Food Chemistry 415 (2023) 135743

Contents lists available at ScienceDirect

Food Chemistry
journal homepage: www.elsevier.com/locate/foodchem

Review

Hard-to-cook phenomenon in common legumes: Chemistry, mechanisms

and utilisation
Dilini Perera a, Lavaraj Devkota a, Gil Garnier a, Joe Panozzo b, Sushil Dhital a, *
a
Department of Chemical and Biological Engineering, Monash University, Clayton Campus, VIC 3800, Australia
b
Agriculture Victoria Research, Horsham, Victoria 3400, Australia

A R T I C L E I N F O A B S T R A C T

Keywords: Future dietary protein demand will focus more on plant-based sources than animal-based products. In this
Common beans scenario, legumes and pulses (lentils, beans, chickpeas, etc.) can play a crucial role as they are one of the richest
Hard to cook sources of plant proteins with many health benefits. However, legume consumption is undermined due to the
Mitigation strategies
hard-to-cook (HTC) phenomenon, which refers to legumes that have high resistance to softening during cooking.
Phytochemicals
This review provides mechanistic insight into the development of the HTC phenomenon in legumes with a special
Pectin-cation-phytate
focus on common beans and their nutrition, health benefits, and hydration behaviour. Furthermore, detailed
elucidation of HTC mechanisms, mainly pectin-cation-phytate hypothesis and compositional changes of mac-
ronutrients like starch, protein, lipids and micronutrients like minerals, phytochemicals and cell wall poly-
saccharides during HTC development are critically reviewed based on the current research findings. Finally,
strategies to improve the hydration and cooking quality of beans are proposed, and a perspective is provided.

1. Introduction
The common beans (Phaseolus vulgaris L.) are legumes belonging to
the Fabaceae family and one of the most widely cultivated legumes in
tropical and temperate agricultural lands (Ramankutty et al., 2018).
Beans are a staple food for human diets, providing as much as 15% of
daily calories and 36% of complete daily protein, emphasizing their vital
role in providing energy and nutrients (Bessada et al., 2019). In addition,
beans are rich in dietary fibres, minerals (average 3 g/kg of Ca, 40 mg/
kg of Fe, and 35 mg/kg of Sn), and vitamins; they also contain phyto-
chemicals and polyunsaturated fatty acids such as linolenic acid, linoleic
acid and Oleic acid essential for human life (Grela et al., 2017). The
Lancet Commission on sustainable food systems and healthy diets have
addressed three critical factors in healthy, nutritious, and sustainable
food production that are the key pillars of the future food system (Willett
et al., 2019). The commission concluded that global food systems can
provide dietary benefits for healthy and environmentally sustainable
outcomes by 2050 and beyond. However, this goal can be achieved by
replacing unhealthy foods with healthy, sustainable foods.
Furthermore, sustainable food production encompasses the envi-
ronmental impact, including phosphorus and nitrogen cycles, climatic
changes, water, land, and biodiversity. Compared to other food pro-
ductions, legume cultivation has the lowest impact on the environment
in all aspects of greenhouse gas emission, land usage, energy con-
sumption, and possible acidification (Willett et al., 2019; Rahman et al.,
2022). Legumes also require less water and low fertile land (Vadez et al.,
2011) and can improve soil quality by atmospheric nitrogen fixation.
Therefore, incorporating legumes into the diet, minimising food loss and
waste, and enhancing food production techniques help to accomplish
the sustainable food system goals.
However, the potential consumption of legumes is undermined by
the requirement to overcome HTC defects manifested during storage
under different temperatures and relative humidity. HTC phenomenon
is a characteristic of stored legumes where the grain does not soften
during a considerable cooking time (Affrifah et al., 2021; Sofi et al.,
2022). Cooking for an extended period requires much energy and can
result in diminished nutritional quality and bioavailability. Therefore, it
is critical to understand the correlation between cooking time and
changes in the physicochemical properties of stored beans. During the
cooking, the structure and physicochemical functions of middle lamella
pectin and proteins are irreversibly altered, loosening the cell walls and
producing a soft texture. However, HTC beans do not soften after

* Corresponding author.
E-mail addresses: dilini.perera1@monash.edu (D. Perera), lavaraj.devkota1@monash.edu (L. Devkota), gil.garnier@monash.edu (G. Garnier), joe.Panozzo@
agriculture.vic.gov.au (J. Panozzo), sushil.dhital@monash.edu (S. Dhital).

https://doi.org/10.1016/j.foodchem.2023.135743
Received 13 September 2022; Received in revised form 15 February 2023; Accepted 16 February 2023
Available online 23 February 2023
0308-8146/© 2023 The Author(s). Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
D. Perera et al. Food Chemistry 415 (2023) 135743

reasonable cooking time since prolonged storage is due to a realignment
of cellular structure and secondary/higher level structure/interaction of
macro-molecules (Rousseau et al., 2020; Sofi et al., 2022). For example,
the cell walls may become more rigid, making it harder for water to
penetrate and soften the beans. This can be due to changes in the
structure of the cellulose and hemicellulose fibers that make up the cell
walls. HTC beans may also experience changes in their protein compo-
sition during storage, which can affect their ability to soften during
cooking. Some HTC beans have been found to have increased levels of a
protein called phaseolin, which can interfere with water absorption and
prevent the beans from softening (Rousseau et al., 2020).
Apart from prolonged storage, other factors such as climate, genetic
differences between species, processing behaviours, and specialised
treatments are known to cause the HTC defect (Yousif et al., 2007;
Mubaiwa et al., 2016). It can be hypothesised that these factors, indi-
vidually or holistically, can contribute to the HTC phenomenon.
Different legumes have been the subject of explanations for the pro-
cesses underlying the HTC phenomenon (Mubaiwa et al., 2016; Siddiq &
Uebersax, 2022; Wafula et al., 2022). The hydration of beans during
soaking and cooking is influenced by various intrinsic factors like size,
the thickness of the seed coat, the chemical composition of cotyledons,
size of the hilum and micropyle, as well as extrinsic factors like tem-
perature, pH level, and presence of solids in the soaking medium (Pen-
icela, 2010; Bassett et al., 2021). Nevertheless, there is no
comprehensive understanding of the phenomenon involved, and it is
still not well understood why some varieties have shorter cooking times
than others.
This review will critically examine recent developments in the HTC
phenomenon, focusing on common bean types, providing mechanistic
insight into the fundamental causes, and recommending approaches to
improve the quality of cooked beans. HTC beans are a significant
concern in food processing as they require longer cooking and can
produce harsh, unpalatable beans. Traditionally, thermal and non-
thermal treatment techniques have been used to shorten the cooking
time of beans. However, recent research has explored new approaches to
address this issue. One such approach involves the development of new
varieties of legumes with reduced levels of phytase enzymes, which are
known to contribute to HTC (Cominelli et al., 2020).
1.1. Bean nutrients and health-related benefits
The consumption of legumes has expanded globally due to their
positive health benefits. The Dietary Guidelines for Americans
(2020–2025) state that a healthy diet should contain half a cup or 130 g
of beans and other legumes (Thompson, 2021). Carbohydrates are the
primary component (50–60% of the dry mass) of beans and are found as
starch and non-starch oligo and polysaccharides (Martinez-Manrique
et al., 2011; Bento et al., 2022). Table 1 summarises the chemical
composition of raw common beans based on recent findings.
Clinical studies and meta-analyses have shown that consuming beans
and other whole legumes directly correlate to lowering metabolic dis-
ease, thus reducing mortality and morbidity (Li et al., 2021a; Binou
et al., 2022). Bean starch is entrapped inside a thick cellular matrix that
limits the diffusion of both enzymes and water. Thermal treatments
enhance the water absorption and swelling of starch granules, which
leads to the disrupt ordered arrangement of amylose and amylopectin
chains and the eventual breakdown of the granules. However, steric
hindrance of the semi-crystalline structure of starch and physical bar-
riers of intact cell walls leads to lower enzymic susceptivity in the small
intestine and a slower starch swelling and degradation rate. The slower
swelling rate allows the amylose and amylopectin chains to maintain
their ordered arrangement for longer. This contributes to a lower gly-
cemic index by slowing down the rate of starch digestion and absorption
in the small intestine and results in a slow and more gradual increment
in blood glucose levels, which helps prevent people with diabetes by
avoiding the detrimental effects of sudden spikes in blood glucose levels
(Dhital et al., 2016; Miao & Hamaker, 2021; Fang et al., 2022).
Many recent studies have shown that cellular structure remains
intact during cooking and processing (Dhital et al., 2016; Li et al.,
2021b; Xiong et al., 2022). The intact cellular structure slows down the
rate at which enzymes reach the substrate, reducing the rate at which
starch and proteins are hydrolysed. This slows down the glucose and
amino acid release rate, reducing the glycaemic and insulinemic re-
sponses to the food. The intact cells are thus passed to a large colon
where colonic microbiota ferment cells to produce a beneficial effect
(Huang et al., 2021; Xiong et al., 2022) to reduce the risk of breast
cancers, prostate cancers and colon cancers (Chen et al., 2020).
Beans contain a significant proportion of oligosaccharides and non-
starch polysaccharides (NSP). Cellulose, hemicellulose, and lignin are
examples of non-starch polysaccharides. The insoluble NSP fraction is
known to be slowly fermented in the colon, whereas the soluble NSP is
comparatively fast fermented (Comino et al., 2018; Gidley & Yakubov,
2019). Due to their small size, the oligosaccharides are rapidly fer-
mented (Li et al., 2021b). In addition, insoluble fibres are essential to
material movement in the digestive system (Tosh & Yada, 2010; Deb-
nath et al., 2019; Fang et al., 2022). Galactomannan gums, oligosac-
charides, and glucans are soluble fibres that help control blood sugar
and decrease cholesterol (Li, Chen, Bui, Xu, & Dhital, 2021; Rodriguez,
Jimenez, Fernandez-Bolanos, Guillen, & Heredia, 2006; Tosh & Yada,
2010). A significant amount of viscous soluble and insoluble dietary
fibre and resistant starch are major contributors to health benefits
(Xiong et al., 2022). Beans provide the “composite” fibres with a varied

