xanthine
oxidase.

et al. (2019), which explored 55 genotypes grown in Kabete field
(Kenya). Furthermore, some efforts have been contributed in unravel-
ing the polyphenol profile of pigeon pea during the past decade. Like
its leaves, pigeon pea (the seed) mainly has two types of polyphe-
nols, that is, flavonoids and stilbenes. The flavonoids include sev-
eral flavonoid aglycones (quercetin, luteolin, apigenin, isorhamnetin,
pinocembrin, and genistein), flavonoid-O-glycosides (quercetin-3-O-β-
D -glucopyranoside, biochanin A, and genistin), flavonoid-C-glycosides
(orientin, vitexin, and isovitexin), methylated flavonoids (pinostrobin,
cajanol, and cajanin), and prenylated flavonoid (isowighteone). The
main stilbene in pigeon pea is cajaninstilbene acid (CSA) (Gai et al.,
2022; Hassan et al., 2016; Nix et al., 2015; Tungmunnithum &
Hano, 2020). Thereinto, cajanin (18.11 ± 0.27 mg/100 g) and cajanol
(11.64 ± 0.17 mg/100 g) might be the most abundant polyphenols,
whereas CSA is found to be unique in pigeon pea (Tungmunnithum
et al., 2021).
Literature data have highlighted that cajanin is the major anti-
inflammatory compound in pigeon pea and can be a promising

8 WU ET AL .
candidate for the treatment of inflammatory diseases. More specifi-
cally, cajanin attenuated LPS-induced expression of nitric oxide (NO;
IC50 = 19.38 ± 0.05 µM), IL-6 (IC50 = 7.78 ± 0.04 µM), and TNF-α
(IC50 = 26.82 ± 0.11 µM) in RAW 264.7 macrophages (Hong et al.,
2022). This is achieved probably by preventing the phosphorylation
and nuclear translocation of nuclear factor kappa-light-chain-enhancer
of activated B cells (NF-κB). For cajanol, it possesses anti-carcinogenic
potential and can be developed into a new natural anticancer agent
(Anwar et al., 2010; Luo et al., 2010). Cajanol can block the cell cycle
of human breast cancer cell MCF-7 (a breast cancer cell line) in G2/M
phase and induce cell apoptosis via reactive oxygen species–mediated
mitochondria-dependent pathway (Luo et al., 2010); it can induce
severe cell death in human lung carcinoma cells A549 but express
moderate cytotoxicity in noncancer cells such as osteoblast precur-
sor cell line MC3T3-E1 and RAW 264.7 macrophages (Zhao et al.,
2013). In addition, cajanol has anti-bacterial potential, which inhib-
ited Escherichia coli proliferation through DNA damages and prevented
Staphylococcus aureus by affecting lecithin and phosphates on cell mem-
branes (Liu et al., 2011). For CSA, it is found to have antioxidant,
anti-inflammatory, antibacterial, anti-depressant, and neuroprotec-
tive potentials. Compared to resveratrol (a famous antioxidant and
also a typical stilbene), CSA showed higher capabilities in prevent-
ing DNA damages via its antioxidant pathway (Wu et al., 2011). It
inhibited the migration of neutrophils and macrophages in injured
zebrafish larvae through mitogen-activated protein kinases/NF-κB sig-
naling pathway (Huang et al., 2016). Besides, derivatives modified
from CSA showed remarkable antibacterial activity and low cytotoxi-
city against Gram-positive bacteria and methicillin-resistant S. aureus
(Geng et al., 2015). The antidepressant effect of CSA may be real-
ized by regulating tryptophan metabolism, promoting the expression
of brain-derived neurotrophic factor and postsynaptic density protein-
95, and activating the cortical protein kinase B (Akt)/mechanistic
target of rapamycin pathway (Zhang et al., 2020). Recently, our group’s
findings have exemplified the significant neuroprotective activity of
CSA for the prevention and treatment of ischemic stroke. Mechanis-
tic analyses revealed that CSA can regulate AMP-activated protein
kinase/nuclear factor erythroid 2–related factor 2 signaling pathway
so as to decrease oxidative damages and mitochondrial dysfunction
both in vitro (oxygen–glucose deprivation/reoxygenation and tert-
butyl hydroperoxide–injured human neuroblastoma cell line SH-SY5Y)
and in vivo (middle cerebral artery occlusion/reperfusion (Xu et al.,
2021).
