Trends in Food Science & Technology 19 (2008) 309e319

Review

Texture changes
of processed fruits
and vegetables:
potential use
of high-pressure
processing
Daniel N. Silaa,
Thomas Duvettera,
Ans De Roecka, Isabel Verlenta,
Chantal Smouta,
Graham K. Moatesb,1,
Brian P. Hillsb,1,
Keith K. Waldronb,1,
Marc Hendrickxa,* and
Ann Van Loeya
a

Center for Food and Microbial Technology, Faculty of

Bioscience Engineering, Katholieke Universiteit
Leuven, Kasteelpark Arenberg 22, B-3001 Heverlee,
Belgium (Tel.: D32 16 32 15 72; fax: D32 16 32 19 60;
e-mail: marc.hendrickx@biw.kuleuven.be)
b
Institute of Food Research, Norwich Research Park,
Colney, Norwich NR4 7UA, UK

In processed fruits and vegetables, changes in texture are

strongly related to transformations in cell wall polymers due
to enzymatic and non-enzymatic reactions. A major challenge
is how to use recent advances in processing technologies and
to adjust raw materials, ingredients and processes to improve
texture of processed plant based foods.
* Corresponding author.
1
Tel.: 44 1603 255000; fax: 44 1603 507723.

0924-2244/$ - see front matter ! 2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.tifs.2007.12.007

This review focuses on the plant cell wall structure and the
processing dependent changes in plant cell walls with focus
on enzymatic and non-enzymatic degradation of pectin.
Stability as well as catalytic activity of two major plant endogenous pectin degrading enzymes, namely pectinmethylesterase and polygalacturonase, towards elevated pressure and
temperature is reviewed. Finally, the effect of processing on
texture of plant based foods and different approaches to
improve the texture of processed plant based foods (i.e.
preheating, phenolics, washing/dipping/infusion pretreatments, high-pressure pretreatments and genetic modification)
are discussed.

Plant cell wall structure in relation to

texture of plant based foods
Understanding of the plant structure is highly relevant
for developing new approaches for improving the quality
of plant based foods. Therefore, an in depth evaluation of
the interactions involved from molecular (polymeric) to tissue/organ level is needed. Models of the primary cell walls
have remained relatively unchanged since the work of Albersheim et al. (Keegstra, Talmadge, Bauer, & Albersheim,
1973; McCann & Roberts, 1991). While adopting the same
framework, more recent models project the macromolecular organization of the wall, including the transient
hierarchy from the molecular to the tissues/organs level
(Fig. 1) (Waldron, 2004).
Insight can be gained at molecular, microscopic/mesoscopic and macroscopic level by monitoring the modification in the structural features of the system which is
paralleled by alteration in the functional properties. An integrated approach which correlates the changes at different
levels is important in order to determine the key influencing
factors.
In most models, the cell wall is represented as a heterogeneous and dynamic polymeric unit comprising of a threedimensional inter linked fibrous structure of cellulose
embedded in a complex matrix of pectin, hemicellulose,
structural proteins and some phenolics (Carpita & McCann,
2000). The wall is highly hydrated (w65% water) and
contains various dissolved solutes, ions and soluble proteins
including enzymes. The relation between the basic structural models and physical properties has been described
using the sticky network, multi-coat network and hybrid
models (Cosgrove, 2000; Thompson, 2005). None of the

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D.N. Sila et al. / Trends in Food Science & Technology 19 (2008) 309e319

ORGAN

TEXTURE

TISSUE

CELL

CELL WALL

POLYMER
Physiological change

Fig. 1. The hierarchy of structures within plants (Waldron, 2004).

models explains all the measured physical properties of

plant materials (Brummell, 2006). The unique interaction
of these components and the changes during growth,
maturation and processing in relation to the mechanical
properties are still open for discussion.
To solve processing dependent changes in plant based
foods, it is important to understand the structureefunction
relationship of cell wall polymers. The interaction between
the structural components of plant based foods and the transient changes involved during post-harvest storage and/or
processing are very important in relation to their texture.
Mostly, in view of structure and functional properties, pectin is regarded to be the most interesting cell wall polymer
because of its abundance, its solubility and its sensitivity to
chemical reactions (Van Buren, 1979). The ubiquitous presence of pectin in fruits and vegetables coupled with its thermal sensitivity emphasizes its critical importance in
determining the texture of many processed plant derived
products. Specific biochemical and/or chemical reactions
occur on pectin which may be valuable or deleterious depending on the intentions of the process and the (pre-)processing steps involved.