Table 1
Chemical composition of raw common beans at (% dry basis).
Sample Ash Lipids Protein Total Total Insoluble Soluble Total Reference
description Carbohydrate starch dietary fiber dietary fiber dietary
fiber

Dark bean 3.8 ± 1.1 ± 17.7 ± 77.4 – 21.5 ± 3.4 5.8 ± 1.1 27.2 (Duenas et al., 2016)
(Tolosa) 0.1 0.1 0.5
Common beans 3.2–5.5 1.4–2.7 20.9–24.2 54.3–70.3 41.8–45.6 13.9–32.8 2.9–7.7 1.7–4.0 (Liu et al., 2013; Chavez-
Mendoza & Sanchez, 2017)
Black beans 4.0–4.6 1.5–1.7 20.3–26.8 65.4–73.4 – – – 2.3 (Telles et al., 2017; Chen
et al., 2021)
Carioca beans 3.9 0.7 22.5 53.2–72.9 – – – (Telles et al., 2017)
Red Kidney 3.3–4.2 1.9–2 25.3–27.9 50.4 – 29.9 8.3 38.2 (Kan et al., 2017)
beans
Haricot bean 2.8–4.3 0.7–2.8 17.9–22.1 56.5–61.6 – – – – (Kitum et al., 2020)
White kidney 3.5 1.4 25.6 59.6 – – – (Guzel & Sayar, 2012; Li
beans et al., 2014)
Pinto bean 4.9 1.8 30.1 57.3 – – – – (Sai-Ut et al., 2009; Audu &
Aremu, 2011)
Navy beans 4.14 2.6 18.1 54.3 – – – – (Deb-Choudhury et al., 2021)

2
D. Perera et al.
fermentation rate throughout the colon due to their diverse fibre profile.
This will allow fermentation and pH to drop throughout the colon,
which is considered a critical component in colonic health (Gidley,
2013).
Beans are rich in phytochemicals, minerals (iron, zinc, copper,
phosphorous, aluminium), and vitamins, such as thiamine, riboflavin,
niacin, pyridoxamine, biotin, folate and tocopherols (Campos-Vega
et al., 2010; Hayat et al., 2014). Phytochemicals like polyphenols can act
as functional and nutraceutical ingredients and possess antioxidant and
anticarcinogenic properties (Campos-Vega et al., 2010; Chavez-
Mendoza & Sanchez, 2017). A significant concentration of these
phenolic chemicals is found in the bean seed coat and fibres, enhancing
bioavailability (Nicolas-Garcia et al., 2021).
Common beans include cholesterol-free protein, vitamin B com-
plexes, vitamin E, minerals, fibre, resistant starch, and phytonutrients,
making them suitable for people of all ages. Their nutrient composition
corresponds to the nutritional requirements of growing children, teens,
and adults. However, the HTC issue may compromise the quality of
these vital nutrients and impact their bioavailability, which demands
further investigation.
2. Structural organisation of beans
Legume seeds have two main parts, the seed coat and the cotyledon.
Within the seed coat, a small opening called the micropyle can be found,
while on the surface of the seed, a scar called the hilum remains after it is
separated from the pod. The microstructural changes in beans can be
easily observed using transmission (TEM) and scanning electron mi-
croscopy (SEM). There are several techniques used for sample prepa-
ration for SEM and TEM. In general, samples are first fixed in a
glutaraldehyde or formaldehyde solution and then freeze-dried to
remove the water. Then dried samples are coated with a thin layer of
platinum or gold to make them conductive and to prevent charging
during the SEM imaging process (Pathan et al., 2010; Miranda et al.,
2019). Embedding is commonly used to prepare bean samples for TEM.
Once samples are fixed in a solution as explained, and then embedded in
a resin. The resin is then cut into thin sections, stained and examined
using TEM (Ayache et al., 2007).
2.1. Structure of the seed coat
The seed coat is a cotyledon protective covering that shields the seed
from the outside environment (Penicela, 2010). SEM analysis can
identify reproducible patterns or fingerprints in each seed coat (Hughes
& Swanson, 1985; Zablatzka et al., 2021). In Fig. 1A, the left column
illustrates fresh beans’ microstructure, whereas the right column in-
dicates the HTC beans. As shown in Fig. 1 A (a), the cross-section of a
legume seed coat includes three distinct layers of a palisade, subepi-
dermal, and parenchyma cells. The outermost seed coat layer is made up
of column-shaped palisade cells. The short, columnar subepidermal
cells, known as hourglass cells, are found directly under the palisade
layer (Sofi et al., 2022). Both cell types are densely packed and grow in
length as the seed grows; nevertheless, no substantial structural changes
occur throughout maturity (Hughes & Swanson, 1985; Zablatzka et al.,
2021).
In contrast, the third layer of parenchyma cells comprises 10–15
loosely bound layers, a cell with intercellular spaces (Sofi et al., 2022).
The micropyle, hilum and seed coat individually contribute to the water
absorption, and the thickness and length of the palisade layer, the
micropyle width, and the cell layer thickness attached to the micropyle
play essential roles in bean hydration (Mandizvo and Odindo, 2019).
Further, Fig. 1A (b) illustrates that HTC legumes have a more rigid and
more organised seed coat structure than fresh beans (Sharafbafi, 2008),
which prevents water absorption through the seed coat during cooking
(Nakalema, 2015; Sofi et al., 2022).
3
Food Chemistry 415 (2023) 135743
2.2. Cotyledon structure
Cotyledon comprises the majority of the mass and size of legume
seeds and has a highly organised cell structure. It contains large granules
of starch in size 10–50 µm and small spherical protein bodies surrounded
by lipoprotein membranes and consists of phytin-rich crystalline glo-
boids (Nakalema, 2015). All of those are embedded in a protein matrix,
as illustrated in Fig. 1A (c). The xylem and phloem of the plant rapidly
transport water, ions, and nutrients to the cotyledon storage cells.
Generally, the cotyledon cell wall contains more pectic substances
than the seed coat cell wall (Fig. 1B). The middle lamella contains
pectin, the primary component in holding the cotyledon cells together
and providing structural support to the tissues (Pirhayati et al., 2011;
Sofi et al., 2022). A clear cotyledon cell separation can be observed in
fresh beans, whereas cells are not separated in HTC beans (Fig. 1A (c)
and (d)). Tricellular junctions with intercellular spaces and the middle
lamella can be visible predominantly in HTC beans due to middle
lamella pectin, which binds adjacent cells together (Fig. 1A (d), (f), and
(h)), and lignified cell walls are highlighted compared to the fresh beans
(Gwala et al., 2019). HTC beans take more time to hydrate since the cells
in the hilum area get compacted and reduce cell separation. Thus, this
prevents water diffusion through the hilum and delays the cotyledon
hydration.
The 30–36 long chains of 1,4-linked glucose molecules make up
cellulose fibrils and are established by hydrogen bonds with one another
to create an impermeable and insoluble crystalline structure around the
cotyledon cells (Zhao et al., 2019). As illustrated in Fig. 1B, hemicellu-
lose, including xylan, glucuronoxylan, arabinoxylan, glucomannan, and
xyloglucan, are connected to that cellulose via hydrogen bonds and by
embedding in the pectin matrix (Albersheim et al., 2010). Rhamnoga-
lacturonan I (RGI) is the most prevalent pectin, which is formed by
modifying rhamnose and galacturonic acid, as well as the side chains of
galactans and arabinans (Doblin et al., 2010). The changes in structure
and composition of the cotyledon cell wall can be linked to the HTC
phenomenon in legumes. (Shiga and Lajolo, 2006; Parmar et al., 2017a;
Uebersax et al., 2022).
Cotyledon cell wall polymers solubilise and depolymerise to soften
the tissues during cooking. Therefore alterations in those polymers
during aging may be responsible for the HTC defect. Cell walls are
complex structures of polysaccharides, proteins and phenolic compo-
nents held together by ionic and covalent bonds (Wainaina et al., 2021;
Sofi et al., 2022). They are primarily composed of two cellulose and
hemicellulose networks with separate pectin networks. The pectic
network is primarily composed of acidic rhamnogalacturonan and
homogalacturonan, as well as neutral arabinan, galactan, and arabino-
galactan polymers (Sofi et al.,2022). Depolymerisation and solubilisa-
tion of pectin soften tissues and increase cell separation during cooking
(Rousseau et al., 2020). With the storage, cell separation becomes more
difficult due to the formation of secondary interactions with other
molecules and cell wall polymers. For example, the demethoxylation of
pectin and the formation of calcium pectate and magnesium pectate may
lower cell separation during cooking (Sofi et al., 2022).
Kyomugasho and colleagues (2021) have used state diagrams to
explore the relationship between glass transition temperature (Tg) and
the high-temperature stability of beans, explicitly examining the role of
the cotyledon. They found that the Tg of starch had the maximum value
at a specific moisture content, followed by protein isolate and cell wall
components. The findings indicate that cell wall components are most
unstable at high storage temperatures. Thus, it is likely that the forma-
tion of high-temperature stability in beans is strongly related to the
formation of HTC through cell wall materials.
Based on the pectin-cation phytate hypothesis (section 4), phytate is
in globoids inside the protein matrix. Tg of proteins is greater than that
of cell wall fraction, and high temperatures are required to release cal-
cium ions from the hydrolysis of phytate (Chigwedere et al., 2019a).
Recent research on HTC has found no significant impact on the degree of
D. Perera et al. Food Chemistry 415 (2023) 135743