4 PHYSICAL STRATEGIES AS A PRIMARY
APPROACH TO INCREASE THE DIETARY
CONSUMPTION OF PIGEON PEA
Being simple, flexible, and convenient, physical strategies have been
applied in the processing of pigeon pea in home cooking and in the sci-
entific research. These physical strategies include thermal, extruded,
and ultrasonic-assisted treatment. Thermal treatment is usually at the
top of the priority list, especially for household wives. Extrusion of
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pigeon pea emerged during the past two decades, which is a favorable
means to promote its application in snack manufacturing and other
packaged foods (Fasoyiro et al., 2017). Moreover, some researchers
have attempted to develop ultrasound-assisted methods for the effi-
cient and green extraction of the functional components in pigeon pea
(Cui et al., 2015; Pan et al., 2021, 2023). Besides, efforts have also been
contributed in the combination of these three physical treatments with
or without other types of pre-processing steps, such as grinding, wall-
breaking, and drying (Pan et al., 2023; Tiwari et al., 2008; Vásquez et al.,
2021).
4.1 Thermal treatment
The basic effect of thermal treatment is to increase the nutritional
value by eliminating harmful substances in pigeon pea. Generally
speaking, legume seeds contain high amounts of anti-nutritional fac-
tors, including trypsin inhibitors, phytic acid, and thiocyanate. These
anti-nutritional factors are usually located in the germ and periph-
eral regions of the seed, interfering with the absorption of nutrients in
human body. The heating process can effectively destroy the structure
of these substances, which consequently improves the digestibility and
absorption rate of nutrients. Moreover, published data have revealed
the effects of thermal processing at different levels (i.e., <100, 100,
120, 160–180◦ C) on the detoxification of tannins, alkaloids, enzyme
inhibitors, cyanogenic glycoside, and hemagglutinin (Kuraz Abebe,
2022; Onwuka, 2006). Of note, moderate heating (<100◦ C) can inac-
tivate enzymes (e.g., lipoxygenase, lipase, amylase, and protease) but
promote the digestibility of proteins. Boiling (100◦ C) decreases the
content of tannins and trypsin inhibitors in pigeon pea, lowers the
foaming capacity, but increases the emulsion stability of pigeon pea
flour. Toasting pigeon pea flour at 100◦ C showed similar trends. After
1 h of toasting, the water and oil absorption capacity slightly increased,
whereas emulsifying activity, foaming capacity, and bulk density sig-
nificantly decreased. Additionally, the toasting process resulted in a
significant reduction in anti-nutritional factors, reducing cyanide levels
by 66% and phytic acid levels by 16.7% (Onimawo & Akpojovwo, 2006).
The combination of soaking (for 18 h at room temperature) and boil-
ing shows better effect than boiling alone, which can make the content
of anti-nutrients below detection (Onwuka, 2006). The most obvious
benefit of toasting at 120◦ C is increasing the content of minerals (such
as Fe, Zn, Ca, and Mg), ash, and proteins. But this might be ascribed
to water loss because toasting significantly reduces the moisture con-
tent. Other than that, toasting had similar effects with boiling (Akubor,
2017).
The performance of more modern household cooking methods
that can offer a higher temperature (e.g., 160 and 180◦ C) such as
microwave on pigeon pea was also investigated. For instance, Sun et al.
found a raised in vitro protein digestion rate of pigeon pea flour after
microwave treatment, probably due to the reduction of β-structure
(e.g., β-fold and β-corner) and the increase in random curl conformation
in the protein. β-Structure was reported to be negatively associated
with the in vitro digestibility of soy proteins, whereas random curling

WU ET AL .
indicated the formation of a more unfolded structure with higher struc-
tural flexibility (Sun et al., 2020). In addition, microwave treatment
can make pigeon pea break into small particles. This action increased
the reaction surface area of pigeon pea, which might help to promote
the protein–protease interaction and thus partly improve the protein
digestibility (Crévieu et al., 1997). Cooking might also affect the sta-
bility and bioavailability of polyphenols (Cao et al., 2021) and other
phytochemicals; however, studies related to this issue are very limited.
