Current Views On Pectin Substances: Yu. S. Ovodov

ISSN 1068-1620, Russian Journal of Bioorganic Chemistry, 2009, Vol. 35, No. 3, pp. 269284. Pleiades Publishing, Ltd.
., 2009.
Original Russian Text Yu.S. Ovodov, 2009, published in Bioorganicheskaya Khimiya, 2009, Vol. 35, No. 3, pp. 293310.

269

INTRODUCTION
Over the last ten years since the publication of our
review devoted to plant polysaccharides and their struc-
tures and properties [1], great progress has been made
in studies of natural compounds of this class.

2

Particu-
lar attention has been given to pectin substances, one of
the most complex classes of plant polysaccharides.
This review concerns just these compounds, and the
data reported for the most part after 1998 are presented.
The references to earlier works are made only in the
historical aspect.

Denition and the Main Properties

Pectin substances (pectins) are assigned to the large
group of glycanogalacturonans, acidic plant polysac-
charides in which the backbone consists of 1,4-linked

-

D

-galacturonic acid residues [25]. Pectin sub-
stances involve protopectin, pectin polysaccharides,
and the concomitant arabinans, galactans, and ara-
binogalactans [6].
Protopectin is an insoluble high-molecular-weight
pectin complex; together with cellulose and hemicellu-
loses it forms the backbone of a cell wall and, when
treated with diluted acids, gives soluble pectin
extracted usually from plant material [7, 8]. There are
only scarce data regarding protopectin structure.
Pectin polysaccharides include both components of
insoluble protopectin and soluble biopolymeric constit-
uents of plant juices.
The concomitant arabinans, galactans, and ara-
binogalactans have, as a rule, a complex branched
structure.
Pectin substances are found in all higher owering
plants. They are the constituents of cell walls and,
together with other components, determine the rigidity
and elasticity of cell walls, turgor, and the resistance of
plants to drought and low temperatures. Pectin sub-
stances provide watersalt exchange, exhibit a high
gelation capacity, play an important role in the nutrition
of humans as components of food bers, and have a
wide spectrum of physiological activity [112]. The
structure of pectin substances depends on many factors
and can substantially vary during the growth and devel-
opment of the plant. It is not surprising that pectin
polysaccharides are considered as one of the most com-
plicated and structurally dynamical classes of biopoly-
mers.

Methods of Structural Study of Pectins

For establishing the structures of pectin polysaccha-
rides, all known classic and modern methods of struc-
tural studies of polysaccharides are widely used. The
primary structure is determined using complete and
partial acid hydrolysis, enzymatic digestion, periodate
oxidation (Smith degradation), the methylation analy-
sis, all types of NMR spectroscopy (one- and two-
dimensional), and others. In the last few years, AFM
has been used to determine the spatial arrangement of
carbohydrate chains [1323].
We will dwell more closely only on the last method.
AFM was rst used in studies of structures of pectin
substances in 1997 [13]. The method enables one to see

Current Views on Pectin Substances

Yu. S. Ovodov

1

Institute of Physiology, Komi Science Centre, The Urals Branch, Russian Academy of Sciences,
ul. Pervomaiskaya 50, Syktyvkar, 167982 Russia

Received September 22, 2008; in nal form, November 10, 2008

Abstract

This review concerns pectin substances, the most complex class of plant polysaccharides. For the
most part, the data reported after 1998 are presented; the references to earlier works are made only in the his-
torical aspect. New data on the structure of pectin substances, their physiological activity, their role in plants,
and their valuable physical properties are surveyed.

Key words: plant polysaccharides, pectin polysaccharides, pectins, pectin rheology, physiological activity.

DOI:

10.1134/S1068162009030017

1

Corresponding author; phone/fax: (8212) 24-10-01; e-mail:
ovoys@physiol.komisc.ru

2

Abbreviations: AceA, aceric acid (3-

C

-carboxy-5-deoxy-

L

-xylo-
furanose); AFM, atomic force microscopy; DHA, 3-deoxy-

D

-

lyxo

-heptulosaric acid; DM, degree of methylesterication;
KDO, 2-keto-3-deoxy-

D

-manno-octonic acid; LM, low degree of
methylesterication; LMA pectin, amidated pectin with a low
DM; HM, high degree of methylesterication; RG-I and RG-II,
rhamnogalacturonans I and II; GIT, gastrointestinal tract.

REVIEW

270

RUSSIAN JOURNAL OF BIOORGANIC CHEMISTRY

Vol. 35

No. 3

2009

OVODOV

the pectin molecule as a whole, i.e., its backbone and
side chains. An AFM image gives immediately a pic-
ture of all molecules [13]. Using AFM images, it is pos-
sible to characterize mixtures of pectin molecules and
individual molecules inside a particular mixture on the
basis of their dimensions and shape. It was proposed
using this method that pectin from the cell walls of a
green tomato has a branched galacturonan backbone
[14]. Later, this suggestion was conrmed experimen-
tally for comaruman, a pectin from the marsh cinque-
foil

Comarum

palustre

L. [17].
By comparing the AFM images of two pectin frac-
tions extracted from a tomato by different methods, a
correlation between the length of side chains and the
content of neutral monosaccharides was established. In
pectin fractions, mixtures of separate polymer mole-
cules and their aggregates were found [14]. Aggregates
are observed even after strong dilution, when only a few
macromolecules are discernible. This indicates that
pectin substances exist as a multipolymer complex in
which individual components are coupled by intermo-
lecular interactions. An aggregate of separate polymers
reects the heterogeneity of a pectin polysaccharide
sample. Linear polymers with a chain length from tens
to hundreds of nanometers are clearly seen, together
with branched polymers carrying the side chains.
The methylation analysis provides information
about the middle member of a mixture of macromol-
ecules. It should be born in mind that some branches are
present as short chains which are difcult to detect by
AFM. In this case, the results obtained by the methyla-
tion studies are also of great importance [13, 14].
A study of the physical structure of pectin from a
fresh sugar beet by AFM showed for the rst time that
the pectin extract contains a mixture of linear pectin, a
branched pectin polysaccharide, and a complex of pec-
tin with a protein linked to one end of the pectin chain
[24]. Previously [25], it has been proposed that there is
a covalent link between pectin and the protein. The data
obtained by AFM conrm this proposal but are not
direct evidence for the presence of this link: the possi-
bility of the physical association between pectin and the
protein is not ruled out. Further studies are needed to
identify the type of the proteinpectin bond in com-
plexes and determine the role of pectin and the protein
in the complex and the plant cell wall [24].
Finally, by using AFM it is possible to obtain struc-
tural information regarding supramolecular nano-sized
particles of a native pectin polysaccharide that are eas-
ily observable by this method [26, 27].
Most recently, AFM has been used to study the
structure of pectin (from the albedo of an orange peel)
in aqueous solutions (water and citrate buffer) and of
gel networks formed by pectin [28]. Varying the con-
centration of pectin in solutions has resulted in greatly
different AFM images. The results obtained have led to
some important conclusions. Thus, pectin in pure water
at concentrations above 10

g/ml forms aggregates of
network structures; at lower concentrations, molecular
network structures dissociate to their constituent com-
ponents. These components tend to expand due to inter-
charge repulsion along the carbohydrate chain, a phe-
nomenon known as the polyelectrolyte effect. At a
concentration of 6.5

g/ml, small bars, segment rods,
rings, screwed bars, branched molecules, and dense
ring-shaped bodies are observed. The same structures
are also distinguishable at a concentration of about
13.1

g/ml, at which pectin macromolecules form
molecular networks in both pure water and a citrate
buffer. At a concentration of 10

g/ml, pectin in the gel
can be xed in an unstable state due to insufcient sal-
vation of the pectin network in the presence of a high
concentration of sucrose in acidic citrate buffer [28].
A very important area of the application of AFM is
a study of pectin substances as food bers for the food
industry, which involves qualitative and quantitative
assays of the structures of food bers and depicts the
properties of food bers and their changes during pro-
cessing and storage [26].

A General Scheme of Pectin Structure

A general structure al pattern of pectin polysaccha-
rides is given in many papers, e.g., in [29, 30].

The linear region

of homogalacturonan consists of
1,4-linked

-

D

-galactopyranosyluronic acid residues.
These regions are joined by one or two

-

L

-rhamnopy-
ranose residues which are involved in the linear chain
by a 1,2-linkage. The backbone of many pectins has
this very structure. They differ only by the length of the
chain [2931].