Fig. 1. [A] Schematic diagram of changes in the structural organisation of fresh and HTC beans; [B] Schematic diagram of bean cell wall structure with cell wall
composition (Shiga & Lajolo, 2006; Albersheim et al., 2010; Zhao et al., 2019).

4
D. Perera et al.
methyl esterification (DM) of pectin with storage (Chigwedere et al.,
2019c; Toili et al., 2022). As a result, the Tg of a protein can be asso-
ciated with HTC, which is connected to phytate hydrolysis, as long as
Ca2+ ions are released and can cross-link with cell wall pectin.
Apart from that “lignification-like hypothesis” suggests the deposi-
tion of phenols and lignin interpolymer cross-linking in the cell wall. In
this case, the Tg of the cell wall may be critical in determining the rate of
HTC development during storage. The phenolic compounds are pri-
marily concentrated in the seed coat and could migrate to the cell walls
during storage. Further, the mechanisms and theories of HTC are arti-
culated in-depth in section 4.
3. Hydration of beans
Hydration of dry beans is a crucial initial step in legume processing
that significantly impacts the final product’s quality and influences
subsequent processing success. Hydration occurs during soaking and
cooking, promoting bean softening by activating cell wall enzymes and
lowering the polymerisation of rhamnogalacturonan I (Martinez-Man-
rique et al., 2011; Holland et al., 2020). Moreover, it improves the sol-
ubility of polygalacturonan and galactan, as well as other
polysaccharides, and ultimately saves cooking time (Miano & Augusto,
2018b). Furthermore, hydration induces starch gelatinisation and pro-
tein denaturation homogeneously; and contributes to a uniform texture
throughout the whole grain (Njoroge et al., 2015). Complete hydration
of the beans enhances the heat transfer and deactivates anti-nutritional
compounds such as saponins, lectins, alkaloids and phytates, as well as
indigestible oligosaccharides and protease inhibitors (Miano & Augusto,
2018b).
“Hard to Cook” and “Hard shell” are two prominent textural defects
caused by prolonged bean storage under adverse conditions. HTC beans
do not soften due to a lack of hydration of cotyledon that reduces the cell
separation during cooking, whereas hard shell beans do not absorb
water through seed coat during soaking (Siddiq & Uebersax, 2022; Sofi
et al., 2022). However, proper hydration is critical in bean processing,
such as canning.
Grains and legumes have two hydration patterns: downward concave
shape (DCS) and sigmoidal, reflective of their water uptake pathway.
The bran, the outer layer of cereal grains including wheat, rice, barley,
millet, corn and oat, is more permeable to water, allowing water to
diffuse across the surface area (Miano & Augusto, 2018a). However,
starch is the main reserve component in grains and is usually more
compact in the endosperm; thus, it takes excessive time to complete
hydration (Miano et al., 2018b). This hydration kinetics behaviour is
known as downward concave shape (DCS), which includes an initial
exponential (II) and later equilibrium stage (III), as explained in Fig. 2.
Legumes, such as common beans, possess an impervious seed coat,
Fig. 2. Typical cereal grains and beans hydration behaviour and pathways; I:
lag phase, II: exponential phase, III: equilibrium phase (Figure concept taken
from Miano and Augusto (2018a)).
5
Food Chemistry 415 (2023) 135743
and the bean hydration behaviour is somewhat different from the grain
hydration because water permeability can vary according to the com-
plex structure, different tissues and cell sizes, structure, composition,
and zones of beans (Miano et al., 2015). Therefore, the hydration pro-
cess of beans has a sigmoidal shape which involves an initial lag stage
(I), then an exponential stage (II), and finally an equilibrium stage (III),
as in Fig. 2. Here, water must diffuse through the hilum via the hilar
groove and tissues before reaching the cotyledon since the bean seed
coat is impermeable to water (Miano & Augusto, 2018a). Water is
subsequently diffused into the gap between the cotyledon and the seed
coat, increasing the seed coat’s permeability. Then the water begins to
diffuse via the seed coat. In the last step of water uptake, the cotyledon is
hydrated via diffusion and capillary flow and achieves the equilibrium
moisture level (Miano & Augusto, 2015). Hence, the impermeability of
seed coats in beans causes a lag phase (stage I). Nevertheless, if the seed
coat of the legume seed is porous, water can pass to the exponential
stage (II) by avoiding the lag phase (I), shown in Fig. 2.
4. Hard-to-cook phenomenon
Beans are grown in tropical or temperate climates and, once har-
vested, stored under controlled conditions until consumption. Prolonged
storage under hot (>25 ◦ C) and high relative humidity (>65%) condi-
tions led to the developing HTC beans, characterised by the prolonged
cooking time to soften the cotyledon. The changes in HTC beans are
irreversible and mainly associated with structural and compositional
changes within the cotyledons. This phenomenon can be explained by
several mechanisms, such as the formation of insoluble pectates,
oxidation of lipids, hydrolysis of phytates, tannin and polyphenol
migration (Njoroge et al., 2016; Chigwedere et al., 2019c). Recent
studies on the HTC phenomenon are summarised in Table 2, with
possible mechanisms reviewed in detail.
4.1. Pectin cation phytate hypothesis
This hypothesis is known as the duel enzyme mechanism, which is
described by combining two enzymes: phytase and pectin methyl
esterase (PME). Generally, plant cells contain phytic acid (PA), a potent
chelating agent of calcium and magnesium ions, as illustrated in Fig. 3A
(a). During storage, the phytase enzyme hydrolyses PA to liberate
phosphate and magnesium into the cotyledon cells (Fig. 3A (b)). As a
result, PA’s chelation potential decreases, and calcium and magnesium
ions increase cross-links with middle lamella pectin (Wainaina et al.,
2022c). Meantime, middle lamella pectins are hydrolysed to pectinic
acid and methanol by PME, forming calcium pectate and magnesium
pectate that make beans hard (Siddiq & Uebersax, 2022). The resultant
calcium and magnesium pectates are not readily soluble in boiling,
which reduces cellular separation and consequently produces HTC
beans. Mattson et al. (1950) have experimentally proven that freshly
harvested peas contained more PA than HTC peas. The same results were
reported by Kruger et al. (2015) for cowpea samples and Panayotou
et al. (2008) for common beans and lentil samples. The PA reduction is
most likely caused by the inherent phytase activity of beans and is
compatible with the phytate-phytase, and insoluble pectin concept,
which claims that PA is dephosphorylated during storage leads to HTC
development (Kruger et al., 2015; Wainaina et al., 2022c).
Recent research has found that the activity of PME is not affected by
the HTC phenomenon (Njoroge et al., 2015; Chigwedere et al., 2019c).
They found that the degree of methyl esterification (DM) did not vary
considerably during bean storage, implying that DM alterations do not
play a significant role in HTC formation. Due to the activity of PME, DM
of pectin should decrease during storage. Furthermore, they discovered
that throughout ageing, the water extractable pectin (WEP) fraction
declined while the sodium carbonate extractable pectin (NEP) fraction
increased, implying that loosely bound WEP was exchanged for cova-
lently bonded NSP. In contrast, the chelator extractable pectin (CEP)
D. Perera et al. Food Chemistry 415 (2023) 135743