Changes in cell wall components during

processing of plant based foods
Cellulose and hemicellulose show minimal changes in
structure and composition within the thermal processing
domain ("150 # C) of most plant based foods (Williams
& Besler, 1996). Most of the changes observed in thermally processed plant based foods are ascribed to transformations in pectin structure and composition. These
changes are strongly influenced by the raw material
properties, the (pre-)processing steps and conditions.
Generally, changes in pectin structure and composition
may arise from enzymatic and/or non-enzymatic reactions. Both reaction mechanisms are imperative for

processed plant based foods. However, the identification

and the quantification of the changes are hindered to
some extent as it can be extremely difficult to extract
native cell wall polysaccharides without modifying their
structure, and it is even more challenging to isolate
polysaccharide complexes for in vitro assays without
disrupting inter- and intramolecular associations or generating artefacts.
Enzymatic cell wall degradation and disassembly
In plant cell wall, each structural component/aggregate
of matrix components is targeted by multiple modifying enzymes. It is probable that one modifying mechanism synergistically alters the action or the efficacy of another (Rose,
Saladie, & Catala, 2004). Numerous ways exist in which
such synergies might occur, especially given the structural
specificity of many cell wall hydrolases and lyases. In that
perspective, the principal pectin degrading enzymes are
pectinmethylesterase (PME E.C. 3.1.1.11), polygalacturonase (PG E.C. 3.2.1.15) and pectate lyase
(PL E.C. 4.2.2.2) (Willats, Knox, & Mikkelsen,
2006). However, other known and unknown enzymes may
be important. PME, an abundant pectinase in plant based
foods, catalyzes the demethoxylation of pectin resulting
in the formation of carboxylated pectin with release of
methanol. The demethoxylated pectin molecule can (1)
cross-link with divalent ions like calcium forming supramolecular assemblies and/or gels (firming effect) and (2) be
a substrate for PG depolymerization (softening effect).
PG, synthesized by some higher and lower plants, bacteria
and fungi, catalyses the hydrolytic cleavage of the O-glycosyl bond of demethoxylated a-D(1e4)polygalacturonic
acid. PME and PG are not the sole determinants of softening, although their concerted action is revealed in many
quality related aspects. As indicated earlier, pectin can be
degraded by the successive demethoxylation and depolymerization by PME and PG, respectively. The activity of
these enzymes is restricted in planta, with maximum hydrolytic potential expressed only in response to tissue disruption/wounding and ripening. For plants with high PG
levels like tomatoes and avocadoes, quality (texture and
rheology) loss due to pectin depolymerization is eminent.
In other plants like carrots, bananas, strawberries and
melons, the endogenous PG activity is negligible (Tucker,
2004). Therefore it is of no consequence with regard to
the mechanical properties of such products. On the other
hand, PL, which has mostly been associated with pathogenesis (bacteria and fungi infestation), is nowadays acknowledged to be expressed in plant tissues (Chourasia, Sane, &
Nath, 2006; Goulao, Santos, de Sousa, & Oliveira, 2007;
Medina-Escobar, Cardenas, Moyano, Caballero, & Munoz-Blanco, 1997; Payasi & Sanwal, 2003). PL depolymerizes pectin following the b-elimination reaction
mechanism. It can either be endo- or exo-acting and the activity may be restricted to short oligosaccharides. Other enzymes like b-galactosidases/exo-galactanases can occur

D.N. Sila et al. / Trends in Food Science & Technology 19 (2008) 309e319

endogenously, but exhibit in vitro activity far below the anticipated level for cell wall loss/degradation. To date, the enzymatic degradation of the cell wall remains complex and it
appears that a multiple class of enzymes and probably other
proteins are responsible.
Non-enzymatic cell wall degradation and disassembly
Pectin shows a high stability in aqueous solutions at
pH 3.0e4.0. Nonetheless, three processes are known so
far that lead to non-enzymatic pectin degradation. First,
the fragmentation of the polymer can occur in planta
due to highly reactive hydroxyl radicals (Fry, Dumville,
& Miller, 2001). Increasing evidence suggests that these
reactions are part of the mechanisms that drive wall restructuring, but their significance in the degradation of
processed plant based foods is not known. During thermal
processing, pectin may undergo either acid or base catalyzed depolymerization. At low pH ("3.5), acid hydrolysis is proposed despite some recent doubts (Krall &
McFeeters, 1998). Under these conditions, which are not
common during regular food handling and processing,
low methoxy pectin depolymerizes faster than high methoxy pectin. At higher pHs (%4.5) and at elevated temperatures (>80 # C), a prevalent condition in most
thermally processed plant based foods, base catalyzed depolymerization (b-elimination reaction) of pectin is quite
relevant (Albersheim, Neukom, & Deuel, 1960; Greve,
McArdle, Gohlke, & Labavitch, 1994). A prerequisite
for the b-elimination reaction is the presence of methyl
ester group at the C-6, which renders H-5 sufficiently
acidic to be removed by an alkali. The reaction rate increases as pH increases, because hydroxyl ions initiate
the reaction (Neukom & Deuel, 1958).
Since the b-elimination reaction is highly dependent on
the methyl ester content of the pectin in plant tissues
(Sajjaanantakul, Van Buren, & Downing, 1989; Waldron,
Smith, Parr, Ng, & Parker, 1997), targeted manipulation of
the degree of methylesterification of pectin can lead to
controlled b-elimination. This can be achieved in vivo
through genetic engineering or by in situ activation of endogenous or exogenous PME using pretreatment conditions (cfr.
later). The beneficial effect of the latter has been shown for
thermally processed carrots (Vu et al., 2004). Brine ingredients affect the b-elimination reaction in different ways and