Except decreasing anti-nutritional factors, thermal treatment can
modify the texture and flavor of pigeon pea to make it more palat-
able. The fibers of legume seeds become softer and easier to chew after
thermal treatment (Tiwari et al., 2008). Meanwhile, heating also pastes
the starch in the seeds. Maninder et al. found that pigeon pea flours
had a higher pasting temperature, higher viscosity at peak, though, and
breakdown point but lower viscosity at setback point. This provides
pigeon pea with better stability, viscosity, and bonding ability during
cooking, thereby improving its texture, mouthfeel, and offering richer
flavor (Maninder et al., 2007). Furthermore, the coat of legume seed
often contains unpleasant flavor, whereas thermal treatment might
work as a practical dehulling method. By examining the effect of soak-
ing (40–70◦ C), blanching (100◦ C), and roasting (120◦ C), Fasoyiro et al.
(2010) found that soaking at 60◦ C for 30 min efficiently achieved
dehulling of pigeon pea, and the dehulling efficiency was higher than
pure higher temperature processing (e.g., 100 and 150◦ C) (Fasoyiro
et al., 2010). Their results indicated the significance of the soaking
process in dehulling. Similar conclusion was reported by Tiwari et al.
(2008), who found that the highest dehulling rate of pigeon pea was
achieved when pretreated by the hydrothermal method.
4.2 Extrusion
The application of extrusion to legume food processing is its ability
to simultaneously achieve multiple effects, such as heating, expan-
sion, and protein modification. Compared to other physical meth-
ods, extrusion is advanced in providing efficient, comprehensive, and
multifunctional effects with unparalleled industrial processing abili-
ties. Extrusion is often accompanied by high-temperature processing;
therefore, it also has the benefits of thermal treatment mentioned in
Section 4.1 (e.g., the removal of harmful substances, increased nutri-
tional value, and improved texture and flavor). During extrusion, pigeon
pea went upon high pressure and shearing, which resulted in a puffing
effect with a light texture. This helped in the development of pigeon
pea snacks (Banki et al., 2021; Okpala, 2018; Ozioma et al., 2023).
Noteworthy, in the extrusion process, pigeon pea was usually mixed
with other foods or food ingredients. For example, the mixture of 25%
unripe plantain and 75% pigeon pea was extruded at 120 rpm and
100◦ C. Significant increase was observed in the mineral content of the
mixture with a decrease in anti-nutrients (Anuonye, 2012). Banki et al.
investigated the effect of different extrusion conditions on rice and
pigeon pea mixture. Results showed that the optimal variables were
screw speed at 220 rpm, mixture moisture content at 30%, mix com-
position at 25% pigeon pea and 75% rice, with the maximum expansion
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9
index at 9.98, bulk density at 0.12 g/mL, and water absorption index
at 6.41. This formulation was better in terms of physical properties,
minerals, and essential amino acid contents (Banki et al., 2021).
The temperature during extrusion is a critical control point, which
showed significant effect on the color of the materials and affected
the acceptability of food products. In particular, Chakraborty et al.
investigated the effect of extrusion on color by evaluating the head
temperature (160–200◦ C), moisture content (12%–24%), and screw
speeds (100–140 rpm) of a pigeon pea–millet mixture. The optimum
processing conditions were as follows: head temperature 171◦ C, mois-
ture content 24%, screw speed 104 rpm, and bean flour content 19%. In
addition, temperature was found to be the most important parameter
influencing color when the composition of the mixture was determined
(Chakraborty et al., 2014). In another case, Kaushal et al. (2019) added
5% jackfruit flesh flour into the mixture of rice and pigeon pea flour
and concluded that extrusion temperature significantly changed water
solubility index, color parameters (L* and b*), water, and oil absorp-
tion capacity. Furthermore, due to the high temperature and pressure
condition during the extrusion process, the enzymatic activities in the
pigeon pea mixture were effectively reduced, thus extending the shelf
life of the food products (Adeleye et al., 2020). On this basis, extrusion
can be used as an industrial processing method with high utility (Shukla
et al., 2021), which is suitable to promote the application of pigeon pea
in food industry.