Table 2
Recent studies on HTC phenomenon in legumes with main findings and conclusions.
Bean type Experiment Conclusion Reference

Carioca beans Evaluated thermal properties and crystallinity of bean starch Beans stored under a nitrogen-modified atmosphere exhibited (Rupollo et al.,
stored for 360 days. reduced crystallinity and gelatinization enthalpy. Therefore, using 2011)
a nitrogen-modified atmosphere can help preserve the thermal
properties of starch.
Cowpea Cowpeas were stored for 21 days (42 ◦ C and 67% RH). Proton-induced X-ray emission (PIXE) analysis of cowpea found (Kruger et al.,
that Ca and Mg bind to wall-middle lamella pectin. 2015)
Common beans Investigated the changes in non-starch polysaccharides in The study revealed a significant difference in the non-starch (Yi et al., 2016)
beans stored for eight months (35 ◦ C and 75% RH). polysaccharides of the seed coats and cotyledons of HTC beans,
and it was found that the interaction between pectin
polysaccharides and divalent ions plays a critical role in the
development of HTC.
Kidney bean Studied the changes in protein and starch microstructure. HTC beans had higher levels of 31, 30, and 27 kDa polypeptides (Parmar et al.,
and a more significant amount of β-sheets and α-helix in their 2017b)
secondary structure. It also contained more damaged starch
granules, contributing to their high peak viscosity.
Carioca bean Beans were stored at 40 ◦C and humidity at 75%. Different genotypes show varying degrees of HTC intensity. More (Siqueira et al.,
susceptible genotypes had high water absorption and peroxidase 2018)
activity, and oxidoreductases remained active during storage, with
an increase in total phenol content.
Common beans Seed coat and cotyledon powders were incubated to obtain a Tg-moisture relation was confirmed that development HTC beans (Chigwedere et al.,
water activity range of 0.1–0.9, and Tg was determined. stored above Tg. 2019a)
Carioca beans Beans were stored for 240 days at different combinations of Beans stored at 36 ◦ C showed signs of HTC defect after 80 days, (Demito et al.,
temperature (12, 20, 28, and 36 ◦ C) and moisture content and the defect increased after 240 days. Refrigerated storage at 2019)
(16.7% and 13.8%). 12 ◦ C provided the best preservation of beans during storage, with
the highest digestibility indexes obtained under these conditions.
Canadian wonder Beans were stored at 35 ◦ C and 80% RH for 20 weeks, and the Postharvest storage reduced the bioaccessibility of Ca, Fe, and Zn, (Rousseau et al.,
beans bioaccessibility of Mg, Ca, Fe, and Zn of cooked beans was which was linked to the increased chelation of minerals by cell 2020)
studied. wall polymers.
Cowpea HTC in cowpea was investigated using two steaming Steam treatment for 4–6 min effectively prevents HTC in cowpeas. (Affrifah et al.,
temperatures (100 ◦C, 121 ◦C) and three steaming times (2, 4, 2021)
and 6 min).
Red haricot beans Beans were stored for three months The aged beans showed a more rigid cell wall with strong cross- (Chen et al., 2021)
(35 ◦C and 83% RH). links of Ca2+_ pectin and feruloylated pectin. This suggests that
aging causes changes in the structure and composition of the cell
wall, leading to increased rigidity.
Carioca beans Beans were stored for three and six months (27.5 ◦C and 60% Kaempferol levels may distinguish with darkening properties, but (Bento et al., 2021)
RH), and the changes in phenolics and saponins were these levels decrease during storage. Additionally, the higher
observed. concentration of saponins could be associated with the darkening
of carioca beans.
Red haricot, rose coco, The storage stability of beans was examined for varying Beans stored at temperatures above their Tg resulted in the (Kyomugasho
red kidney, and durations (2, 4, 6, 8, and 12 months) (20 ◦C, 25 ◦C at 75% RH; development of the HTC defect. The Tg values were observed to et al., 2021)
pinto bean 35 ◦C at 75 %RH; 45 ◦C at 75% RH; and 35 ◦C at 83% RH). vary in the following order: Tg of starch > Tg of protein > Tg of the
cell wall. Below, the Tg beans are in a glassy state characterised by
high viscosity and immobilisation of cellular components, and
above, the Tg will be promoted molecular mobility, and HTC
defect was observed.
Common beans Beans were stored for three months (35 ◦C and 83% RH). The NIR spectra can be used to predict the cooking time of fresh beans (Wafula et al.,
susceptibility to developing the HTC defect was predicted and their vulnerability to the HTC defect when exposed to 2021)
using NIR spectra. unfavourable storage conditions.
Red kidney beans Beans were stored at 25 ◦C, 30 ◦C, 35 ◦C and 42 ◦C for six The volatile marker compounds found are connected mainly to the (Wainaina et al.,
months and studied the release of volatiles. breakdown of proteins and the oxidation of lipids. 2022a)
Red kidney beans Beans were stored for up to 6 months, at 25 ◦C, 30 ◦C, 35 ◦C Storing the beans at or below their Tg can help maintain their (Wainaina et al.,
and 42 ◦C, at five moisture contents (6.9%, 8.7%, 11.1%, textural stability and cooking properties even after prolonged 2022b)
12.8%, 14.5%) and textural stability and cooking behaviour storage.
were studied.
Rose coco, Nabe, Beans were stored at 35 ◦C and 83% RH for three months to The aging process resulted in a significant reduction in the InsP6 (Wainaina et al.,
Carioca, Red haricot induce the development of HTC. The levels of pectin, cell level, and phytate hydrolysis was observed to be the predominant 2022c)
wall-bound calcium, and inositol hexaphosphate (InsP6) were factor, whereas pectin demethylesterification appeared to have
measured before and after aging. little or no impact.
Red kidney beans Beans were stored at 25, 30, 35, and 42 ◦C and studied the Phytate hydrolysis resulted in high InsP5 content. Temperature (Wainaina et al.,
phytate hydrolysis. played a more significant role than moisture content in this 2022d)
process.
Red haricot beans Beans were stored at 35 ◦C and 83 % RH for three months, and Phytate hydrolysis happens more frequently during storage than (Chen et al.,
determine the phytates (inositol phosphates), Ca and Mg were during bean soaking. Aged beans with firmer textures after 2022a)
during post-harvest storage and subsequent soaking of the cooking may have more Ca-pectin cross-linking.
beans.
Red kidney beans Beans were stored at 35 ◦C and different RH (6, 54, 66, 72, Beans can be stored at low temperatures or dried to reduce (Aravindakshan
and 82%) for 15 weeks. moisture content below their Tg values. et al., 2022)
Andean commercial Studied the PME enzyme contribution to HTC defect using PME-encoding genes were strongly expressed in the easy-to-cook (Toili et al., 2022)
bean molecular tools. bean, whereas their inhibitors were more abundant in the HTC
bean.
Common beans Beans were stored under RH at 70% at a temperature of 30 ◦C Changes in the 37 phenolic chemicals were discovered during (Nicolas-Garcia
Jamapa variety and studied phenolic compounds and their relationship. storage of 270 days. Condensed tannins were observed in the seed et al., 2022)
coat after 90 days.
(continued on next page)