311

extents, however, increasing the additive concentration

increases the b-elimination reaction constant and the texture
degradation rate constant, while the residual texture value
and the degree of methylesterification of the sample decrease
(Vu, Smout, Sila, Van Loey, & Hendrickx, 2006).
Sequential extraction using chemical solvents as
described by Redgewell and Selvendran (1986) can be
used to assess changes in the relative solubility of different
classes of polysaccharides following thermal processing.
The polysaccharides released by the sequential extractions
are predominantly pectic in nature.
The dissolution of the wall polymers is concurrent with
tissue softening which involves cell separation (Lecain, Ng,
Parker, Smith, & Waldron, 1999; Ng & Waldron, 1997)
(Fig. 2). The strength of cell adhesion is likely to be dependent,
to a large extent on the strength of cellecell interactions in the
middle lamella adjacent to the intercellular spaces. Indeed,
maximum tissue softening only occurs after separation in
this zone. Numerous studies have been carried out to compare
and correlate cell separation and dissolution of pectic polysaccharides (Sila, Doungla, Smout, Van Loey, & Hendrickx,
2006) often involving mathematical modeling of the texture
(Tijskens, Waldron, Ng, Ingham, & van Dijk, 1997).
Current thermal processing methods such as pressure
cooking have been shown to have a major effect on the
extractability of cell wall polysaccharides (Lecain et al.,
1999). Substantial increases were observed in total pectic
polysaccharides within the water-soluble, salt-soluble and
imidazole-soluble fractions; accompanying decreases
were seen in the total pectic polysaccharides solubilised
by cyclohexane-trans-1,2-diaminetetra-acetic acid, sodium
bicarbonate and 0.05 M potassium hydroxide and, in
particular, a relatively large decrease in the level of total
pectic polysaccharides in the residue. These results indicate that the heat-treatment-induced modifications occur
in most, if not all, pectic components in the cell wall,
and are probably due to an increase in heat degradation
through b-elimination resulting in base-catalysed
depolymerisation.
Extrusion cooking also results in an increase in the
solubility of the pectic polysaccharides and hemicelluloses
as well as an increase in the swelling of the cell wall (Ng,
Lecain, Parker, Smith, & Waldron, 1999). Thibault, Ralet,
Axelos, and Della Valle (1995) have demonstrated the

Fig. 2. Light micrographs of onion outer scale leaves: (A) fresh, (B) pressure cooked (20 min) and (C) pressure cooked (50 min).

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D.N. Sila et al. / Trends in Food Science & Technology 19 (2008) 309e319