4.3 Ultrasound-assisted treatment
Ultrasound is applied in food processing primarily owing to its non-
thermal effect, which achieved the purpose of food manufacturing
but ultimately retained the nutrients and appearance of foods. In
addition, compared with traditional heat treatments, ultrasonics usu-
ally requires shorter processing time and lower temperature. This
partly promoted the application of ultrasound technologies, espe-
cially ultrasound-assisted green extraction applications in the study of
pigeon pea. Cui et al. (2015) developed a deep eutectic solvent-based
microwave-assisted extraction method based on ultrasound for the
extraction of genistin, genistein, and apigenin from pigeon pea roots,
which increased the extraction rate by 1.50-, 1.27-, and 1.64-folds,
respectively (Duan et al., 2013). Pan’s group from Northeast Forestry
University has also made some contributions in this field. In 2021, Pan
et al. first developed a CO2 -based switchable-solvent method based on
ultrasound (as a tool for quick clean extraction) for the enrichment of
phytochemicals from the leaves of pigeon pea, and the results showed
that the extraction rates of vitexin and CSA were 1.06 and 1.49 times
higher than those of the traditional method (65% ethanol-ultrasound-
assisted extract), respectively. Moreover, the system can be recycled
five times while ensuring a stable extraction rate, making it a sustain-
able method (Pan et al., 2021). In 2023, Pan et al. further developed
CO2 -responsive surfactant-functionalized multi-walled carbon nan-
otubes with a microwave ultrasound assistant method for extracting
a variety of active substances from various sources, including pigeon
pea leaves. The content of target compounds was increased by 1.1–20

10
times. Meanwhile, the green surfactant can be recycled with no signif-
icant loss in six operation cycles, which is an environmentally friendly
method (Pan et al., 2023).
5 CHEMICAL STRATEGIES AS A PRACTICAL
APPROACH TO IMPROVE THE FUNCTION OF
PIGEON PEA FLOUR
Pigeon pea can be ground into flour and then applied as a food ingre-
dient to other products (Yadav et al., 2011). As starch is the major
component in pigeon pea flour, special attentions have been paid to
it (Maninder et al., 2007). In particular, compared to potato and rice
starch, pigeon pea starch has a higher content of amylose and a lower
level of amylopectin. This results in a low starch digestibility index but
high yield stress and solid-like behavior (Acevedo et al., 2020). There-
fore, pigeon pea starch has the potential to act as a supplement to
traditional starch. However, pigeon pea starch has relatively low solu-
bility and functional properties compared to traditional flours available
in the market. These properties strictly restrain its application in food
industry. Direct modifications of starch by chemicals and enzymatic
hydrolysis have been conducted in recent years. Noteworthy, protein
is another main component in the pigeon pea flour. Enzymes might also
affect proteins in the pigeon pea flour and digest them into functional
peptides, potentially improving its nutritional value.
5.1 Chemical modification
Chemical modification (e.g., hydroxypropylation and acetylation) of
pigeon pea starch has been proposed. These modifications are
reported to increase its properties, including decreasing gelatinization
point, but increasing storage stability and moisture holding capac-
ity. For instance, Lawal et al. prepared hydroxypropyl derivatives of
pigeon pea starch and characterized its functional properties by com-
parison with natural pigeon pea starch. After hydroxypropylation, the
shape and size of the starch granules were basically unchanged, but
the swelling parameter was increased. This was because, after hydrox-
ypropylation, the introduced hydroxyl groups can cause inter- and
intramolecular repulsion, enhancing the water absorption ability of
modified starch and making it more suitable for the preparation of food
thickeners and hydrogels. Owing to similar reason, the turbidity of the
starch was decreased with the increment of the hydroxypropyl level.