The ramied region

consists of three subunits: RG-I,
arabinogalactan, and xylogalacturonan, which can be
present in different ratios, as it is shown for apple pectin
[32]. The data obtained for pectins from a peach, carrot,
onion, leek, and potato are fully consistent with this
structure [31]. In some cases (zosteran, a pectin from
sea phanerogram, and lemnan, a pectin from duck-
weed), the polymer contains an apiogalacturonan frag-
ment [1, 6].
The RG-I fragment considerably varies in different
pectins. Its backbone consists of alternating residues of
1,4-linked galacturonic acid residues and 1,2-linked
rhamnose residues partially substituted for by single
galactose residues linked to rhamnose residues by 1,4-
linkages. Also, the RG-I subunit can have long arabinan
and galactan side chains. Arabinose residues can be ter-
minal 1,3- and 1,3,5- linked residues. RG-I can also
contain either a xylogalacturonan unit in which single
xylopyranose residues are 1,3-linked to the backbone,
as it is in apple pectin [31], or an apiogalacturonan frag-
ment in which single or 1,3-linked

D

-apiose residues
are linked to the

D

-galacturonic acid of the core by 1,2-
and/or 1,3-linkages [1, 6].
A general model for apple, citrus, and beet pectins
has been suggested which has an alternating linear 1,4-


Vol. 35

No. 3

2009

CURRENT VIEWS ON PECTIN SUBSTANCES 271
Linear region Linear region
Branched region
Branched region

:

arabinan, galactan,
Smooth regions
Ramied (hairy) regions
Homogalacturonan
Weakly ramied region

GalA
RG-I
Rha
Gal Ara

Rhamnogalacturonan
Branched (hairy) regions

RG-I

Xyl
Fer

of homo-
galacturonan
of rhamno-
galacturonan
of rhamnogalacturonan

I (RG-I)

and arabinogalactan.
Weakly branched region:
xylogalacturonan

Fer
Xyl Xyl Gal
Ara
Ara
Ara
Ara
Gal Gal Gal Ara

Schematic structure of pectin polysaccharides. Fer, ferulie acid residue.

linked

-

D

-galacturonan chain and a branched region
containing most of the neutral monosaccharides [33].
Original pectins from these sources differ by molecular
mass (citrus > apple > beet) and content of rhamnose
(beet > apple > citrus). From apple, beet, and citrus pec-
tins, homogalacturonan with a polymerization degree
of 72120, 91108, and 114138 kDa, respectively,
were isolated. Sugar beet pectin was found to contain
ferulic acid (Fer) residues which are linked to the neu-
tral monosaccharides of side chains (mainly to

L

-ara-
binofuranose residues) by an ester bond [34].
The structure of a molecule of apple and beet pec-
tins, as well as related pectins from other vegetables
and fruits, can be represented by a scheme from [31].

RG-II

, a minor component of primary cell walls, is
distinguished by a very complex structure [1, 30, 35
37]. RG-II is resistant to

-1,4

-

endo

-polygalacturonase
and is isolated after the preliminary treatment of pectins
with this enzyme. Among the constituent sugar residues
of RG-II, the widely occurring monosaccharides such
as

D

-galacturonic acid,

L

-rhamnose,

D

-galactose,

L

-
arabinose,

D

-xylose,

D

-glucose,

L

-fucose,

D

-mannose,
and

D

-glucuronic acid have been identied [37,38].
However, along with these residues, rather unusual
monosaccharides have been found as follows:

D

-
apiose,

2-

-methyl-

L

-fucose, 2-

O

-methyl-

D

-xylose,
AceA, DHA, and KDO. AceA was rst found in pectin
substances as an unidentied sugar residue of RG-II
from cell walls of a suspension of a sycamore culture in
1978 [39] and identied in 1983 as an acidic branched
monosaccharide [40].
H
H
OH
OH
CH
2
OH
O
OH
H
3
C
OH
COOH HO
O
H, OH
O
OH
OH
CH
2
OH
OH
COOH
HO
O
OH
OH
COOH
HO
HOOC
AceA
KDO DHA
D-Api

272


Vol. 35

No. 3

2009

OVODOV

DHA was identied by combined gas-liquid
chromatography-mass spectrometry and proton reso-
nance spectroscopy in 1988 [41]. The information
about the presence of KDO in pectins was rst reported
in 1978 [42] and later entirely conrmed [41]. This was
the rst evidence of the presence of KDO in pectin sub-
stances of plants, in particular in RG-II of primary cell
walls. Earlier, KDO had been detected only in bacterial
lipopolysaccharides. It was proposed that the three
above mentioned acidic monosaccharides (AceA,
DHA, and KDO) are the necessary components of RG-
II and should be present in any plant.
RG-II was rst isolated from the cell walls of the
sycamore

Acer

pseudoplatanus

[39]. Also, RG-II was
found in cell walls of rice, Douglas spruce, onion, kiwi,
radish, roots of the falcate hares ear

Bupleurum

falca-
tum

, and leaves of

Arabidopsis

thaliana

(see [1]). It was
also isolated from the pulp of a sugar beet, the commer-
cial enzyme preparation Pectinol AC, the products of
processing of a grape vine as the major component of
red wine polysaccharides, as well as from the juices of
apple (

Malus

domestica

), carrot (

Daucus

carota

), and
tomato (

Solanum

licopersicum

) treated with enzyme
preparations. RG-II is present as a dimer (see below) in
pectin polysaccharides of black currant and blueberry
(in cell walls, juice, and fruit press residues). In RG-II
located near the plasma membrane, the galacturonic
acid residues of the main chain are not esteried by
methanol, whereas RG-II located in primary cell walls
contains a great number of methoxyl groups, as was
shown by immunochemical methods. Interestingly,
after the softening of fruits and vegetables during ripen-
ing, RG-II becomes a dominant polysaccharide in
apple, tomato, and carrot juices. Thus, RG-II that accu-
mulates in juices during the processing of fruits and
vegetables is an important pectin polysaccharide in the
industrial production of fruit juices [43].
RG-II is a comparatively small-sized polysaccha-
ride of extremely complex structure which has not yet
been completely established, although some variants
have been suggested [35].
It was found that RH-II contains boron, which forms
boratediol esters that cross-link RH-II molecules to
form a dimer [38, 43]. It was shown that borate esters
cross-linking two RG-II molecules are localized on

-1,3'

-linked apiose residues, which consequently play
an important role in the growth of plant tissues. The
dimer has acid-labile borate ester bonds which are
hydrolyzed as pH in the cell wall and decrease during
the auxin-induced cell growth [38].
It is known that RG-II is the only polysaccharide in
primary cell walls that has borate bridges which form a
dimer from RG-II [38, 4446]. This borate complex is
a part of a macromolecular pectin complex consisting
of homogalacturonan, RG-I, and RG-II; borate esters of
RG-II form molecules of cross-linked macromolecular
pectin [46]. However, it is not ruled out that homogalac-
turonan, RG-I, and RG-II are covalently linked to each
other without the participation of borate [29, 47], since
polysaccharides are not separated from each other by
gel-permeation chromatography.
Although RG-II is contained in the cell wall in
minor amounts, it plays a key role in its architectonics
[48]. The structure of RG-II is rather conservative for
different plant species [41], indicating the importance
of its biological functions. RG-II not only inuences
plant growth, but is also involved in the interactions of
plants with phytopathogens by affecting the specic
transport systems during the passage of amino acids
through membranes. Of importance for the realization
of the biological functions of RG-II are rather long
galacturonan regions in its macromolecule (of seven
and more galacturonic acid residues) and, in the opin-
ion of the authors of [49], the residues of such unusual
monosaccharides as KDO and apiose.

Commercial Pectins

Commercial pectins nd use in the food industry
owing to their gelling capacity. Pectins obtained by acid
extraction from vegetables and fruits (sugar beets,
apples, citrus fruits) are predominantly homogalactur-
onans and contain minor amounts of neutral side
chains.
It is just these pectins that form a group of com-
mercial pectins [50, 51]. According to European
rules, commercial pectins intended for food products
must contain more than 65% of GalA per dry weight
of a sample, and according to American pharmaco-
poeia, more than 74%. Homogalacturonans differ
from one another by substituents: GalA residues can
contain free carboxyls or methanol-esteried car-
boxyls. Also, at C2 and C4 positions, GalA residues
can be acetylated, as is the case in pectins from sugar
beet and potato tubers. The gelation properties of
O
O
CH
2
O
O
O
O
H
2
C
O
O
O
O
B


Vol. 35

No. 3

2009


pectins with a low DM (LM pectins) can be enhanced
by chemical amidation in methanol, which leads to
the occurrence of amide groups at position C6 of
GalA residues (LMA pectins).