possibility of obtaining gelled systems directly during

extrusion cooking of citrus fibre. Their results show that
extrusion cooking is less degradative towards the sidechains than conventional acid treatment.
Stability and catalytic activity of pectinases
towards elevated temperature and/or pressure
Process stability of pectinases
Enzymes, like other proteins, are stable within a certain
pressureetemperature domain. Exceeding these limits disturbs the forces stabilizing the three-dimensional protein
structure, causing unfolding and denaturation of the
molecule, and hence, inactivation. The temperature and
the pressure inactivation kinetics of enzymes have been
described by several models depending on the source
and the pressureetemperature domain studied. The
models most commonly applied are the first-order model,
the distinct-isozyme model and the fractional conversion
model. The temperature dependence of the inactivation
rate constant k (at constant pressure) is often expressed
as activation energy, as indicated by the Arrhenius equation. The pressure dependence of the inactivation rate constant can be expressed as an activation volume, as
indicated in the Eyring model (Van Loey, Indrawati,
Smout, & Hendrickx, 2002).
The combined temperatureepressure dependence of the
inactivation rate can be presented by isorate contour plots,
connecting temperatureepressure combinations resulting in
the same inactivation rate. These, most often elliptical,
pressureetemperature kinetic diagrams have been modeled
on a thermodynamic basis (e.g. Heremans & Smeller, 1998;
Van Loey et al., 2002). For protein denaturation and
enzyme inactivation often an antagonistic effect of pressure
and temperature is observed, namely moderate pressure
retards thermal denaturation/inactivation. This phenomenon
is attributed to counteracting effects of pressure and
temperature on the formation and disruption of intramolecular bonds and/or to their opposing effects on interactions
between protein and solvent (Heremans & Smeller, 1998;
Knorr, Heinz, & Buckow, 2006; Smeller, 2002).
Pressureetemperature inactivation
of pectinmethylesterase
Many studies have been performed on the pressuree
temperature inactivation of PME from plant sources: pepper (Castro, Van Loey, Saraiva, Smout, & Hendrickx,
2006), tomato (Crelier, Robert, Claude, & Juillerat,
2001; Fachin et al., 2002), apple (Denes, Baron, & Drilleau, 2000), orange (Nienaber & Shellhammer, 2001),
white grape fruit (Guiavarch, Segovia, Hendrickx, &
Van Loey, 2005), banana (Ly Nguyen, Van Loey, Fachin,
Verlent, & Hendrickx, 2002) and carrot (Ly Nguyen
et al., 2003) PME. In case of microbial PMEs, detailed
conformational stability and inactivation studies on recombinant Aspergillus aculeatus PME during combined
pressureetemperature treatments have been outlined

(Dirix, Duvetter, Van Loey, Hendrickx, & Heremans,

2005). Generally, PMEs are rather heat labile enzymes
since most of the (purified) enzymes readily inactivate
at temperatures below 70 # C. Contrary to their temperature sensitivity, however, PME can be regarded as a pressure resistant enzyme. Pressures higher than 700 MPa
are usually required to induce short-term inactivation at
room temperature. However, some PMEs do not lose activity even after treatments at 900 MPa. As reflected by
the elliptical shape of the pressureetemperature isorate
contour plots, pressure and temperature exhibit an antagonistic effect on the PME stability at moderate pressure
("300 MPa). In other words, moderate pressure retards
the thermal inactivation of PME (e.g. Ly Nguyen et al.,
2003).
Pressureetemperature inactivation
of polygalacturonase
Detailed thermal and pressure stabilities of tomato PG
isozymes, including in vitro conversion reactions with bsubunit (a protein that converts PG2 to PG1), indicate
that PG1 is more thermo-resistant, however, both isozymes
are pressure labile. Conversely, the b-subunit protein is
extremely resistant to heat and pressure as indicated by
its activity after severe thermal and pressure treatments
(Peeters, Fachin, Smout, Van Loey, & Hendrickx, 2004).
It is important to note that tomato PG is more resistant to
thermal denaturation at atmospheric pressure when
compared to tomato PME, however, it is easily inactivated
on moderate temperatureepressure combinations (Crelier
et al., 2001). Therefore, by pressurization, PG can selectively be knocked out of a food system without significantly
affecting the PME content.
Catalytic activity of pectinases
Considering the functional roles of pectinases, the
kinetics of PME/PG catalytic activity is important. To determine the catalytic activity of the enzymes during pressureetemperature treatments, the reaction products of
pectin conversion can be quantified. For PME, the amount
of methanol produced during the demethoxylation of pectin
can be measured. In case of PG catalytic activity, the
formation of reducing groups during thermal or pressuree
temperature treatment can be followed.
Pectinmethylesterase activity at elevated
pressureetemperature conditions
The effect of pressureetemperature combinations on
enhancing PME catalyzed conversion reactions has been
reported for tomato (Verlent, Van Loey, Smout, Duvetter,
Ly Nguyen, & Hendrickx, 2004), carrot (Sila, Smout,
et al., 2007) and recombinant A. aculeatus PMEs (Duvetter
et al., 2006). Substrate conversion rate for purified tomato
PME is optimal at about 45.0 # C at pH 8.0 and about
60.0 # C at pH 4.4 at atmospheric conditions. When
exposed to pressureetemperature combinations, the