This is helpful in improving the transparency and light transmittance
rate, which might consequently promote the customer acceptance of
pigeon pea starch (Lawal, 2011). For acetylation, the main character-
istic is that it promoted the content of amylose (which may be due to
the molecular de-polymerization of branched amylopectin after acety-
lation), which was slower in digestion comparing with amylopectin. This
increase in the amylose level of pigeon pea starch is able to decrease
glycemic index, thereby improving glucose tolerance and insulin level
in diabetic patients (Olagunju et al., 2020).
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WU ET AL .
5.2 Enzymatic hydrolysis
Enzymatic reaction can improve the texture and flavor of the food
by directional transformation of the components in the food while
maintaining its original characteristics. The application of enzymatic
hydrolysis in pigeon pea has helped in dehulling, shortening cooking
time, and improving the properties of starch and proteins. As exam-
ples, Sangani et al. optimized the enzymatic hydrolysis conditions of
pigeon pea by using a combination of xylanase, pectinase, and cellu-
lase. They found a 13.81% increase in dehulling rate and a 19.77%
decrease in cooking time with higher overall acceptability (Sangani,
Patel, Davara, et al., 2014) (Sangani, Patel, Bhatt, et al., 2014). Benítez
et al. (2021) conducted response surface methodology analysis so as to
improve starch digestibility and obtain desirable functional attributes
at nutritional level (e.g., viscosity, water retention capacity, and solubil-
ity). In addition, recent efforts have also been contributed in developing
pigeon pea into functional foods for malnourished people through the
degrading of proteins into small functional peptides. Olagunju et al.
(2018) obtained a variety of bioactive peptides with different molec-
ular weights by the utilization of enzymes (e.g., lipase, trypsin, pepsin,
and trypsin) on pigeon pea. Some of the peptides exerted obvious
antioxidant properties, along with favorable inhibitory activity against
ACE/renin and reduced systolic blood pressure in hypertensive rats
(Olagunju et al., 2018). Later, the same research group fractionated
these peptides by membrane dialysis and investigated their functional
activities (e.g., antioxidant, anti-hypertensive, and antidiabetic) by in
vitro models (Olagunju et al., 2021).
6 BIOLOGICAL STRATEGIES AS A PROMISING
APPROACH TO PROMOTE THE NUTRITIONAL
BENEFITS OF PIGEON PEA
In the review of literature data, we noticed the trend of employing
biological methods, especially sprouting and fermentation, as a promis-
ing strategy in this field. Sprouting entails several notable advantages,
including enhanced nutritional value, reduced anti-nutrient content,
and improved taste and flavor (Akubor, 2017; Chinma et al., 2022;
Rizvi et al., 2022). Additionally, it contributes to an augmentation of
bioactive components, thereby increasing antioxidant properties and
improving storage stability (Boateng et al., 2008; Chinma et al., 2022).
In the context of fermentation, its prominent role lies in leveraging
diverse microbial fermentation processes to enhance taste, eliminate
undesirable flavors, and even intensify the depth and complexity of
taste and flavor profiles (Permana et al., 2020).
6.1 Sprouting
Sprouts are a new concept that has emerged in recent years. Sprouts
are basically germinated seeds and tender seedlings, which are usually
harvested after 3–5 days from seed hydration (Ebert, 2022; Rouphael

WU ET AL .
FIGURE 3 The effects and benefits of pigeon pea sprouting.
et al., 2021). The sprouting process is reported to reduce the content
of anti-nutrients while increasing the level of several nutrients and
bioactive components (mainly polyphenols) in pigeon pea (Figure 3).