The physical properties of commercial pectins
depend on their molecular masses, and primarily on
DM. DM is determined by the number of moles of
methanol per 100 moles of galacturonic acid [50]. The
modern methods of the direct quantitative analysis of
methanol involve gasliquid chromatography (GLC) or
HPLC. HPLC requires much less time compared with
the earlier methods, in particular ion-exchange chroma-
tography on DEAE cellulose [52]. A method for deter-
mining DM by Fourier transform IR spectroscopy has
been elaborated: the value of the absorption band of the
carboxyl at 1756 cm

1

correlates well with the mean
value of DM [53]. The method is rapid but requires the
maintenance of a strictly specied pH value and precise
calibration using the characterized pectins. Recently, a
method of quantitative analysis of methoxyl groups in
pectins by GLC on columns with a special attachment
has been developed [54]. The results of this assay fully
agree with the data of HPLC.
In addition, a method for the simultaneous determi-
nation of the content of methoxyl and acetyl groups in
pectin polysaccharides has been devised. The method
consists in that methanol and acetic acid that split out
after alkaline hydrolysis are separated by GLC in a
Chrompak PoraPlot Q capillary column and are
detected by mass spectrometry. By using this proce-
dure, methanol and acetic acid are readily determined
quantitatively in 1 mg of pectin [55].
A well-suited method for determining DM and the
homogeneity of pectin fractions is capillary electro-
phoresis [56]. It was used to determine the mean value
of DM of a pectin sample since there is a direct relation-
ship between the electrophoretic mobility of the mole-
cule and the mean value of the charge per one monosac-
charide residue [57]. Subsequently, this method was
used to study the distribution of galacturonic acid
methyl esters in pectins [5880]. Samples were frac-
tionated by gel chromatography according to molecular
masses and then by electrophoresis according to
charge. The results conrmed the assumption that elec-
trophoretic mobility depends on molecular mass and
DM and showed that capillary electrophoresis provides
real information regarding the homogeneity and charge
density distribution in the pectin chain [56].
Pectins with a high DM (HM pectins) contain 50%
or more esteried GalA residues. LM pectins are
obtained by deesterication of HM pectins under par-
ticular controlled conditions (pH, temperature, and
time). Both groups of pectin polysaccharides form gels,
but under different conditions: LM pectins at low pH
values or in the presence of calcium cation, and HM
pectins form gels by hydrophobic interactions, espe-
cially in the presence of sucrose [51].
As a rule, commercial pectins are obtained using cit-
rus fruit peels and apple wastes, byproducts of the man-
ufacture of juices, and sucrose from sugar beet, and, to
a lesser degree, byproducts of the manufacture of starch
from potatoes, sunower heads in oil production, and
onions.
The pectin extraction procedure should be rapid to
prevent the degradation of polysaccharides by enzymes
in the starting materials. The degradation of pectins by
enzymes during the storage of the initial materials can
lead to samples with quite different gel forming proper-
ties. The starting material should be dried immediately
after processing to avoid degradation [51].
Pectins from the byproducts of sugar beet process-
ing have a low gelation capacity owing to their low
molecular masses and a high content of acetyl groups.
Treatment with acidic methanol removes acetyl groups
and increases DM but substantially lowers the molecu-
lar mass [61]. Nevertheless, acetylated pectins nd
application owing to their emulsifying properties. The
gelation of acetylated pectins can be enhanced by treat-
ment with pectinesterase and pectinacetylesterase [62].
As mentioned above, sugar beet pectin contains fer-
ulic acid residues [63] linked by ester bonds primarily
to O2 residues of arabinose and O6 residues of galac-
tose of neutral carbohydrate chains. The side chains of
arabinan and galactan consist of

1,5

-linked

-

L

-arabi-
nose and

1,4

-linked

-

D

-galactose residues, respec-
tively. When treated with a mixture of

H

2

O/peroxidase
or ammonium persulfate, these molecules are easily
joined to form dimers, thereby increasing the viscosity
and gelling of sugar beet pectin. About 20% of the total
number of ferulic acid residues in pectin are present as
dimers. The structural identication of dimeric oli-
gosaccharides from sugar beet pectin, linked by ferulic
acid residues, showed the presence of covalent (intra-
O
OH
OH
COOCH
3
O O
O
OH
OH
COOH
O
O
OH
OH
CONH
2
O