6
D. Perera et al.
Table 2 (continued )
Bean type Experiment
Red haricot beans Beans were stored under 35 ◦C and 83% RH conditions for
three months, and their cell wall composition and phenolics
were analysed.
fraction increased nonsignificantly (Chigwedere et al., 2019c). Based on
this knowledge, phytate breakdown and subsequent cation migration to
the cotyledon is a more plausible explanation of the pectin-cation-
phytate hypothesis than the previously held concept of PME activity.
Chen et al. (2021) studied the effect of humidity and temperature on the
microstructural changes of pectin in stored red haricot beans. According
to their observations, the cell walls of HTC beans are bound with greater
cross-links with calcium and magnesium pectates than fresh beans.
Several recent studies have linked changes in inositol phosphate con-
version and subsequent migration of Mg and Ca cations to be the most
prominent explanation of the phytate-cation hypothesis (Chen et al.,
2022a; Wainaina et al., 2022c).
4.2. Lipid oxidation and polymerisation
Beans contain 1–2% lipids present predominantly as phospholipids
and triacylglycerols components, whereas diacylglycerols, hydrocar-
bons, sterol esters, and hydrocarbons are available in minor amounts
(Hayat et al., 2014; Rodriguez et al., 2022). Beans contain essential
polyunsaturated fatty acids, and linolenic acid is the most prominent
(Yoshida et al., 2008; Hayat et al., 2014).
Chemistry of those fatty acids changes significantly during storage
conditions (Nasar-Abbas et al., 2008). Storing the legumes at 4 ◦ C
minimises lipid degradation, even when ground or damaged. The high
temperatures and humidity induce lipid oxidation. Lipid degradation
occurs in beans storage at 30 ◦ C and humidity at 83%, making the beans
difficult to cook (Reyes-Moreno et al., 1993; Ferreira et al., 2017). Also,
oxidation and polymerisation of membrane lipids are linked to alter-
ations in moisture absorption, which may impact texture (Chigwedere
et al., 2018). Cell membranes are comprised of a continuous phospho-
lipid bilayer which serves as a fundamental structure and barrier to
material movements across the membrane, and proteins embedded in
continuous bilayers (Liu and Bourne, 1995; Dhital et al., 2016). In
contrast, membrane proteins act as specialised receptors, transporters
and enzymes. Since membrane lipid bilayers are impervious to most
polar molecules, their movement is assisted by unique carrier proteins,
each of which is accountable for transporting a specific solute across the
bilayer (Liu and Bourne, 1995).
Fig. 3B illustrates lipid oxidation and polymerisation related to the
development of HTC beans. The fatty acid (FA) composition of the HTC
bean membrane changes to an increased content of saturated fatty acids
(SFA) and a decline in polyunsaturated fatty acids (PUFA). C14:0 FA
primarily increased, whereas C18:3 was found to decrease most signif-
icantly (Reyes-Moreno et al., 1993). As shown in Fig. 3B, PUFA oxidised
to form free radicals. The membrane integrity is lost due to increased
peroxidation within the cytoplasm (Mubaiwa et al., 2016). Lipid
oxidation, polymerisation, and membrane degradation have been
recorded and found to be directly associated with HTC beans (Chigwe-
dere et al., 2019b; Chen et al., 2021). The hypothesis suggests that a
damaged cell membrane allows the free movement of cytoplasmic sol-
utes and increases the mobility of ions. However, noticeable leakage of
solutes occurred in stored beans, signifying irreversible degradation of
the cell membrane. Biochemically, cell membrane consistency is
required for proper cellular metabolism (Liu & Bourne, 1995). Conse-
quently, membrane degradation upon storage reduces the quality of
beans and is linked with the HTC legumes.
Following the pectin-cation-phytate hypothesis, membrane
7
Food Chemistry 415 (2023) 135743
Conclusion Reference
An increase in interactions between cell wall components was (Chen et al.,
noted, specifically through arabinose and galactose side-chains of 2022b)
RG-I pectin and high phenolic interactions with cellulose and
hemicellulose were observed.
breakdown in stored beans not only permits the movement of calcium
and magnesium cations from hydrolysed phytic acid to cell wall pectin
but also reduces turgor pressure, the force required to keep cells sepa-
rated from each other (Richardson & Stanley, 1991; Liu & Bourne,
1995). Structural changes of the plasma membrane and alterations in
SFA composition in membrane lipids suggest the possibility that mem-
brane degradation is the key factor contributing to the initiation of HTC
beans.
4.3. Lignification
Lignin is a complex and high-molecular-weight natural phenolic
polymer and a key component of cell walls. Lignin is biosynthesised
initially in the cell corners of the primary cell wall, followed by the
middle lamella and secondary wall (Zhong et., 2019). Cell wall lignifi-
cation obliterates the differences between primary cell walls and the
intermediate lamella of neighbouring cells (Mubaiwa et al., 2019;
Siqueira et al., 2018). It is reasonable to hypothesise that the HTC
development in common beans is caused, at least in part, by a lignifi-
cation process.
Lignin content in bean cell walls can change under high temperatures
and humidity during storage (Shiga, 2004; Nasar-Abbas et al., 2008).
Legume seed coat contains phenolic compounds, and those can convert
into lignin, leading to seed coat impermeability during seed develop-
ment (Sofi et al., 2022). The lignin deposition on cotyledon cells con-
tributes to the hardening of the cell wall, which decreases water
absorption and swelling, thus making cooking more challenging (Nasar-
Abbas et al., 2008). During storage, lignification of the middle lamella of
legumes can occur, further reducing cookability. Studies have reported
that the acid-detergent fibre and lignin content of faba beans increased
dramatically after 12 months of storage at temperatures higher than
37 ◦ C (Nasar-Abbas et al., 2008).
Peroxidase enzymes in cotyledon cells facilitate the oxidation and
polymerisation of phenolic substances to produce lignin monomers
(Grabber, 2005). Cross-linking of the cell wall may occur during initial
lignification due to the synthesis of lignin-protein complexes (Zhong
et al., 2019). The bean tissues contain phenolic precursors, such as free
phenolic compounds and seed coat polyphenols. Higher moisture level
supports the bioactivity of enzymes and lignifies the middle lamella and
cotyledon cell walls (Grabber, 2005).
According to Pirhayati et al. (2011), the lignin found in cotyledon
cell walls may impact the cooking characteristics of legumes. Although
cotyledon contains less lignin than the seed coat, this little lignin may
also contribute to the HTC phenomenon. The seed coat contains lignin
and its phenolic precursors, and polymerising these phenolic monomers
may lead to HTC beans (Reyes-Moreno et al., 1993). Rodriguez and
Mendoza (1990) reported that HTC mung bean contained 12% more
fibre, 23% more silica and seven times more lignin than fresh beans.
Therefore, a high amount of lignin, silica and cellulose content affects
the HTC phenomenon of legumes and water permeability through the
seed coat. The HTC legumes can be explained using a lignification-like
theory that involves polymerising phenolic acids.
4.4. Deposition of phenolic compounds and tannins
Changes in phenolic compounds and their derivatives also associate
with the HTC phenomenon of common beans. According to Parmar et al.
D. Perera et al.
Fig. 3. [A] Mechanism of formation insoluble pectate (a) Phytic acid (PA) activity of fresh cotyledon cells. (b1) phytase enzyme hydrolyses PA into inorganic
phosphate and magnesium inside the cotyledon cells. (b2) Pectin methyl esterase (PME) hydrolyses pectin in the middle lamella to pectinic acid and methanol. (b3)
Formation of insoluble pectates due to the activity of phytase and PME. [B] Phospholipid bilayer of cell membrane with the mechanism of membrane poly un-
saturated fatty acid oxidation to form free radicals and loss of membrane integrity due to increased peroxidation in the cytoplasm.
(2017), there is variation in the total phenol content of kidney beans in
both easy-to-cook (ETC) and hard-to-cook (HTC) grains. The researchers
found that the phenolic content of the seed coat changed during HTC
development. Changes in cell wall components lower the release of
phenolic compounds during soaking in HTC beans compared to ETC
beans (Nasar-Abbas et al., 2008; Pirhayati et al., 2011). They proposed
that the oxidative degradation of phenolic compounds was a primary
source of these component reductions (Nasar-Abbas et al., 2008).
Further, this may be due to the extra cross-linking of phenols with
proteins and carbohydrates during adverse conditions to inhibit water
absorption and reduce cell separation.
Under high humidity and temperature conditions, stored beans can
produce more free phenolic acids, like ferulic acid, which can bind with
proteins in the middle lamella of the cotyledon (Machado et al., 2008).
This interaction can increase protein hydrophobicity, which reduces
seed hydration during cooking and may limit cell separation. In addi-