Moreover, sprouting may offer better texture and flavor with low-
ered pasting viscosity but improved nutrient (e.g., protein and starch)
digestibility, owing to the enzymatic degradation of macromolecules
into small molecules in the legume (Akubor, 2017; Chinma et al.,
2022; Rizvi et al., 2022; Uchegbu & Ishiwu, 2016). In addition, this
consequently promotes the bioactivity of pigeon pea. For instance,
Uchegbu and Ishiwu reported that dietary intake of germinated pigeon
pea reduced fasting blood glucose level in diabetic rats, dramatically
decreased lipid peroxidation, but promoted glutathione level in liver
(Uchegbu & Ishiwu, 2016). Noteworthy, the decrease of anti-nutrient
levels is sometimes the main purpose of legume sprouting. Soaking
and germination can reduce the level of phytates by 16.54% and
57.97%, respectively, while decreasing the level of tannins by 22.94%
and 63.05% (Rizvi et al., 2022). Phytates are usually water-soluble and
located in the seed coat; thus, they might dissolve into water dur-
ing soaking. The natural husking of pigeon pea during sprouting can
also contribute to the lower content of phytates because they are
dominantly located in the seed coat. The reduced content of tannins
probably ascribes to the formation of complex with proteins (Chinma
et al., 2022; Olagunju et al., 2020; Rizvi et al., 2022).
In terms of macronutrients, soaking and germination promoted the
content of pigeon pea proteins by 6.34% and 15.41%, respectively.
This might be owing to the rapid synthesis of proteins as a large quan-
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11
tity of metabolic enzymes is needed during sprouting (Chinma et al.,
2022; Rizvi et al., 2022). Furthermore, in vitro protein digestibility was
also improved by 11.64% and 28.34% in soaking and germination, and
the increased level was higher than in thermal treatment (Akporhonor
et al., 2006). According to Khatoon and Prakash (2006), after activa-
tion by the sprouting process, proteases increased the flexibility of
protein chain via structural modification of proteins and promoted the
degradation of stored proteins into small chain peptides and free amino
acids, thereby raising the in vitro protein digestibility of legumes. More-
over, germination reduces the level of total and digestible starch in
pigeon pea, probably owing to the degradation of starch into oligo- and
mono-saccharides by activated amylase. This process provides energy
for the growth of sprouts and makes germinated pigeon pea suitable
for individuals with special dietary needs. Germination is also found to
increase the level of resistant starch, which might be prebiotic (Akubor,
2017; Chinma et al., 2022). In addition, sprouting can increase water,
ash, and mineral contents but decrease fat level in pigeon pea (Aku-
bor, 2017). Collectively, sprouted pigeon pea has great potential to be
developed into healthy foods.
In the study of bioactive phytochemicals, sprouting is reported to
increase the concentration of total phenols (by 0.97% in soaking and
5.34% in germination) and total flavonoids (by 11.13% in soaking and
63.88% in germination) (Rizvi et al., 2022). Two reasons might be
responsible for these changes. One is that germination promotes the
degradation of insoluble polymers into soluble small molecules (e.g.,
simple phenolic acids) and converts bounded polyphenols into their

12
free form. The other is that germination may disrupt the cell wall dur-
ing the degradation of insoluble polymers, leading to the release of
cytosol phenolic compounds. This biosynthetic and releasing process
certainly adds to the accumulation of total phenols and total flavonoids
(Boateng et al., 2008), thereby increasing pigeon pea’s metal chelating
activity by 64.16% and antioxidant capability by 76.15%. In addition,
prolonged germination time (48 h) and higher temperature (35◦ C) are
preferred due to the increased content of these bioactive phytochem-
icals (Chinma et al., 2022; Rizvi et al., 2022; Sharma et al., 2019).
Another interesting thing is that the mature leaves of pigeon pea plant
are high in polyphenol contents with more polyphenols discovered in
them (Kong et al., 2010; Lai et al., 2012; Liu et al., 2022; Wu et al., 2009).
For instance, pinostrobin, cajanololactone A, and longistylin A and C
are identified in the mature leaves, but not in the seed so far. Among
them, pinostrobin is used as a quality indicator of pigeon pea leaves for
its high content and characteristic existence (Wang & Li, 2022). High
level of bioactive phytochemicals often suggests high potential in the
prevention/treatment of pathological conditions and diseases, includ-
ing anti-osteoporosis (Lai Lijin, 2020; Linan, 2014; Liu et al., 2022),
anti-inflammatory (Huang et al., 2016; Lai et al., 2012), anti-cancer
(Abo-Zeid et al., 2018; Luo et al., 2010), anti-bacterial (Kong et al.,
2010), anti-obesity (Solverson et al., 2022), and anti-viral (Nwodo et al.,
2011; Zu et al., 2010). On this basis, we discover a research gap that no
data are reported for pigeon pea about the changing trend of any single
polyphenols during sprouting, microgreens, and mature leaves. Future
work is warranted in this field.