274


Vol. 35

No. 3 2009
OVODOV
and intermolecular) cross-links of pectin arabinans and
galactans through diferuloyl bridges in sugar beet cell
walls [63].
Owing to valuable physical properties, commercial
pectins are widely used as thickening agents, gels,
adhesives, emulsiers, and stabilizers of solutions.
Sources of Pectins Investigated in the Last Century
(19982008)
Pectin polysaccharides of the European North of
Russia, which were studied in the Department of
Molecular Immunology and Biotechnology of the Insti-
tute of Physiology (Komi Science Centre, The Urals
Branch, Russian Academy of Sciences), such as
silenan, a pectin from the catchy Silene vulgaris
(Moench) Garke (Oberna behen L.), tanacetan, a pectin
from the tansy Tanacetum vulgare L., lemnan, a pectin
from the lesser duckweed Lemna minor L., comaruman
from the marsh cinquefoil C. palustre L., and some oth-
ers have been described in great detail in review [6]. We
will not dwell on these compounds.
Plants of the genus Lupinus can serve as sources of
valuable pectins. It has been established that they con-
tain from 1.5 to 7% of pectic substances depending on
the variety, year, and vegetation stage. Varieties con-
taining about 7% of pectins are promising for industrial
production of pectins. Lupin pectins have perfect swell-
ing and water-absorbing abilities and form a stable gel
at a 2% concentration [64]. The molecular mass of pec-
tins varies between 100 and 400 kDa, and DM is 65
85%. The basis of a pectin macromolecule is D-galac-
turonic acid; of other monosaccharides, galactose and
arabinose have been identied as the main components.
Pectin has a complex branched structure and represents
a mixture of linear galacturonans and highly-branched
pectin polysaccharides [65].
It was proposed to use mandarin orange peels to
expand the spectrum of pectin sources. The composi-
tion and properties of pectins from mandarin orange
peels depend on the conditions of growth and the vari-
ety of the fruits [66].
It was also proposed to use burdock (genus Arc-
tium) as plant row material for obtaining pectin. Bur-
dock is characterized by a high content of pectic sub-
stances [67]. The highest content of pectin is in leaf cut-
tings (1.5%). In dry press residues of burdock, 21% of
pectic substances accumulate, whereas in dry apple
marcs, only 11%. Consequently, burdock leaf cuttings
can compete with traditional row material for obtaining
pectin. Unfortunately, burdock pectin gelatinizes
poorly, which is related to a low DM.
Pumpkin pectin was isolated by extraction with
0.1 M HCl after preliminary digestion using three
enzyme preparations: cellulase from Trichoderma vir-
ide, hemicellulase from Aspergillus niger, and a gly-
cosidase complex from Xanthomonas campestris [68].
The method was optimized as follows: pectin was
obtained from the pumpkin pulp preliminarily treated
with an enzyme preparation from Asp. awamori [69].
The main function of the enzymes is to digest cellulose
and other insoluble components of plant tissue. The
treatment with puried enzymes of a known specicity
enables one not only to isolate pectin but also to obtain
additional information about its structure and links with
other cell wall components.
It should be noted that pumpkin is a nontraditional
but a very interesting and promising source of pectin.
Pumpkin pectin forms gels at concentrations much
lower than the commercial citrus pectin. The use of
enzymes substantially increases the yields of pectin
substances and leads to the complete solubilization of
plant material.
Pectins of amaranth were isolated from different
species of this widely distributed plant [70]. On the ter-
ritory of Russia, 17 amaranth species exist. Amaranthus
retroexus L. or prostrate amaranth, known as a plant,
occurs most widely. Many amaranth species nd appli-
cation in nontraditional medicine. Thus, an aqueous
infusion of leaves of this plant is used in the treatment
of diseases of the GIT and as a hemostatic. A decoction
of the apices of Am. cruentos is used for the treatment
of rheumatism, womens diseases, and as an antitussic.
Amaranth decoctions are used to remove radionuclides.
Several methods of isolating pectin from amaranth have
been developed. Good results with a high efciency of
isolation have been obtained by the method using a
rotor-pulsation device. The raw material is thoroughly
divided by the device and is subjected to cavitation and
ultrasound treatment, which results in the disintegra-
tion of the raw material and the highly effective extrac-
tion of pectin [71].
Amaranth pectin is characterized by a high content
of D-galacturonic acid residues (more than 80%) and a
high DM (75%). Structurally, this is a typical represen-
tative of this class of polymers: it has a linear region of
galacturonan (rhamnogalacturonan) and a ramied
region of RG-I in which the side chains are composed
primarily of arabinose and galactose residues [70].
Amaranth pectin is used in medicine as a hypoten-
sive drug, in nutrition, and as a regulator of growth and
development of plants [70].
A new pectin has been isolated from leaves of the
Thailand ligneous vine Cissampelos pareira (family
Menispermaceae) which widely occurs in warm
regions of Asia, East Africa, and South America [72].
The plant is considered to be a medicinal plant and is
used in the treatment of asthma, dysentery, urological
diseases, and post-traumatic pains. Aqueous extracts of
leaves form gels which are used as a dessert in Thai-
land. These gels can serve as a source for obtaining nat-
ural polysaccharides, in particular pectin. Pectin con-
sists predominantly of galacturonic acid residues (70
75%) with a low DM and minor amounts of neutral
monosaccharides. Pectin forms a good gel whose sta-
RUSSIAN JOURNAL OF BIOORGANIC CHEMISTRY Vol. 35 No. 3 2009
bility depends on the presence of Na
+
, Ca
2+
, and
sucrose.
From berries of the cherry Prunus dulcis, a bio-
logically active arabinan-rich pectin polysaccharide
with a molecular mass of 762 kDa has been isolated
[73]. The highly-branched arabinan moiety contains the
backbone of 1,5-linked arabinose residues; other link-
ages are in the ratio of 3 : 2 : 1 : 1 for t-Araf : 1,5-Araf :
1,3-Araf : 1,3,5-Araf, respectively, where t is the termi-
nal residues.
It was shown that the crude extract activates D and
T lymphocytes. Acidic and neutral fractions also stim-
ulate T lymphocytes.
Sunower pectin isolated from sunower heads
and stalks after the removal of seeds belongs to LM
pectins [74]. It is obtained with higher yields than pec-
tins from traditional sources such as citrus fruits and
apples. The esterication degree of the LM pectin is
11%, the content of galacturonic acid varies from 77 to
85%, and the content of acetyl groups is about 2.5%.
Aqueous solutions of pectin are rather viscous, and it
forms gels comparable in rigidity with commercial
gels.
From the iris Iris pseudoacorus, a polysaccharide
fraction has been isolated, which, according to the data
obtained, is pectin [75]. The sugar composition of the
pectin fraction is represented by galacturonic acid
(46.7%), galactose, and arabinose as predominant
monosaccharides. Also, rhamnose and xylose are
present in minor amounts. The polysaccharide is solu-
ble in water and forms viscous solutions.
From tubers of the potato Solanum tuberosum,
pectin RG-I has been isolated [76] which has been
shown to regulate the development of potato tubers
[77]. Side chains representing 1,4-linked -D-galactan
and 1,5-linked -L-arabinan are attached to rhamnose
residues at position 4 [47]. A further study of potato
pectin was performed using enzymatic treatment with
endo--1,4-galactanase and endo-1,5-arabinase or with
their combination. The resulting products were sepa-
rated and analyzed using monoclonal antibodies to pec-
tin polysaccharides. The analysis showed the presence
of a great number of 1,5--galactan side chains to some
of which short side chains containing arabinose were
coupled. Also, 1,5--L-arabinan side chains substituted
for by 1,4--D-galactan oligomers (polymerization
degree >4) and 1,5--L-arabinans resistant to enzy-
matic digestion were found [76].
From the pea Pisum sativum L., xylogalacturonan
has been isolated which was characterized using mon-
oclonal antibodies. It has been shown that the epitope to
monoclonal antibodies specic for xylogalacturonan
represents the region of xylogalacturonan highly
enriched with xylose [78].
Pectin polysaccharides from olives, particularly
those extracted by diluted alkali [79], contain a large
amount of arabinose. Samples obtained by extraction
with cold buffer solutions have a low degree of branch-
ing. These polysaccharides contain predominantly 1,5-
linked arabinose residues of which about 30% in the
arabinan backbone are substituted at position 3. The
amount of 1,3,5-linked arabinose residues increases to
70% if pectin polysaccharides are extracted using more
potent solvents (e.g., aqueous solution of ammonium
oxalate on heating to 6070).
Pectin polysaccharides from olives contain two
types of galactans [80]. A great portion of galactans
with 1,3,6-linked galactose residues is often associated
with hydroxyproline-containing proteins. However,