tion, phenolic components can bind with insoluble polymers like lignin
and migrate to the cotyledon.
Tannins, a group of high-molecular-weight compounds that contain
multiple phenolic hydroxyl groups, can react with proteins to form
persistent cross-links through peroxidase and free radicals produced by
the breakdown of cell membranes. This reaction decreases the extract-
ability of proteins (Kruger et al., 2015). Further, condensed tannins in
common beans decrease with storage time, and a strongly negative
significant relationship between tannin concentration and HTC devel-
opment was discovered (Parmar et al., 2017a). Condensed tannins link
with macromolecules to change their function and cellular structure
(Giuberti et al., 2020). According to Srisuma et al. (1989), HTC beans
have high amounts of hydroxycinnamic acids, specifically ferulic acid
and hydroxycinnamic acids, leading to discolouration and increasing the
development of HTC (Mubaiwa et al., 2016; Siddiq and Uebersax, 2022).
Recently Chen et al. (2021) immunolabelled different pectin ex-
tractions and observed highly cross-linked pectin with calcium ions and
ferulic acid, and these complexes are concentrated in the intercellular
8
Food Chemistry 415 (2023) 135743
spaces of the cotyledon cells. The results support the pectin-phytate
hypothesis and the involvement of phenolic compounds in the cross-
linking of pectin during the development of the HTC phenomenon in
stored common beans.
5. Structural and biochemical changes during storage
Beans and other legumes undergo physical and metabolic changes
that impair their cooking properties when stored for prolonged periods.
Storage in adverse conditions develops the HTC phenomenon charac-
terised by incomplete hydration of the cotyledon during cooking,
causing it to remain rigid.
5.1. Starch
Structural and biochemical changes in the starch of HTC beans can
be elucidated in various length scales. At the supermolecular level,
thermal stability, short-range order, and crystalline structure are ana-
lysed by differential scanning calorimetry (DSC), Fourier transforms
infrared spectroscopy (FTIR), and X-ray diffraction (XRD), respectively.
Microscopic techniques observe the granular or tissue level changes as in
Fig. 1. For example, microscopic images (Fig. 4A) illustrate that cooked
HTC beans with lower cell separation in cotyledon and starch granules
are quite noticeable, whereas they appear fused in fresh beans (Miranda
et al., 2019).
Starch originates as semi-crystalline granular grouped in a radial
pattern and composed of amylose and amylopectin molecules (Yousif
et al., 2007). Amylose is assumed responsible for the amorphous
component, whereas amylopectin is considered for granular crystallinity
(Junejo et al., 2022). The legume starch is classified as type C, combi-
nation cereal starches (type A) and tuber starches (type B) (Junejo et al.,
2022). Loss of crystallinity results in starch solubilisation, which in-
creases the starch suspension viscosity and the pasting properties.
Generally, crystalline areas hold starch granules together, preventing
D. Perera et al.
swelling and individual molecules’ dispersion.
When starch is subjected to heat under excess water, the amorphous
regions have extensive hydration and swelling to expand the matrix held
together with unbroken crystals. This is referred to as gelatinisation
(Mohamed, 2021). The gelatinisation of legume starch occurs typically
within a limited range of temperatures at 76 ◦ C and results in endo-
thermic transformation. This enthalpy gain shifts the crystalline order to
a disordered state, causing the crystal stabilising forces to be disrupted
(hydrogen bonding). The crystalline starch structure changes during
storage, increasing the gelatinisation temperature. The HTC develop-
ment influences the quantity of water absorbed by common beans dur-
ing cooking (Yousif et al., 2007). However, complete starch
gelatinisation is achieved in fresh and HTC beans after 30 min, while the
aged beans required a far longer time to get softened due to the delay of
pectin solubilisation during cooking (Chigwedere et al., 2018).
The effect of storage conditions on starch gelatinisation and protein
denaturation can be analysed using DSC. Yousif et al. (2003) found that
storage at 30 ◦ C increased the peak (Tp) and onset (To) temperatures of
starch gelatinisation for adzuki beans. The increased gelatinisation
temperatures can be attributed to alterations in the helical structure
leading to a difference in the thermal stability of the starch (Hohlberg &
Stanley, 1987; Yousif et al., 2003). On the other hand, HTC bean starch
can be damaged by activating the amylase enzyme, and damaged starch
absorbs more water (Jukic et al., 2019). Bambara groundnuts exhibiting
the HTC phenomenon showed increased cell breakage and the release of
cytoplasmic contents readily accessible to amylases during digestion
(Gwala et al., 2019; Duijsens et al., 2021). As a result, the starch content
in HTC bambara groundnuts was digested faster than in fresh seeds
under the same cooking conditions (Gwala et al., 2019).
FTIR also reveal the possible differences between starch structures
during storage. Bands at 1022 and 1047 cm−1 describe amorphous and
crystalline indices of starch, respectively. The ratio of 1047/1022 can be
used to identify the relative crystallinity of starch (Sevenou et al., 2002).
Furthermore, the ratio decreased slightly during high-temperature
storage, suggesting that starch can lose its short-range ordered struc-
ture during storage. These findings suggest that storage conditions
significantly impact starch structure and organization.
Fig. 4. [A] Top:SEM images of navy beans stored at 4 ◦ C for 24 months. Bottom: Stored at 37 ◦ C for 24 months, showing “hard-to-cook” (Uebersax & Bedford, 1980;
Uebersax et al., 2022); [B] Changes in protein structures during storage to make protein network around the starch granules.
9
Food Chemistry 415 (2023) 135743
5.2. Protein
Benas are rich in globulins-type storage proteins, and tissue pH can
be used to indicate storage-caused HTC defects. Fig. 4B illustrates how
storage proteins are denatured and coagulated due to the sudden pH
drop within the cells (acidification).
Heat induces the changes in protein structures that lead to unfolding,
aggregation, coagulation, or gelation. Before starch gelatinisation,
stored bean proteins coagulate and create a network over the starch
granules, thus limiting water absorption for starch gelatinisation (Wang
et al., 2020). Structural changes in proteins can be further explained by
molecular reactions that may occur during storage. The absorption
peaks of FTIR spectra at wavelengths 1400 and 1800 cm−1 are charac-
teristic of amide bonds. The structural changes in proteins can be
identified by observing the bands between 3500 and 3300 cm−1 and
1649 and 1550 cm−1, which correspond to the deformation of primary
amide NH groups. Strong vibrations at 1157 and 1080 cm−1 are also
associated with these changes in protein structure. (Kaptso et al., 2015;
Warren et al., 2016).
The activity of proteinase enzymes can be increased, and non-
enzymatic hydrolysis of proteins can be induced at high humidity and
temperature (Coelho et al., 2013). As a result, proteins can break down,
which could potentially cause a loss of membrane permeability. Mean-
while, intermolecular disulphide linkages of carbon - nitrogen terminal
of cysteine amino acid residues cause protein aggregation and eventu-
ally reduce total water-soluble proteins (Yousif et al., 2007). In common
beans, soluble protein fraction, mainly albumin and globulin, appear to
be in an overall decline, whereas insoluble protein may promote in HTC
beans (Coelho et al., 2013). Yousif et al. (2007) detected a substantial
increment in the high molecular weight (HMW) fraction and a decre-
ment in the low molecular weight (LMW) fraction of bean proteins while
suggesting the aggregation and coagulation of protein during storage. In
addition, protein denaturation enthalpy decreases, significantly
reducing the thermal stability of proteins.
Further, SEM images of cytoplasmic structures of cotyledon cells
appear collapsed; for example, partially autolysed protein bodies can be
observed (Berrios et al., 2006). Prolong storage increases the lignified
D. Perera et al.
protein level of cotyledon cells, and there is a strong association between
cooked bean textural quality and lignified protein concentration (Reyes-
Moreno et al., 1993; Nasar-Abbas et al., 2008). As explained in Fig. 1A
HTC column, lignified cell edges, middle lamella, and secondary walls of
HTC beans appear in SEM images but not for the fresh beans.
5.3. The effects of HTC on nutrition quality
Beans are rich in essential nutrients, such as starch, protein, fibre,
minerals, vitamins, and phytochemicals. Hence the consumption of
beans is undermined by the HTC phenomenon, which changes the di-
gestibility and bio accessibility of those essential nutrients. Legumes can
develop the HTC phenomenon after extended storage above 25 ◦ C and
humidity levels exceeding 65%. HTC beans need to prolong cooking