Except direct consuming, sprouted pigeon pea has been reported to
be incorporated in cookies and in the extraction of bioactive ingredi-
ents. In one study, pigeon pea was soaked in water for 10 h, germinated
for 4 days, dried at 50◦ C, milled, and 100-mesh sieved. Then the
sprouted pigeon pea flour was blended with fermented sorghum and
cocoyam flours (Okpala et al., 2013). The purpose of this study was
to replace wheat flour with the blend so as to restrain energy intake.
In another study, Balasubramanian et al. (2013) extracted urease (also
known as urea amidohydrolase EC 3.3.1.5) from sprouted pigeon pea
(germinated for 5 days at 37◦ C in sterile environment) and confirmed
its antifungal activity.
6.2 Fermentation
Literature data have highlighted the great potential of fermentation in
the attenuation of anti-nutrients in pigeon pea (Figure 4). Specifically,
fermentation is reported to decrease the content of γ-galactoside by
82% and phytic acid by 48% and reduce the activity of trypsin inhibitor
by 39% (Anaemene & Fadupin, 2022; Balogun et al., 2021; Torres et al.,
2006). However, the impact of fermentation on nutrients is contro-
versial in different reports. Anaemene and Fadupin (2022) found that
natural (wild) fermentation under 28◦ C for 72 h could increase the con-
tent of Ca by 56% in pigeon pea. Balogun et al. (2021) demonstrated
an increase in ash, water content, and crude protein but a decrease in
pH value, crude fiber, crude fats, and carbohydrates when fermented at
26438429, 0, Downloaded from https://onlinelibrary.wiley.com/doi/10.1002/fft2.381 by Cochrane Philippines, Wiley Online Library on [19/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
WU ET AL .
30 ± 2◦ C for 96 h. Meanwhile, Torres et al. (2006) reported that natural
fermentation of pigeon pea under 42◦ C for 48 h significantly increased
the level of fats and total soluble carbohydrates but decreased the con-
tent of proteins, dietary fibers, minerals (e.g., Ca, Na, K, Mg, and Zn), and
vitamins (e.g., VB2 and VE). It seems the temperature during fermenta-
tion is a critical parameter, but more research is needed to get a final
conclusion. In addition, a recent study revealed the microbes involved
during natural fermentation of pigeon pea. They included Lactobacil-
lus species, Bacillus species, Enterobacter faecalis, E. coli, S. aureus, and
Pseudomonas aeruginosa, with Lactobacillus and Bacillus species being
the predominant species (Balogun et al., 2021; Olasupo et al., 2016).
Thus, the Lactobacillus and Bacillus species were then selected to per-
form a controlled fermentation in pigeon pea, which received higher
scores in color and texture attributes than those produced in natural
fermentation (Balogun et al., 2021). Furthermore, Permana et al. used
molds (i.e., Aspergillus oryzae and Rhizopus oligosporus) in the controlled
fermentation to prepare pigeon pea sauce. The results showed that
the mold-fermented sauce was acceptable in color, flavor, and overall
acceptance (Permana et al., 2020).