they may represent the side chains of RG-I or single
homopolysaccharides. About 2030% of galactose res-
idues are present as 1,4-linked galactan.
During ripening of olive fruits, the content of pec-
tins substantially increases, their sugar composition
changes, and the amount of methyl esters and acetyl
groups decreases. The molecular-weight distribution of
polysaccharides in this case changes little [79, 80].
The byproducts of olive oil manufacture serve as a
valuable source of pectic substances. An extract with a
high content of arabinans extracted from this raw mate-
rial was separated into two fractions, after which the
structure of the main arabinan ( 8.4 kDa) was deter-
mined by the methylation analysis and NMR spectros-
copy [81]. The backbone of its molecule consists of
1,5-linked -L-arabinofuranose residues. Single
-L-arabinofuranose residues are linked to the back-
bone by a 1,3-linkage. 1,5-Linked terminal -L-ara-
binofuranose residues and disaccharide units of 1,3-
linked -L-arabinofuranose, which are linked at posi-
tion 3 of the backbone by -linkages, are present in
minor amounts. The pectin polysaccharide from the
fruits of the olive Olea europea has this structure [79,
82]; its content reaches 40%, and its backbone is com-
posed of units of highly-branched RG-I enriched with
arabinose.
Soy bean pectins are represented by two groups
[83]. The rst group includes pectins with 1,4-linked
galacturonic acid residues as the backbone. Among
these are homogalacturonan, RG-II, and xylogalactur-
onan.
The second group involves pectins with the back-
bone of RG-I and side chains of arabinan, galactan, or
arabinogalactan. The ratio of rhamnose to galacturonic
acid in this group of pectins varies between 0.05 : 1 and
1.0 : 1.0.
Diospyros kaki, a Chinese plant of the Ebenaceae
family, is widely distributed in the tropics and subtrop-
ics. The plant is used in Chinese traditional medicine
mainly in the treatment of hiccups. Pectin has been iso-
lated from leaves of the plant by extraction with hot
water followed by chromatography on DEAE cellulose
and gel ltration on Sephacryl S-300 [84]. Pectin rep-
resents a heteroglycan with the backbone consisting of
repeating disaccharide residues [ 4)--GalAp-
(1 2)--Rhap-1 ] with a 58.7% substitution at
position O4 of rhamnopyranose residues by -1,4-
276
OVODOV
linked xylopyranose and -1,3- and -1,6-linked galac-
topyranose residues (galactan). In addition, the side
chains contain arabinofuranose residues of xylose resi-
dues linked at position 2 by a -1,4-linkage and of galac-
tose residues linked at position 3 by a -1,6-linkage.
Pectin stimulates the proliferation of B lymphocytes
induced by a lipopolysaccharide but does not stimulate
the proliferation of T lymphocytes induced by con-
canavalin A. It was shown that arabinose residues in the
side chains are not necessary for immune activity. Of
primary importance is the backbone of galacturonan.
Centella asiatica is used in China and India as a sed-
ative and an antiulcerous preparation, as well as in the
treatment of lepsory. Pectin isolated from this plant
contains Rha, Ara, Gal, Glc, and GalA in the molar
ratio of 1.0 : 0.6 : 1.5 : 0.2 : 1.1 [85]. It was found that
the backbone of this pectin consists of the repeating dis-
accharide unit 4)--D-GalpA(1 2)--L-Rhap-
1 and hence belongs to the RG-I group. The side
chains are predominantly linked to the rhamnopyranose
residues of the core by a 1,4-linkage. These chains con-
sist of residues of arabinose and galactose, or arabinose
residues in combination with galactose residues; also,
short glucan chains are present. About 45% of Rhap res-
idues in the core are substituted for by side chains. Ara-
binose residues form for the most part oligoarabinosyl
side chains and are occasionally attached to 1,6-linked
galactose residues.
Pectin possesses no immune activity. However,
some of its derivatives, e.g., those devoid of acetyl
groups and containing a lesser amount of arabinose res-
idues, show immunostimulating activity. Treatment
with periodate and the removal of 1,6-linked Gal resi-
dues also lead to a derivative with immunostimulating
activity. Derivatives containing the core with a minor
amount of side chains have the highest activity.
Some species of cactus (family Cactaceae) nd
practical application [8689]. Thus, the opuntia Opun-
tia cus indica Mill is cultivated in Chile as a fruit
plant, and young shoots of this cactus are also used in
some countries by humans as a food product. The pec-
tin polysaccharide obtained from the fruit peel has the
main homogalacturonan chain of -1,4-linked galactu-
ronic acid residues; some regions of homogalacturonan
are joined by 1,2-linked -L-rhamnose. The side chains
are composed of 1,5--L-arabinan and 1,4--D-galac-
tan. Also, the endosperm of opuntia seeds was found to
contain pectins rich in arabinan.
From the ginseng Panax ginseng C.A. Mey (family
Araliaceae), a series of pectin polysaccharides have
been isolated and thoroughly studied by Japanese
investigators [90]. The structures and physiological
activity of some of these polysaccharides have been
considered in detail in a recently published review [91].
Most recently [92], one more acidic pectin-like
polysaccharide has been isolated from ginseng. It has a
high antiadhesive activity against pathogenic bacteria,
in particular against the adhesion by epithelial cells of
the GIT of the bacterium Helicobacter pylori, one of the
main ethiological agents of chronic gastritis, ulcers of
stomach and duodenum, stomach carcinoma, and lym-
phoid tumors. Pectin polysaccharide has a molecular
mass of 12 kDa and consists of residues of galacturonic
and glucuronic acids, rhamnose, arabinose, and galac-
tose. A partial hydrolysis of the polysaccharide by pec-
tinase furnishes an oligosaccharide fraction whose
activity is comparable with the activity of the original
biopolymer.
Hares ear Bupleurum falcatum (family Umbel-
liferae) is a medicinal plant widely distributed in Japan,
China, and Primorye Territory. Its roots are tradition-
ally used in the treatment of chronic hepatitis, nephro-
logical syndrome, and various autoimmune diseases.
The physiologically active pectin polysaccharide from
this plant, named bupleuran 2IIc, has been studied in
detail by a group of Japanese investigators headed by
Prof. Yamada [90, 93]. They have established the struc-
tures of bupleuran 2IIc and its antigenic epitope. Pectin
contains approximately 70% of 1,4-linked -D-galacto-
pyranosyluronic acid, of which 30% is esteried by
methanol. Some galacturonic acid residues are the
points of branching of the carbohydrate chain of the
pectin polysaccharide. Also, bupleuran 2IIc has a typi-
cal ramied region of RG-I. The polysaccharide also
contains a minor region of RG-II which contains the
same rarely-occurring monosaccharide residues:
apiose, AceA, KDO, and DHA. It was shown that
bupleuran 2IIc has high antiulcerous and anticomple-
mentary activities. Intravenous injection of pectin
results in the formation of antibodies, and the polysac-
charide is found in the liver. After peroral administra-
tion, the polysaccharide is also detected in the liver and
Peyers patches. It was found that the antigenic epitope
of antiulcerous bupleuran 2IIc is located in the external
moiety of the ramied region and represents two differ-
ent oligosaccharides with the terminal side residues of
glucuronic or 4-O-methylglucuronic acids, which are
linked to the galactan chain by a 1,6-linkage [94]:
Also, bupleuran 2IIc has a mitogenic action: it pro-
liferates B cells in the absence of macrophages, and
activated B cells are transformed to antibody-forming
cells in the presence of interleukin IL-6. Pectin also
enhances the secretion of macroglobulin IgM from nor-
mal murine B cells. As a result, long-term cell prolifer-
ation is induced. When B cells are stimulated by
bupleuran 2IIc, the protein of retinoblastoma is phos-
phorylated and the expression of cyclins regulating the
cell cycle is enhanced, which correlates with cell prolif-
eration. The ramied region of pectin enhances the
secreting activity of IL-6, thereby causing an increase
in macroglobulin secretion [95].
3Galp1 3Galp1 3Galp1
6
GlcpA1
or
4MeGlcpA1
Structure of the epitope
of bupleuran 2IIc
From leaves of the plantain Plantago major L.
(family Plantaginaceae), two polysaccharides show-
ing a high activity toward the complement have been
isolated: arabinogalactan RMI and pectin polysaccha-
ride PMII [96]. The rst polysaccharide belongs to ara-
binogalactans of AG-II type. It consists of the 1,3-
linked galactan core with branching 1,6-linked galactan
side chains coupled to the backbone by a 1,6-linkage.
Terminal and 1,5-linked arabinofuranose residues are
linked to these side chains via a 1,3-linkage. Also, PMI
contains a protein component characteristic for AG-II.
The pectin polysaccharide PMII has a typical structure
of RG-I [91].
A study of the interaction between the human com-
plement and PMII has shown that PMII is a potent acti-
vator of both the classical and alternative pathway of
complement activation. It induces the secretion of
human immunoglobulin IgG, which is directly related
to the well-known wound-healing effect of leaves of the