time to soften the cotyledon, thus increasing the solid leaching and
destroying heat-sensitive vitamins and bioactive compounds.
HTC development of common beans increases the starch di-
gestibility, reducing protein’s digestibility. HTC beans had a faster rate
of cell damage and release of cell cytoplasmic content to easily digested
by amylases. As a result, following a specified cooking period, HTC
Bambara groundnuts digested starch faster than fresh seeds (Duijsens
et al., 2021). However, HTC beans significantly decrease protein di-
gestibility due to the complex formation of a protein with other proteins,
starch, hemicellulose, and minerals during storage (Duijsens et al.,
2021). Also, protein-phytate interactions cause blockage of proteolytic
enzymes and decrease protein digestibility and solubility (Ruiz-Ruiz
et al., 2010). During storage, protein coagulates and aggregates due to
the pH drop, which may lead to their solubility and thermal stability,
thus potentially decreasing protein digestibility (Duijsens et al., 2021).
Furthermore, this can be related to lignification, like the hypothesis of
HTC beans. During storage, tannins and lignin monomers can poly-
merise with proteins and carbohydrates, which is a reason for reduced
protein digestibility.
HTC phenomenon also may negatively affect essential minerals such
as calcium, iron and zinc (Duijsens et al., 2021). During storage, solu-
bility and bio accessibility of Ca, Zn, and Fe decrease; however, Mg is not
significantly affected since it has a lower affinity of mineral antinutrient
to magnesium (Rousseau et al., 2020). This may be due to increase
calcium and pectin cross-linking. Several researchers have proposed the
pectin-cation-phytate hypothesis to explain these observations. Ac-
cording to this hypothesis, HTC development in beans would result in
mineral relocation (Gwala et al., 2020; Rousseau et al., 2020). It was
proven that Bambara groundnuts and common beans lose some phos-
phorylation with time, causing the release of phytate-chelated minerals.
Overall, minerals are somewhat difficult to absorb in the small intestine
when consuming HTC beans.
6. Strategies to improve the cooking quality of stored beans
Prolonged beans storage under adverse conditions induces changes
in beans’ structural, compositional and biological composition. Most
stored beans delay hydration during soaking due to hard shell defects or
take longer to soften during cooking due to hard-to-cook defects.
However, beans must be adequately hydrated while soaking and cook-
ing to ensure the quality of cooked beans. This section discusses proper
storage conditions to overcome storage-induced defects and improve the
hydration of beans during soaking and cooking.
6.1. Conventional techniques
Optimal storage conditions are crucial in obtaining good quality le-
gumes. Bean quality can be preserved for an extended period under
controlled storage conditions. However, storage at sub-optimal condi-
tions for a prolonged period will result in hard-to-cook beans that take a
long cooking time, nutrient loss, off-flavour, and off-colour
development.
10
Food Chemistry 415 (2023) 135743
Several studies have investigated the effect of humidity, tempera-
ture, and storage duration on HTC development. Reyes-Moreno et al.
(1993) and Mulu-Abay (2021) reported that moisture content above
13% deteriorates beans significantly after six months at ambient tem-
perature, and beans become unsuitable for consumption after 12 months
of storage. However, on the laboratory scale, beans with a moisture
content of less than 10% can be kept for up to two years without losing
quality (Rani et al., 2013). Hohlberg and Stanley (1987) reported that
HTC defects are mainly found in beans with dark seed coat stored for ten
months at 30 ◦ C under relative humidity (RH) at 85% compared to the
15 ◦ C, RH at 35%, and 25 ◦ C, RH at 65%. Uebersax and Bedford (1980)
found that the navy beans stored at 75% RH or less with 20 ◦ C or lower
temperatures had a better quality. Avila et al. (2015) reported that
common beans’ cooking properties at 2 ◦ C were not changed after one
year of storage. Yi et al. (2016) explained that storage at 35 ◦ C and 75%
RH promoted the HTC condition in beans. According to several studies,
the HTC impact is minimised at refrigerated temperatures (0 to 5 ◦ C).
Regardless of the other variables, beans kept at temperatures below
17 ◦ C or with moisture levels below 8 to 9% do not harden significantly
(Ferreira et al., 2017).
Chigwedere and colleagues (2019a) investigated the stability of
common beans concerning storage and cooking quality and proposed
that storing beans below their Tg preserves the quality of beans. They
also established a relationship between the Tg and moisture levels,
which could help identify suitable storage conditions. The matrix of
beans is heterogeneous, leading to local Tg values that depend on the
local reactions. Kyomugasho et al. (2020) confirmed the importance of
these local reactions in determining the local Tg values. Beans with high
moisture levels typically have lower Tg values and must be stored below
Tg value or at low temperatures. It may be challenging to store beans
effectively in tropical climates where temperatures remain high
throughout the year, and cooling storage facilities are scarce. As a result,
it is crucial to reduce the moisture content of the beans to an appropriate
level using safe methods after harvesting (Bradford et al., 2018). Sun
drying or heated-air drying can lower the moisture content, but it is
essential to be cautious with sun drying as prolonged exposure to tem-
peratures exceeding 25 ◦ C and relative humidity above 65% may cause
heat-induced discolouration.
Patil et al. (2007) examined extrusion cooking to enhance the
nutritional profile of HTC beans stored at 42 ◦ C and 80% RH for six
weeks or more than one year. Beans stored in adverse conditions
develop HTC defects, resulting in longer cooking times. Furthermore,
increasing temperature leads to a reduction in both bulk density and
solubility index in both fresh and HTC beans; however, the water ab-
sorption index increased due to the larger quantity of gelatinised starch
in the extruded samples (Uebersax et al., 2022). Extrusion deactivates
the antinutritional factors such as trypsin, lectins, and amylase in-
hibitors in fresh and HTC beans (Chigwedere et al., 2019b; Uebersax
et al., 2022). The extrusion also resulted in a significant rearrangement
of insoluble dietary fibre to soluble, with the primary sugars implicated
being pectic polysaccharides, arabinose, and uronic acids (Patil et al.,
2007; Ruiz-Ruiz et al., 2008).
Mbofung et al. (1999) introduced a novel way of improving and
optimising the consumption and nutritional quality of HTC beans by
processing them into Koki, a healthy cowpea-based food product typi-
cally prepared by steaming. Incorporating HTC beans with cowpeas to
produce Koki resulted in food with high nutritional value. Adding HTC
red kidney beans is a better choice for producing Koki from composite
pastes, as it reduces trypsin and alpha-amylase inhibitors and improves
the in-vitro digestibility of its protein and carbohydrate. Koki produced
from HTC beans are highly nutritious with an enhanced sulphur amino
acid content due to the complementing action of the combination of
legumes (Mbofung et al., 1999). Wet processing followed by drying can
reduce the cooking time of HTC beans. Steaming of beans results in
about 25% softer than non-steam beans (Reyes-Moreno et al., 1993;
Belmiro et al., 2018). Steam can enhance hydration while decreasing
D. Perera et al.
equilibrium moisture content and lag phase time (Miano and Augusto,
2018a). High temperatures at 100 ◦ C in 1 to 2 s can be employed to
inactivate trypsin inhibitors, although this was insufficient to prevent
HTC development. However, wet processing requires high energy and
alters the texture of the beans.
The phytic acid content of beans can be used to predict their cooking
quality (Yousif et al., 2007; Wainaina et al., 2022a). Soaked beans in
sodium phytate solution minimise cooking time because sodium phytate
serves as a chelating agent, eliminating Ca2+ ions from calcium pectates
during cooking, lowering the cooking time and improving cellular sep-
aration (Luo et al., 2012). Soaking beans in sodium chloride, sodium
carbonate, sodium bicarbonate, sodium phosphate, or ethyl-
enediaminetetraacetic acid (EDTA) solutions dramatically lower the lag
phase and total cooking time (Kinyanjui et al., 2017; Mubaiwa et al.,
2017) by boosting pectin water solubility when compared to beans
soaked in pure water. In addition, it increases the cooking rate, thereby
increasing the uniformity in the softening of the bean samples. Salt used
in soaking water increases the hydration coefficient, reducing the
cooking time, HTC development, cohesiveness, and chewiness of the
beans. Avila et al. (2015) reported that 2.5% sodium bicarbonate and
1% potassium chloride effectively increase the hydration coefficient,
decrease the HTC cowpea, and reduce the cooking time. Further light
microscopy images have proven that beans soaked in sodium salts
resulted in better cell separation (Marston and Omana, 1979; Alpos