Fermented pigeon pea has been reported as a food ingredient in
nutrients for fortified pasta and complementary food for children
(Bolaji et al., 2021; Permana et al., 2020; Torres et al., 2006). To be
specific, pasta added with 5%–10% fermented pigeon pea had a longer
cooking time and higher water absorption rate, with improved level of
proteins, fats, dietary fiber, minerals, VE, and antioxidant activity. The
protein digestibility was raised significantly in this pigeon pea pasta
(Torres et al., 2006). In another study, pigeon pea was mixed with plan-
tain, fermented, and squeezed to a final product with high nutritional
and sensory properties. The mixture was also high in edible proteins,
which was suitable in the preparation of complementary foods, espe-
cially for malnourished children in some developing countries (Bolaji
et al., 2021; Permana et al., 2020). Furthermore, pigeon pea was applied
in the production of nattokinase, a serine fibrinolytic enzyme with
strong fibrinolytic activity. It was found that pigeon pea fermented
with Bacillus subtilis 14715 for 32 h had the highest nattokinase activ-
ity. The following animal study of B. subtilis 14715 treated pigeon pea
suggested it could lower diastolic and systolic blood pressures in spon-
taneously hypertensive rats, but non-fermented pigeon pea showed
less effects (Lee et al., 2015). This work highlighted the potential of
fermented pigeon pea as a functional ingredient in the prevention of
hypertensive diseases.
7 CONCLUSIONS AND FUTURE PERSPECTIVES
In the present review, we summarized the potential strategies to pro-
mote the application of pigeon pea in food products. These physical,
chemical, and biological approaches can improve its nutrient content,
flavor, and texture, reduce anti-nutritional factors, and, more impor-
tantly, enhance the production of bioactive phytochemicals and their
health benefits. Generally, the impacts resulting from physical methods
exhibited a dispersing effect, with heat treatment primarily employed

WU ET AL .
FIGURE 4 The fermentation benefits of pigeon pea and its application.
for anti-nutrient reduction and extrusion served for industrial produc-
tion. Chemical methods were less explored and were mainly focused
on modifying the structure of starch and protein. On the other hand,
biological methods emerged as a promising avenue, demonstrating
their potential to comprehensively and simultaneously address mul-
tiple aspects. These aspects included the reduction of anti-nutrients,
the augmentation of health-promoting nutrients, and the obvious
improvement of sensory properties. This notably helped to increase
the customer acceptance of pigeon pea. Given that, there are still some
research gaps to be fulfilled in this area: (1) More mechanistic stud-
ies are needed to achieve a systematic elucidation of these strategies.
(2) Extensive research has been conducted on the medicinal value of
pigeon pea leaves. However, limited data are available regarding the
potential health benefits of sprouted and fermented pigeon pea seeds.
(3) Currently, the sprouting and fermentation processes still need more
research and improvement to clarify the changes in nutrients and plant
secondary metabolites during processing. In addition, we propose that
the main nutritional and food applications for pigeon pea in the future
might be: (1) plant-based protein source with pleasant texture and fla-
vor; (2) functional foods developed with known functional benefits and
to specific person/patients.
26438429, 0, Downloaded from https://onlinelibrary.wiley.com/doi/10.1002/fft2.381 by Cochrane Philippines, Wiley Online Library on [19/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
13
AUTHOR CONTRIBUTIONS
Jun Wu: Conceptualization; data curation; writing—original draft;
visualization. Qian Zhou: Conceptualization; writing—original draft;
writing—review and editing; funding acquisition. Chenhaojin Zhou:
Writing-original draft. Ka-Wing Cheng: Writing—review and edit-
ing. Mingfu Wang: Conceptualization; writing—review and editing;
supervision; project administration; funding acquisition.
ACKNOWLEDGMENTS
This work was supported by the Guangdong Basic and Applied Basic
Research Foundation of China (Grant No. 2022A1515012098), the
National Natural Science Foundation of China (Grant No. 32101935),
and the Shenzhen Science and Technology Program (Grant No.
ZDSYS20220117155800001).
CONFLICT OF INTEREST STATEMENT
The authors declare that there are no conflicts of interest.
ORCID
Qian Zhou https://orcid.org/0000-0003-0138-6761
Mingfu Wang https://orcid.org/0000-0003-1469-3963

14
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How to cite this article: Wu, J., Zhou, Q., Zhou, C., Cheng,
K.-W., & Wang, M. (2024). Strategies to promote the dietary
use of pigeon pea (Cajanus cajan L.) for human nutrition and
health. Food Frontiers, 1–17. https://doi.org/10.1002/fft2.381