greater plantain P. major [91, 97].
It has been shown on mice that regular prophylactic
administration of PMII protects well against pneumo-
coccal infection, and this protective effect is due to the
stimulation of inborn rather than adoptive immunity
[91, 98100].
Argane tree (Argania spinosa L.), family Sapota-
ceae, is an endemic of south-west Morocco [101, 102],
has a hard wood pulp, and gives fruits similar to olives.
It protects soil against water and wind erosion. Differ-
ent parts of the plant nd wide and various applications:
seeds are used to obtain valuable oil for nutrition, cos-
metic, and medicinal purposes (they lower the choles-
terol level and improve blood ow); substances being
part of cell wall polymers are used in food and cosmetic
industries and as drugs.
Along with cellulose, cell walls contain pectins, an
arabinogalactanprotein complex, and xyloglucan.
In the fraction of pectin substances from fruit pulp,
homogalacturonan, RG-I, and RG-II have been isolated
and fractionated. RG-I is the main constituent of pectin
and contains, along with galacturonic acid, a great
amount of arabinose residues and a lesser amount of
galactose residues as branched side chains. The resi-
dues of other monosaccharides: glucose, xylose,
apiose, and glucuronic acid are the minor components
of RG-I. Arabinans and/or arabinogalactans contained
in the pectin fraction are present either in a free state or
as part of RG-I side chains. The presence of galactan
side chains is also not excluded. RG-II is present in the
pectin fraction as a minor polysaccharide in the form of
a dimer. Arabinose residues amount to 34% of all
monosaccharides of this polysaccharide, which consid-
erably exceeds the content of arabinose in RG-II of the
majority of other plants [102]. For comparison, the con-
tent of arabinose in RG-II of Arabidopsis thaliana is
14% [103], and in RG-II of pumpkin, as low as 9%
[104].
Peach (Persica vulgaris L.), family Rosaceae, is
grown as a fruit culture in countries of moderately
warm and subtropical zones. In China, six species of
this plant grow. Many varieties are cultivated in Middle
Asia, in the Caucasus, in southern Ukraine, and in
Moldova. Despite the wide distribution of the peach,
pectin substances from its fruits have been little inves-
tigated. There are only a few articles on the extraction
of pectin from peach fruits [105108]. Temperature,
pH, and the time of acid treatment affect the yield of
pectin from fresh and stored peach grist. The yield of
pectin varies from 2 to 10% depending on pH and tem-
perature. The yield is maximal at a higher temperature
and low pH values. The DM value in all cases remains
high and ranges from 72.1 to 95.4%. It was also found
that the yield of pectin from stored press residues is
substantially higher than that from fresh husks; how-
ever, the DM and specic viscosity of solutions in this
case decrease.
The yield of pectin substances also depends on the
ratio of ethanol and extract during precipitation of pec-
tin. As this ratio increases, a higher yield and greater
DM values are observed. A maximal yield (9.94 2%)
was obtained at the ethanolextract ratio of 1.5 : 1.0 and
with the time of acid treatment of about 120 min [108].
Pectin from spinach (Spinaca oleraceae L.), fam-
ily Chenopodiaceae, has been studied by treating the
cell walls of a suspension culture with the enzyme
driselase [109] which easily hydrolyzes glycosyl bonds
of -D-GalpA and -L-Rhap in homogalacturonan and
RG-I after the preliminary removal of methyl ester
groups by cold diluted alkali. Driselase is also capable
of partially and completely hydrolyzing methylesteri-
ed oligogalacturonides to form free GalA and metha-
nol. However, other esters (acetates and feruloyl ester
groups) are resistant to driselase; as a result, oligosac-
charides carrying -acetyl or -feruloyl ester groups
can result during fermentolysis. In addition, the pres-
ence of -acetyl groups protects one or several methyl
ester groups from hydrolysis by driselase. This makes
it possible to map methyl ester and acetyl ester groups
inside the pectin core. By enzymatic hydrolysis of spin-
ach pectin with driselase, oligogalacturonides and
rhamnogalacturonides were isolated. Five out of six oli-
gosaccharides from homogalacturonan contained 3--
acetyl-GalpA residues, whereas methylesteried GalA
residues were located near both 2--acetyl-GalA and
3--acetyl-GalA. The oligosaccharide GalA-Rha-
GalA-Rha-GalA obtained from RG-I also contained
3--acetyl-GalA residues.
Thus, the distribution of methyl ester and acetyl
groups in galacturonan and RG-I of spinach pectin is
irregular and is still far from being completely under-
stood [105].
278
OVODOV
Physical Properties of Pectins
The gelling properties of pectin polysaccharides
depend mainly on DM [33, 110, 111].
Pectins are widely used as gelling agents, stabilizers
of solutions, thickening agents, adhesives, and emulsi-
ers in many food products. It has been shown that an
important role in the formation of pectin gels is played
by sucrose, which can stabilize the structure of junction
zones of gel formation [1, 112].
The interaction of proteins and peptides with pectins
results in the formation of soluble and insoluble com-
plexes whose structures depend on the ratio of biopoly-
mers and pH [113115]. In this process, covalent bonds
between the tyrosine residues of the protein and the car-
boxyl group of uronic acid of pectin are formed, which
can lead to the formation of a pectin network. There is
an alternative hypothesis, according to which proteins
form ionic complexes with pectins [114, 116].
Globular proteins, such as bovine serum albumin
(BSA) and -lactoglobulin, form gels, and the mixing
of these gels with LM pectin results in the formation of
mixed gels. During the formation of gels, a kinetic
competition between gel formation and the phase sepa-
ration process takes place. The control of relative rates
of these processes by external factors, such as environ-
mental conditions, leads to a wide spectrum of micro-
structures [116, 117]. The formation of gels during the
interaction of pectin polysaccharides and globular pro-
teins has been extensively studied. A variety of struc-
tures and the rheological properties of resulting gels
have been described which differ depending on the
nature of biopolymers and the conditions of gel forma-
tion (see, e.g., [117125]). It has been found that the
presence of Na
+
and Ca
2+
cations and their ratio play a
decisive role in the gel formation of mixtures of globu-
lar proteins with pectins [117]. Increasing the amount
of pectin and calcium concentration makes the mixed
gel more rigid [119].
During the treatment and storage of gels, reactions
between pectin and a protein or a peptide can take
place: hydrolysis of ester groups in pectin, decarboxy-
lation, -elimination, or the hydrolysis of glycoside
bonds due to the presence of pectinase in the mixture.
These processes should be taken into consideration
during the production of pectinprotein gels [126].
The high bioadhesive capacity of pectins was used
to obtain a medicinal preparation based on lactoferrin,
a milk glycoprotein producing the bactericide action in
the treatment of chronic inammation in stomatitis
[127]. It was shown that HM pectins are best suited for
preparing bioadhesive tablets since, along with a high
bioadhesive capacity, they have the property of releas-
ing the active medicinal substance. Moreover, this pro-
cess can be regulated by the concentration of calcium
ions, which cross-react with the galacturonan fragment
of the pectin macromolecule and facilitate the release
of lactoferrin.
It is well known that the gel-forming capacity of
pectins depends on many factors, including DM, the
localization of methyl ester groups, pH, the concentra-
tion of biopolymer and calcium ions, the ionic strength
of solution, temperature, and others [72, 110, 128
130].
LM pectins form gels in the presence of calcium
ions. Commercial LMA pectins require a lesser amount
of calcium for gelation and are less sensitive to synere-
sis induced by a high concentration of calcium ions; in
this case, the gel formation is thermally reversible
[131].
The range of calcium ion concentrations at which
rigid gels are formed is relatively narrow in the case
that gel formation occurs in the absence of other salts or
with the use of NaCl alone. This range is extended
towards higher Ca
2+
concentrations in the presence of
sodium citrate and potassium and sodium tartrates, as it
has been shown in a study of the effect of this mixture
of salts on the gelation of LMA pectin [128]. These
salts make gels more elastic and thermally reversible,
even during mechanical destruction. The treatment of
HM pectins with pectinmethylesterase leads to the for-
mation of LM pectins whose behavior in solutions
greatly differs from the behavior of HM pectins and
strongly depends on the concentration of monovalent
cations [129, 132]. When solved in the presence of
monovalent ions of Li
+
, Na
+
, and K
+
at concentrations
from 0.005 to 0.05 M, the resulting LM pectin exhibits
a higher specic viscosity than HM pectin, whereas
HM pectin has a higher specic viscosity in the pres-