et al., 2021). Therefore, monovalent cations of Na+ and K+ soften
cotyledon tissue, with heat swiftly disrupting the significant linkages
between the pectinic acids and increasing pectin solubility (Yousif et al.,
2007).
Germination is a technique that involves soaking beans to stimulate
the process of sprouting. It has been used as a pre-processing step to
improve the nutritional quality of beans, owing to the resulting
biochemical reactions. Germination has been found to increase the
content of certain nutrients and reduce the levels of anti-nutrients in
beans. However, recent research has indicated that germination can also
significantly impact the cooking time of red beans. The study found that
germinated red beans required a shorter cooking time than non-
germinated beans, which could have implications for food processing
and preparation (Haileslassie et al., 2019).
It is recommended that HTC development can be avoided by
improving the storage conditions of legumes. Bean storage in a modified
atmosphere has been reported as a solution for reducing storage-induced
defects in legumes. Perez et al. (2021) have reported that HTC devel-
opment of beans can be reduced by storage under a CO2 atmosphere.
Legumes stored in CO2 or N2 atmospheres suppress respiration and
inhibit enzyme activity and microbial growth. Further, it reduces
oxidation reactions and biochemical changes during storage. However,
it is also imperative that such arrangements may not always be feasible,
with these adding up to the cost of bean production.
6.2. Use of advanced and emerging technologies
Apart from conventional methods, combining different non-thermal
techniques such as ultrasound, high hydrostatic pressure (HHP) treat-
ment, pulsed electric field (PEF), and irradiation has enhanced the hy-
dration of beans. Cell proliferation by ultrasound waves at a higher
frequency (20 kHz) can accelerate the water flow through the legumes
by directly causing pressure differences by compressing and expanding
the medium, increasing the capillary flow, and improving the hydration
(Guimaraes et al., 2020). Ultrasound can cause structural changes
indirectly through sonic cavitation, causing cell and tissue disruptions.
These cavities may improve water transport in HTC legumes (Miano
et al., 2016; Miano et al., 2018a). Ultrasound pushes water to the space
between cotyledon and seed coat through the hilum without changing
the starch granules (Miano et al., 2016). Therefore, ultrasound is a
promising technology for faster legume hydration. Miano et al. (2016)
investigated the use of ultrasonic technology in the hydration of mung
11
Food Chemistry 415 (2023) 135743
beans. They concluded that ultrasonication shortens the hydration
process by around 25% without affecting starch’s structural or pasting
properties.
Recently, scientists have examined how high-pressure treatment af-
fects legume hydration. Belmiro et al. (2018) studied the impact of high-
pressure processing on the hydration of common beans, and their results
revealed that HHP at 600 MPa accelerated bean hydration by 4.7 times.
Furthermore, due to the higher initial hydration, HHP reduces the
cooking time of the beans by 15 min. Ueno et al. (2015) also examined
the effect of high-pressure treatment at 200 Mpa at 25 ◦ C for 10 min on
kidney beans, adzuki beans, and soybean. They found that high pressure
positively affected water absorption into the cotyledon of adzuki beans
and hypothesised that HHP enhances the hydration of beans; firstly,
high pressure is applied to the capillaries and intercellular gaps to fill
with water, followed by a rapid transition of the seed coat from glassy to
rubber behaviour (Ueno et al., 2015; Belmiro et al., 2018). High-
pressure treatments improve water absorption due to structural modi-
fications and boost bean hydration. However, this could be related to the
spherical crystalline structure being disrupted under high-pressure
conditions, whereas the microcrystalline structure may not be dis-
rupted (Li et al., 2015).
Devkota et al. (2022) investigated the effect of pulsed electric field
(PEF) intensity of 4 kV/cm at 2 Hz frequency, and 15 μs pulse width
assisted hydration of two common bean varieties and concluded that
PEF treatment reduced the hydration time by 3-fold for small red kidney
beans. The thick seed coat of beans is the primary barrier to water up-
take, and the formation of tiny pores can overcome this on the seed coat
via PEF, which increases the water transfer through the seed coat (Alpos
et al., 2022). Therefore, PEF is an emerging technique for improving the
hydration of legumes.
In addition, irradiation has been used to improve non-thermal
technology. Ramaswamy et al. (2005) irradiated navy beans using
60
Co γ -rays and identified that the combination of high pressure and
irradiation treatments improved the hydration of beans during the over-
extended soaking time (>3h), increasing equilibrium moisture content.
The cooking time for dry beans is significantly decreased when irradi-
ation dosages of 1, 5, and 10 kGy (Celik et al. 2004). For carioca beans
exposed to radiation at doses ranging from 1 to 10 kGy, similar results
were reported by Lima et al. (2019). The breakdown of the membrane
during irradiation results in increased water absorption qualities during
cooking, which was thought to cause this reduction in cooking time.
Naviglio et al. (2013) developed a circularly pressured soaking phase
to hydrate cannellini beans at room temperature. This treatment differs
from HHP technology since cyclic pressure is not applied for high
pressure. This novel process reduced the hydration time of the beans to
approximately 60 min and improved water permeability by creating a
pressure gradient inside the grains and enhancing capillary flow.
Furthermore, Demirhan and Ozbek (2015) have reported that micro-
waves can improve the hydration of cowpeas. Microwaves might be
transferred energy to the water molecules by regulating their motions
(Divekar et al., 2017). However, more research is required to investigate
the effects of HHP, PEF, ultrasound, irradiation, microwave and cyclic
pressure application treatments on stored legume hydration.
7. Conclusion
Hard to cook phenomenon contributes to reduced consumption of
beans all over the world. Several hypotheses are proposed to explain the
mechanism behind the HTC phenomenon, with the “pectin-cation-phy-
tate” hypothesis recently becoming the most widely accepted. This hy-
pothesis describes how high storage temperatures and relative humidity
increased phytase activity, resulting in phytate dephosphorylation and a
lower chelate capacity of divalent cations of calcium, iron, and mag-
nesium. However, other mechanisms also deserve further examination.
Given the complexity of the food matrix, it is reasonable to assume that
several mechanisms might play a holistic role in causing HTC legumes
D. Perera et al.
rather than one individual mechanism. This, however, needs to be
further studied on how individual factors combine to result in an HTC
phenomenon that resists hydration and cooking. It also becomes
apparent from the review that macro and micronutrient composition
deteriorate due to HTC leading to lower bioavailability of these nutrients
for human consumption and, therefore, diminished health benefits.
Besides the thermal and non-thermal treatment strategies for miti-
gating HTC, current research is moving towards developing new vari-
eties of legumes with reduced phytase enzyme and examining how it
impacts HTC. Finding a more efficient solution to the HTC phenomenon,
reducing cooking time, enhancing nutritional content, and improving
sensory qualities of HTC beans are becoming significant and will impact
the future food industry and legume consumption. Based on the current
knowledge, it is essential to implement recommended solutions to pre-
vent HTC development and optimise the storage conditions to minimise
the effects. Moreover, a clear focus on elucidating HTC mechanisms
using advanced analytical methods is required to maintain the sustain-
able consumption trend of legumes.
Data availability
The data (cited reference materials) is available on request from the
corresponding author.
CRediT authorship contribution statement
Dilini Perera: Conceptualization, Methodology, Investigation,
Formal analysis, Writing – original draft, Writing – review & editing.
Lavaraj Devkota: Supervision, Conceptualization, Methodology,
Writing – review & editing. Gil Garnier: Supervision, Conceptualiza-
tion, Methodology, Writing – review & editing. Joe Panozzo: Supervi-
sion, Conceptualization, Methodology, Writing – review & editing.
Sushil Dhital: Supervision, Conceptualization, Methodology, Writing –
review & editing.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
Data will be made available on request.
Acknowledgement
The authors would like to acknowledge the support from the Monash
graduate research scholarship (Dilini Perera) and funding from Austra-
lian Research Council Linkage grant LP210200616.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.foodchem.2023.135743.
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