D.N. Sila et al. / Trends in Food Science & Technology 19 (2008) 309e319

demethoxylation of pectin by tomato PME increases significantly (5e7 fold): the optimal conditions at pH 8.0 are
55 # C in combination with 300 MPa and at pH 4.4
60 # C in combination with 400 MPa. Optimal conditions
for carrot PME are reported to be 500 MPa and 50 # C (pH
6.5). In case of recombinant A. aculeatus PME, the optimal
conditions are about 50e55 # C depending on the applied
pressure (200e300 MPa). A likely explanation for the
stimulating effect of pressure on PME activity is based
on the assumption that any phenomenon accompanied by
a decrease in volume at constant temperature is favored
by an increase in pressure. Solvation of the charged groups
created by pectin demethoxylation is accompanied by volume reduction resulting from electrostriction, i.e. the compact alignment of water dipoles owing to the coulombic
field of the charged groups. It is important to note that at
low pH, i.e. pH 4.4 and 4.5 chemical demethoxylation of
pectin is not detected. In contrast, a spontaneous chemical
demethoxylation of pectin, which is accelerated by pressureetemperature treatment, is noticed at pH 8.
Polygalacturonase activity at elevated
pressureetemperature conditions
PG activity at elevated pressureetemperature conditions has only been studied for purified tomato PG
at pH 4.4 (Verlent, Van Loey, Smout, Duvetter, &
Hendrickx, 2004). At atmospheric conditions, purified tomato PG shows optimal catalytic activity at 55e60 # C.
When comparing atmospheric pressure conditions with elevated ones, the optimum temperature at elevated pressure
for PG catalyzed hydrolysis of polygalacturonic acid
shifts to lower values at pH 4.4 probably due to inactivation. Tomato PG does not show any activity at 500 MPa
between 25 and 75 # C.
Effect of processing on texture of plant based foods
Kinetics of texture degradation in plant based foods
Kinetics of texture degradation during thermal
processing has been investigated for several plant based
foods (e.g. artichoke, apple, apricot, asparagus, and carrots)
(Rao & Lund, 1986). The texture degradation models have
continuously been evolving: a first-order degradation mechanism (e.g. Huang & Bourne, 1983), a biphasic degradation
mechanism (e.g. Van Loey, Fransis, Hendrickx, Maesmans,
& Tobback, 1995), a fractional conversion model (e.g.
Rizvi & Tong, 1997), neural network and fuzzy logic
modeling (Xie, Xiong, & Church, 1998), etc. More
recently, in situ micromechanical techniques have been
used (Vanstreels et al., 2005).
Progress in modes of improving the texture
of processed plant based foods
The demand for fresh like processed plant based foods
has prompted progressive research in ways of improving
the texture of thermally processed products. Various approaches have been employed including low temperature
blanching (<70 # C) prior to sterilization (e.g. Roy, Taylor,

313

& Kramer, 2001), calcium infusion before thermal processing (e.g. Gras, Vidal, Betoret, Chiralt, & Fito, 2003; Siliha,
Janh, & Gierschner, 1996), pH adjustment during processing (Van Buren & Pitifier, 1992) and exogenous pectinmethylesterase infusion prior to thermal processing (e.g.
Duvetter et al., 2005). All these pre-processing methods
have a common goal namely reducing texture degradation,
but each of them contributes uniquely to the textural value
of the end product.
Preheating
Preheating at mild temperatures (50e70 # C) typically
for %30 min prior to high temperature processing
(%100 # C) has been used to improve the texture of
thermally processed plant based foods. At these conditions, thermal stimulation of cell wall bound pectinmethylesterase demethoxylates pectic polysaccharides increasing
the chances for formation of ionically cross-linked pectin
complexes and reducing the b-elimination reaction. This
reduces the vulnerability of the product to softening. Preheating also increases the permeability of the plasma
membrane triggering a cation influx which boosts PME
activity (Van Buren, 1973). The availability of divalent
ions (exogenous or endogenous) is important (Ng & Waldron, 1997), hence the frequent combination of preheating
with calcium impregnation (calcium soaking and vacuum
infusion).
Phenolics
Other possible firming mechanisms relate to peroxidase
activity. A small percentage of the sugars in the wall carry
ferulic acid and the associated phenolic groups, which may
be cross-linked by the action of peroxidase and hydrogen
peroxide. These cross-linked structures form intercellular
bridges like diferulate and o,o0 -dityrosine, thus connecting
polymers together in a tight network (Brett & Waldron,
1996).
In bamboo shoots, sugar beets and Chinese waterchestnuts, thermal texture stability is linked to diferulic acid
(Parker & Waldron, 1995; Waldron et al., 1997). In the
case of Chinese waterchestnut, ferulic acid-substituents
on thermally-stable arabinoxylan hemicelluloses are involved in cell adhesion, and are particularly concentrated
at the edge of the cell faces where adhesion is controlled
(Waldron & Brett, 2007). Thermal processing of Chinese
waterchestnuts causes starch gelatinization, and the rounding up of cells, but adhesion and texture is maintained.
Only by modifying the diferulate-cross-linked arabinoxylans by chemical or enzymatic means can the cells be separated. Interestingly, the work of Parker, Parker, Smith, and
Waldron (2003) has indicated that such adhesion is dependent on a particular diferulate (8,80 -diferulic acid, aryltetralyn form). In sugarbeet, thermal stability is also influenced
by diferulic acid, but here it cross-links pectic polysaccharides. The peroxidative cross-linking of pectic polysaccharides was highlighted originally by Thibault (1986). Ng,