ence of 0.2 M monovalent salts. Interestingly, M
w
sub-
stantially increases during the transformation of HM
pectin to LM pectin, which is probably related to the
aggregation of molecules of LM pectin being formed
[129, 132].
Mixtures of HM and LM pectins are often used in
the food industry for the manufacture of products with
particular properties. By varying the conditions of gel
formation, it is possible to substantially change the
rheological behavior and the microstructure of mixed
gels of HM and LM pectins [130, 133, 134]. A strong
synergetic effect in the rheological behavior of mixed
gels was noted in the presence of Ca
2+
and 60% sucrose
at pH 3, i.e., under conditions promoting the gelation of
both HM and LM pectins.
Also, during the production of gels with a low
sucrose content, a strong synergetic effect is observed
in the rheological behavior of mixed gels in which the
concentration of HM pectin is three times as high as the
concentration of LM pectin. In this case, a mixed gel
with a lower sucrose concentration has the same rheo-
logical parameters as HM pectin in the presence of
sucrose of a considerably higher concentration [130].
Interesting results have been obtained in studies of
the gelling properties of natural citrus pectin (DM
64%). Pectin was preliminarily either esteried by
methanol to pectin with a higher DM or saponied to
obtain pectin with a low DM value, and the resulting
samples were then reduced by sodium borohydride
[110]. As a result, two thirds of galacturonan carboxyl
groups were transformed to primarily alcohol groups,
and only one third remained in the form of free car-
boxyl groups. Eventually, the resulting samples gave
solutions with a substantially higher dynamic viscosity
and better rheological parameters. It has been shown
earlier that the gelation and subsequent stability of gels
are determined by hydrogen bonds and hydrophobic
interactions, in particular between the hydroxyl groups
of galacturonan and sucrose and between the methyl
ester groups, respectively [135]. In the absence of
sucrose, these bonds are weak and gelation does not
occur. In the case of citrus pectin modied by saponi-
cation, the number of hydroxyl groups increases, which
leads to the formation of a considerably greater amount
of hydrogen bonds. Therefore, the stabilization of the
remaining hydrophobic interactions and the formation
of gels require a lesser amount of sucrose than in the
case of the original citrus pectin [110].
Gel matrices based on biopolymers, among them
pectin substances, are widely used in food, agricultural,
and pharmaceutical industries as systems for the encap-
sulation and subsequent release of active substances.
One of the trends in the food industry is the release of
aromatic compounds from colloid solutions and gels
[136138]. It has been shown that the intensity of the
odor emitted by colloid solutions and gels weakens
with increasing the concentration and rigidity of the gel
owing to the direct binding of aromatic substances with
polymeric chains [136, 138140]. Gels of citrus HM
pectin at different concentrations for the limonene odor
conservation have been studied, and a correlation
between a decrease in the intensity of the emitted odor
and an increase in gel rigidity has been established
[138].
At present, in the eld of biopolymer research it is
proposed to use biopolyelecrtolyte multilayers for the
creation of encapsulation systems and the preparation
of coatings which enable one to control cell adhesion
[141, 142]. Thus, the formation of multilayers during
the interaction of citrus pectin (DM 36.6%) with the
weak oppositely-charged polyelectrolytes poly-L-
lysine and chitosan has been studied [143145]. Poly-
L-lysine can cross-link molecular networks of pectin
and bind with a high afnity to the surface of a pectin
macromolecule. Pectin and chitosan in the ratio 1.2 : 1
at pH 5.6 form on solid surface alternating layers of
pectin and chitosan; during this process, the binding of
the polymers to the surface takes place. The multilayer
formation is observed at pH values at which both poly-
mers carry a charge on macromolecules. The thickness
of an individual layer depends on the concentration of
biopolymer. Changes in pH that suppress the charge of
one of the polyelectrolytes lead to the destruction of the
multilayer composition, indicating the importance of
electrostatic interactions for the formation and stability
of multilayers [145].
Effect of Pectins on the Growth and Development
of Plants
Pectins fulll important biological functions in
plants [1, 29]. They regulate ion and water exchange,
participate in the structure formation of plant cell walls,
facilitate seed germination and growth of plantlets, pro-
vide plant turgor, etc. Changes in the composition and
structure of pectin substances during the growth and
development of plants have been studied by an example
of pectins from stalks and leaves of the common ax
Linum usitatissimum L. [146]. The content of pectins in
stalks of the plant decreases from 7% in the initial stage
of development to 2% in the stage of wilting, whereas
their content in leaves during the development remains
unchanged (~4%). The content of galacturonic acid in
pectin polysaccharides of the stalk increases from 53 to
66% in the wilting phase, whereas in pectins of leaves
only insignicant changes during plant growth are
observed. The DM of ax pectin polysaccharides is
high, 80100%, and varies slightly during the develop-
ment of the plant. The degree of acetylation plays an
important role in the development of the plant and is
equal to 2340%; it only slightly changes during plant
growth and development. Pectin substances have high
molecular masses and are synthesized at the initial
stage of plant development [146]. The content of solu-
ble -galactans localized in ax stalks falls during the
development of the plant: they are either destroyed or
are xed inside cell walls [147].
It has been shown that during ripening of fruits, the
polygalacturonase-induced solubilization of pectin
polysaccharides and the release of ethylene take place,
which enhances fruit ripening. These processes proceed
differently in different fruits [148153]. Thus, the rate
of ethylene release by the fruits of the Peruvian cherry
(ground cherry Physalis peruviana L.) increases during
ripening and is maximal on day 50 after owering,
which is accompanied by the synchronous release of
CO
2
, and the high level of the polygalacturonase activ-
ity correlates well with an increase in the amount of
released ethylene [152]. It should be noted that fruit
softening is always associated with a decrease in the
amount of the insoluble form of pectic substances,
which is controlled by specic enzymes, primarily by
polygalacturonase [152, 153].
Physiological Activity of Pectins
Pectins have a wide spectrum of physiological activ-
ity [133, 154], including the immunomodulating action
[1, 155]. The gastroprotective action of pectins is also
well known [1, 156, 157].
Pectins have been an integral part of food at all
stages of the evolutionary development of humans,
which has governed the practically ideal adaptation of
the human organisms to them [154].
A property of great importance is the anticarcino-
genic and/or antimetastatic effects of pectins, in partic-
280
OVODOV
ular apple pectin, which have been demonstrated by the
example of apple pectin in experimental models of
intestinal carcinogenesis and liver metastases. The
application of apple pectin in the treatment of intestinal
cancer has been patented in Japan.
It has been shown that pectins with different esteri-
cation degrees, as well as polysaccharides from these
pectins, induce the apoptosis of adenocarcinoma HT29
of the human large intestine cancer in vitro. The great-
est effect has been observed with oligosaccharide prep-
arations from LM pectins [158].
It is assumed that on the surface of cancer cells, pec-
tin polysaccharides bind proteins responsible for the
adhesion of tumor cells by healthy tissues. A commer-
cial preparation which represents modied citrus pectin
suppresses the growth of cancer cells and metastasizing
[159, 160]. It has been shown that it also increases the
prostate-specic antigen doubling time, the major
oncomarker of prostate adenocarcinoma, in 70% of
men suffering from prostate cancer [161].
Also, modied citrus pectin facilitates the removal
of toxic elements out of the organism with urea [160].
The use of pectin as a carrier matrix of biologically
active components or drugs merits attention [162, 163].
Among these are the products of interaction of citrus
pectin with helminthicides. Studying the immobiliza-
tion of the antituberculous drug isoniasid on pectin sub-
stances has shown that the preparation obtained has a
higher tuberculostatic activity than pure isoniasid. The
application of chemically modied pectins as carriers
of drugs also is promising for the treatment of the intes-
tine. Cross-linked pectin, which is less soluble and less
subject to degradation in the organism, as well as cal-
cium pectinate gel [162, 163] containing NaHCO
3
and
3
are effective for these purposes. These gels are
characterized by a high porosity and proper density.