D.N. Sila et al. / Trends in Food Science & Technology 19 (2008) 309e319

Harvey, Parker, Smith, and Waldron (1998) showed that although sugarbeet could be softened by thermal processing
(through cell separation), this could be reduced by incubating tissues in hydrogen peroxide. This stimulated
endogenous peroxidase enzymes to cross-link pectic ferulic
acid, increasing the cross-links (particularly 8-O-40 diferulic acid and 8,50 -diferulic acid (benzofuran form))
within the pectin network and enhancing cell adhesion.
Washing, dipping or infusion treatments
In view of texture stability of plant based foods, firming
agents such as calcium salts, gelling hydrocolloids, PME
have been applied (Saurel, 2002).
Calcium. To counter the deleterious textural effects in
thermally processed plant tissues, calcium treatments
have been used successfully for firming of many fruits
and vegetables, e.g. cauliflower (Hoogzand & Doesburg,
1961), Jalapeno pepper (Howard, Burma, & Wagner,
1994), and carrots (Vu et al., 2004), due to its ability
to bind with demethoxylated polyuronides (Grant, Morris,
Rees, Smith, & Thom, 1973). The formation of crosslinks between free carboxyl groups of a pectin chain
and calcium ions has led to the widespread use
of calcium infiltration techniques. However, a high
concentration of calcium confers undesirable taste to the
product and promotes the b-elimination reaction (Grant
et al., 1973). The effectiveness of calcium as a firming
agent is amplified by increasing PME activity. This is
demonstrated by the extra benefits attained when combining calcium pretreatment conditions with mild heating
(Smout, Sila, Vu, Van Loey, & Hendrickx, 2005; Vu
et al., 2004). Industrially, empirical procedures are employed during the application of calcium due to lack of
precise knowledge on the mechanisms conferring tissue
stability while the functionally relevant pectic polymers
remain unknown.
Polyamines. The preservation of whole fruits by dipping
them in aqueous solutions like plant hormones (polyamines), which is improved by vacuum application,
is not new (Valero, Martinez-Romero, Serrano, &
Riquelme, 1998). For minimally processed plant based
foods, e.g. strawberries, polyamines have been used to
enhance the firmness of the fruit, but they are less effective than calcium treatments (Ponappa, Scheerens, &
Miller, 1993).
Pectinmethylesterase. Exploitation of endogenous PME
activity is renowned in plant based foods unlike exogenous
PME infusion which has been limited to beverages and
ground/macerated foods. With the invention of vacuumimpregnation technology, PME can be incorporated into
porous solid food matrices containing occluded air such
as strawberries (Duvetter et al., 2005; Guillemin, Degraeve,
Guillon, Lahaye, & Saurel, 2006). However, the technique

is less efficient in materials with impermeable skin or

lacking interior voids (Baker & Wicker, 1996). Use of
exogenous PME is aimed at enhancing the existing endogenous enzyme content or replenishing inactivated enzyme.
Exogenous PMEs of plant origin and microbial PMEs
(fungal and bacterial) are used. Some examples of exogenous plant PME applications include citrus PME in peaches
(Javeri, Toledo, & Wicker, 1991), Valencia orange PME in
mangoes (Banjongsinsiri, Kenney, & Wicker, 2004), citrus
PME in processed red pepper (Jensen & Petersen, 2004),
and tomato PME in strawberries (Duvetter et al., 2005),
while exogenous microbial PME treatments include
recombinant A. aculeatus PME in strawberries (Duvetter
et al., 2005; Suutarinen et al., 2002) and Aspergillus niger
PME in apples (Degraeve, Saurel, & Coutel, 2003;
Guillemin et al., 2006). Combining vacuum infusion of A.
aculeatus PME and calcium salts enhances the texture of
high-pressure processed strawberries significantly (Fig. 3)
(Duvetter et al., 2005).
Despite the potential benefits offered by endogenous and
exogenous PME, the presence of substantial quantities of
pectinmethylesterase inhibitor in some fruits and vegetables
may lower the PME effect (Giovane, Balestrieri, Quagliuolo,
Castaldo, & Servillo, 1995). In fact, the trend is now towards
using more and more microbial PMEs than plant PMEs
because microbial PMEs are (1) readily available commercially, (2) more beneficial in texture improvement and (3)
not inhibited by pectinmethylesterase inhibitor unlike plant
PMEs (DAvino, Camardella, Christensen, Giovane, &
Servillo, 2003; Giovane et al., 2004).
Hydrocolloids. The infusion of gelling agents in plant
based foods allows for texture improvement either by generating intercellular bridges in the pores, by complementing
the plant cell wall network or by forming bonds between
the added hydrocolloids and the cell wall components.
However, the intercellular gel can modify the texture response only where the porous structure is high and the

1200

400 MPa non-infused

400 MPa - VI

b b

1000

Hardness (g)

314

600

d d

d d

800

550 MPa non-infused

550 MPa - VI
d

d d

a a

400
c c

200
0

0,1

cc

cc
5

c c

10

15

c c

20

Treatment time (min)

Fig. 3. Hardness of non-infused and vacuum-infused (VI recombinant A. aculeatus PME (100 Units/ml) and CaCl2$2H2O (0.5% w/v))
strawberries after pressure treatment at 400 and 550 MPa at 10 # C. aed
Means with the same letter indicate there is no significant difference (Tukeys HSD test: P < 0.05) between vacuum-infused or non-infused,
treatment temperature and treatment time (Duvetter et al., 2005).

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D.N. Sila et al. / Trends in Food Science & Technology 19 (2008) 309e319

High-pressure pretreatment condition

One of the emerging technologies exhibiting a great
potential in preserving plant based foods quality is highpressure processing. Using high-pressure treatment as the
main processing technology eliminates the adverse effects
of heat severity. In view of enzymatic activity and in
relation to texture preservation, high pressure enhances
the catalytic activity of PME significantly (Duvetter
et al., 2006; Sila, Smout, et al., 2007; Verlent, Van Loey,
Smout, Duvetter, Ly Nguyen, & Hendrickx, 2004). For
thermally processed carrots, adopting it as a pre-processing
tool in combination with mild temperatures diminishes
thermo-softening remarkably (Sila, Smout, Vu, & Hendrickx, 2004; Sila, Smout, Vu, Van Loey, & Hendrickx,
2005). This is exemplified by a 9-fold reduction in the thermal softening rate of high-pressure pretreated (400 MPa,
60 # C, 15 min) calcium soaked carrots (0.5% w/v) compared to conventionally pre-processed samples (90 # C, 4
min) (Fig. 4). More recently, Sila, Yue, et al. (2007)
investigated texture changes of non-pretreated and highpressure pretreated carrots during thermal processing using
a multiscale approach, combining molecular (i.e. pectin
characterisation (solubility, degree of methoxylation, and
b-elimination)), mesoscopic (microscopic analysis of tissue
(Fig. 5) and tissue-failure characteristics) and macroscopic
(i.e. hardness analysis by compression and cutting)
analyses.
Understanding the effects of combined pressuree
temperature treatments on plant texture at the sub-cellular
level is a major challenge. Optical microscopy, as well as
scanning electron microscopy, and atomic force microscopy have been used to visualise the effects of pressure
on cell walls and organelles structures (Hills, Costa,
Marigheto, & Wright, 2005; Marigheto, Vial, Wright, &
Hills, 2004) but texture is also affected by the rupture
of membranes and the accompanying microscopic redistribution of water as well as by the gelatinisation of cell
biopolymers and there are few techniques capable of
monitoring these changes on the sub-cellular distance
scale.
Multidimensional
cross-correlation
NMR

Compression Force (N)

60
115 C
120 C
125 C

50
40
30
20
10

tro
l
co
n

C
90

Ca

C
60

60
+

Ca
Ca

P
H
+

C
60

Ca

Ca

0
H

solute gain is significant, especially as mass transfer phenomena are generally limited by the high viscosity of gelling agent solutions.
Among the well-known applications, the vacuum treatment of button mushroom (Agaricus bisporus) with xanthan
gum before blanching and canning has been shown to improve the weight yield and the organoleptic quality of the
final product (Gormley & Walshe, 1986). Preliminary vacuum impregnation of strawberries in solutions containing
gelling agents was proposed by Cierco (1994) as a new
method for improving the quality of frozen fruits. Matringe,
Chatellier, and Saurel (1999) showed the possibility of
introducing various gelling hydrocolloids (gelatin, pectin,
alginate and starch) through the application of vacuum on
fresh apple pieces before freezing.

Pretreatment conditions
Fig. 4. Effect of pretreatment on the texture of carrot discs after thermal
processing at different temperatures for equivalent processes
(Fo 6 min): HP Ca, high-pressure pretreatment (400 MPa,
60 # C for 15 min) followed by calcium soaking, Ca HP, calcium
soaking prior to high-pressure pretreatment (400 MPa, 60 # C for
15.0 min), 60 # C, preheating at 60 # C for 40 min (low temperature
blanching), 60 # C Ca, low temperature blanching followed by
calcium soaking, Ca 60 # C, calcium soaking prior to low temperature blanching, Ca, calcium soaking only, 90 # C, conventional
high temperature blanching (preheating at 90 # C for 4 min), and
control, non-pretreated samples (Sila et al., 2005).

relaxometry is one particularly promising technique for

monitoring sub-cellular water redistribution and biopolymer gelation. A T1eT2 proton spectrum gives peaks
from water in different plant cell organelles (Hills, Marigheto, Wright, & Benamira, 2004) and has been used to
explore the effects of high pressure on strawberries (Marigheto et al., 2004) and potato (Hills et al., 2005). A onedimensional NMR water proton transverse relaxation
study has also been used to monitor the effects of highpressure processing on cod fish (Lambelet, Renevey,
Kaabi, & Raemy, 1995) and meat (Bertram, Wu, Straadt,
Aagaard, & Aaslyng, 2006).
The pressure-induced denaturation (and gelation) of cell
proteins is another factor influencing texture in these
products and a great deal of work has been done on the
effects of HP on protein structure (Cheftel & Dumay,
1998). Unfortunately this work is usually performed on
purified, extracted biopolymers, and it remains to be seen
whether the multidimensional cross-correlation NMR
methods can succeed in monitor similar to biopolymer
conformational changes in cellular tissue.

Genetic modification of matrix polymers

To date, genetic manipulation of cell wall polymers has
focused on modification of cell wall enzymes. Polysaccharide modification in vivo is based on genes encoding biosynthetic enzymes or components of synthase complexes.
Depending on the intention, up or down regulation of the
activity of the enzymes is possible by gene silencing and
selective knockouts (Brummell & Harpster, 2001). Potato
tuber pectin (rich in galactan) has been modified

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D.N. Sila et al. / Trends in Food Science & Technology 19 (2008) 309e319

Fig. 5. Micro-structural images indicating changes in cell wall area to cell vacuole for (a) non-pretreated, non-cooked carrots, (b) non-pretreated
cooked carrots (100.0 # C, 120.0 min), (c) high pressure pretreated, non-cooked carrots and (d) high pressure pretreated cooked carrots
(100.0 # C, 120.0 min). Each micrograph represents four fused snapshots taken at a magnification (&40) (Sila, Yue, et al., 2007).

significantly by expressing a rhamnogalacturonan I degrading lyase which reduces the levels of arabinan and galactan
in the tubers altering the biophysical properties (Ulvskov
et al., 2004). In tomatoes, introduction of an anti-sense
gene for PME is known to have created high solid fruits
with minimal PME activity (Tieman, Harriman, Ramamohan, & Handa, 1992). The decreased PME activity was associated with a 30e70% decrease in bound Ca2 and Mg2
in transgenic pericarp (Tieman & Handa, 1994). Pectin solubility in transgenic plant cell walls can be controlled by
modulating pectinmethylesterase/endo-polygalacturonase
digestion (Tucker, 2004). A significant control in texture
may be realized by using a combination of enzyme knockouts as opposed to a single enzyme activity, especially if
these are targeted to different domains within the wall.
Conclusion
Knowledge on the effect of processing on the texture of
plant based foods and the related correlations with pectin
conversions is still inadequate. Whether the future for
texture improvement of processed fruits and vegetables
will depend on chemical and/or enzymatic approaches or
the in planta genetic modification approach is not clear.
Also a combination of approaches (e.g. combining the
infusion of PME with the impregnation of gelling and/or
firming agents) may be adopted in the future to improve
the mechanical properties of plant based foods. Although

it has clearly been demonstrated that a high-pressure

pretreatment significantly improves texture of thermally
sterilized vegetables, the effects of high-pressure sterilization on the texture of plant based foods and the related
pectin conversion reactions remain to be fully explored.
Acknowledgments
This literature review has been carried out with financial
support from the Commission of the European Communities, Framework 6, Priority 5 Food Quality and Safety,
Integrated Project NovelQ FP6-CT-2006-015710.
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