The acidication of the gelling medium increases the
pore sizes, which provides a good and quick release of
the medicinal preparation in the proper place. On the
other hand, the method makes it possible to regulate the
realization of the drug and provide its prolonged release
[163].
Pectins attract increasing interest in gastroenterol-
ogy as peroral preparations and therapeutical means in
the treatment of diseases associated with mucosal
inammation. Also, pectins are used as a physical bar-
rier to protect epithelium against opportunistic micro-
organisms during stress [164167]. Among different
aspects of the action of pectins on the GIT, its interac-
tion with mucus is the subject of most intensive study;
thus, it has been shown that pectins bind to mucin, the
main component of the mucus of the GIT, to form a gel
network [167]. In this manner, pectin polysaccharides
enhance the protective barrier properties of mucus and
hence can be applied to treat lesions of the GIT and
infectious diseases. By varying the molecular charac-
teristics of pectin, it is possible to change the rigidity of
the resulting gels and the pattern of pectin distribution
in the pectinmucin complex. This enables one to reg-
ulate the interaction of pectin with mucin to meet the
demands for drug delivery and clinical therapy [168].
Conclusions
Pectic substances represent a manifold class of com-
plex plant polysaccharides which constitute a function-
ally important moiety of primary cell walls. At present,
great advances of fundamental importance have been
made, especially in the elucidation of the main features
of the chemical structure, physicochemical properties,
and biological activity of pectins. Nevertheless, many
problems are yet to be studied to use this big class of
natural compounds purposefully and with awareness
for treating various diseases and increasing the duration
of active human life.
ACKNOWLEDGMENTS
This work was supported by the program: Leading
Scientic Schools (NSh-2383.2008.4), the Russian
Foundation for Basic Research (project no. 06-04-
48079), the integration project of basic research of the
Urals and the Siberian Divisions of the Russian Acad-
emy of Sciences, the Programs of the Presidium of the
Russian Academy of Sciences: Fundamental Sciences
to Medicine and Molecular and Cellular Biology,
and the Foundation for the Promotion of National Sci-
ence.
The author is grateful to the senior engineer of the
Institute of Physiology (Komi Science Centre, The
Urals Branch, Russian Academy of Sciences)
Yu.A. Ovchinnikova for help in the preparation of the
review.
REFERENCES
1. Ovodov, Yu.S., Russian J. Bioorgan. Chem., 1998,
vol. 24, pp. 423439.
2. Kertesz, Z.I., The Pectic Substances, New York: Inter-
sci. Publ., 1951.
3. Hirst, E.L. and Jones, J.K.N., Adv. Carbohydr. Chem.,
1946, vol. 2, pp. 235271.
4. Aspinall, G.O., Adv. Carbohydr. Chem., 1959, vol. 14,
pp. 429468.
5. Ovodov, Yu.S., Pure Appl. Chem., 1975, vol. 42,
pp. 351369.
6. Ovodov, Yu.S., Sever: nauka i perspektivy innovatsion-
nogo razvitiya, (The North: Science and Prospects of
Innovative Development), Lazhentsev, V.N., Ed., Syk-
tyvkar: Izd. Komi Nauch. Tsentr. Ural. Otd. Ros. Akad.
Nauk, 2006, pp. 236255.
7. Henglein, F.A., Handbuch der Panzenphysiologie,
1958, vol. 1, pp. 405464.
8. Joslyn, M.A., Adv. Food Res., 1962, no. 11, pp. 125.
9. Endress, H.-U, in The Chemistry and Technology of
Pectin, Walter, B.H., Ed., San Diego: Academic Press,
1991, pp. 251268.
10. Voragen, A.G.J, Pilnik, W, Thibault, J.-F, Axelos, M.A.V,
and Renard, C.M.C.G, in Food Polysaccharides and their
Applications, Stephen, A.M., Ed., New York: Marcell Dek-
ker, 1995, pp. 287339.
11. Behall, K and Reiser, S, in Chemistry and Function of
Pectins, Fishman, M.L. and Jan, J.J., Eds., Washington,
DC: Am. Chem. Soc., 1991, pp. 248265.
12. Schols, H.A and Voragen, A.G.J, in Pectins and their
Manipulatio, Seymour, G.B. and Knox, J.P., Eds.,
Oxford: Blackwell Publ., 2002, pp. 129.
13. Round, A.N., MacDougal A.J., Ring S.G., Morris V.J,
Carbohydr. Res., 1997, vol. 303, pp. 251253.
14. Round, A.N. and Rigby, N.M., MacDougal A.J.,
Ring S.G., Morris V.J, Carbohydr. Res., 2001, vol. 331,
pp. 337342.
15. Mimmo, T., Marzadori, C., Montecchio, D., and
Gessa, C., Carbohydr. Res., 2005, vol. 340, pp. 2510
2519.
16. Cozzolino, R., Malvagna, P., Spina, F., Giori, A., Fuzza-
ti, N., Anelli, A., Garozzo, D., and Impallomeni, G.,
17. Ovodova, R.G., Popov, S.V., Bushneva, O.A.,
Golovchenko, V.V., Chizhov, A.O., Klinov, D.V., and
Ovodov, Yu.S., Biochemistry (Moscow), 2006, vol. 71,
pp. 538542.
18. Zandleven, J., Beldman, G., Bosveld, M., Schols, H.A.,
and Voragen, A.G.J., Carbohydr. Polym., 2006, vol. 65,
pp. 495503.
19. Zandleven, J., Sorensen, S.O., Harholt, J., Beldman, G.,
Schols, H.A., Scheller, H.V., and Voragen, A.G.J., Phy-
tochemistry, 2007, vol. 68, pp. 12191226.
20. Balance, S., Borsheim, K.Y., Inngjerdingen, K.,
Paulsen, B.S., and Christensen, B.E., Carbohydr.
Polym., 2007, vol. 67, pp. 104115.
21. Einhorn-Stoll, U., Kunzen, H., and Dongowski, G.,
Food Hydrocoll., 2007, vol. 21, pp. 11011112.
22. Winning, H., Viereck, N., Norgaard, L., Larsen, J., and
Engelsen, S.B., Food Hydrocoll., 2007, vol. 21,
pp. 256266.
23. Yapo, B.M., Lerouge, P., Thibault, J.-F., and Ralet, M.-C.,
Carbohydr. Polym., 2007, vol. 69, pp. 426435.
24. Kirby, A.R., MacDougal A.J., Morris V.J, Food Bio-
phys., 2006, vol. 1, pp. 5156.
25. Keegstra, K., Talmadge, K., Bauer, W.D., and Alber-
sheim, P., Plant Physiol., 1973, vol. 51, pp. 188199.
26. Yang, H., Wang, Y., Lai, S., An, H., Li, Y., and Chen, F.,
J. Food Sci., 2007, vol. 72, pp. 6575.
27. Fischman, M.L., Cooke, P.H., and Cofn, D.R., Bio-
macromolecules, 2004, vol. 5, pp. 334341.
28. Fischman, M.L., Cooke, P.H., Chau, H.K., Cofn, D.R.,
and Hotchkiss, A.T., Jr, Biomacromolecules, 2007,
vol. 8, pp. 573601.
29. Willats, W.G.T., Mc, CartneyL., Mackie, W., and
Knox, J.P., Plant. Mol. Biol., 2001, vol. 47, pp. 927.
30. Neill, M.A and York, W.S, in The Plant Cell Wall,
Rose, J.K.C., Ed., Oxford: Blackwell Publ. Ltd. Ann.
Plant Rev, 2003, pp. 154.
31. Oosterveld, A., Beldman, G., Schols, H.A., and Vora-
gen, A.G.J., Carbohydr. Res., 1996, vol. 288, pp. 143
153.
32. Schols, H.A., Bakx, E.J., Shipper, D., and Voragen, A.G.J.,
33. Novoselskaya, I.L., Voropaeva, N.L., Semenova, L.N.,
and Rashidova, S.Sh., Khim. Prir. Soedin., 2000, no. 1,
pp. 311.
34. Guillion, F. and Thibault, J.-F., Carbohydr. Res., 1989,
vol. 190, pp. 8596.
35. Vidal, S., Doco, T., Williams, P., Pellerin, P., York, W.S.,
ONeill, M.A., Glushka, J., Darvill, A.G., and Alber-
sheim, P., Carbohydr. Res., 2000, vol. 326, pp. 277
294.
36. Strasser, G.R. and Amado, R., Carbohydr. Polym.,
2002, vol. 48, pp. 263269.
37. Neill, M.A, Albersheim, P, and Darvill, A.G, in Methods
in Plant Biochemistry, vol. 2: Carbohydrates,
Dey, P.M., Ed., London: Academic Press, 1990, pp.
415441.
38. ONeill, M.A., Warrenfeltz, D., Kates, K., Pellerin, P.,
Doco, T., Darvill, A.G., and Albersheim, P., J. Biol.
Chem., 1996, vol. 271, pp. 2292322930.
39. Darvill, A.G., Mc, Neill M., and Albersheim, P., Plant
Physiol., 1978, vol. 62, pp. 418422.
40. Spellman, M.W., Mc, Neill M., Darvill, A.G., Alber-
sheim, P., and Henrick, K., Carbohydr. Res., 1983,
vol. 122, pp. 131153.
41. Stevenson, T., Darvill, A.G., and Albersheim, P., Carbo-
hydr. Res., 1988, vol. 179, pp. 269288.
42. Karkhanis, Y.D., Zeltner, J.Y., Jackson, J.L., and
Carlo, D.J., Anal. Biochem., 1978, vol. 85, pp. 595601.
43. Hilz, H., Williams, P., Doco, T., Schols, H.A., and Vora-
gen, A.G.J., Carbohydr. Polym., 2006, vol. 65, pp. 521
528.
44. Ishii, T. and Matsunaga, T., Carbohydr. Res., 1996,
vol. 284, pp. 19.
45. Ishii, T., Matsunaga, T., Pellerin, P., ONeill, M.A., Dar-
vill, A.G., and Albersheim, P., J. Biol. Chem., 1999,
vol. 274, pp. 1309813104.
46. Ishii, T. and Matsunaga, T., Phytochemistry, 2001,
vol. 57, pp. 969974.
47. Ridley, B.L., ONeill, M.A., and Mohnen, D., Phy-
48. ONeill, M.A., Ishii, T., Albersheim, P., and Darvill, A.G.,
Ann. Rev. Plant Biol. 2004, vol. 55, pp. 109139.
49. Aldington, S. and Fry, S.C., J. Exp. Bot., 1994, vol. 45,
pp. 287293.
50. Guillotin, S.E., Studies on Intra- and Intermolecular
Distributions of Substituents in Commercial Pectins,
The Netherlands: Wageningen Univ., 2005, pp. 424.
51. Guillotin, S.E., Bakx, E.J., Boulenguer, P., Mazoyer, J.,
Schols, H.A., and Voragen, A.G.J., Carbohydr. Polym.,
2005, vol. 60, pp. 391398.
52. Guillotin, S.E., van Loey, A., Bonlengner, P., Schols, H.A.,
and Voragen, A.G.J., Food Hydrocoll., 2007, vol. 21,
pp. 8591.
53. Gnanasambamdam, R. and Procter, A., Food Chem.,
2000, vol. 68, pp. 327332.
54. Huisman, M.M.H., Oosterveld, A., and Schols, H.A.,
55. Savary, B.J. and Nunez, A., J. Chromatogr., 2003,
vol. 1017, pp. 151159.
282
OVODOV
56. Strm, A., Ralet, M.-C., Thibault, J.-F., and Wil-
liams, M.A.K., Carbohydr. Polym., 2005, vol. 60,
pp. 467473.
57. Zhong, H.-J., Williams, M.A.K., Goodall, D.M., and
Hansen, M.E., Carbohydr. Res., 1998, vol. 308, pp. 18.
58. Jiang, C.-M., Wu, M.C., Chang, W.-H., and Chang, H.-M.,
J. Agr. Food Chem., 2001, vol. 49, pp. 55845588.
59. Williams, M.A.K., Foster, T.J., and Schols, H.A., J. Agr.
Food Chem., 2003, vol. 51, pp. 17771782.
60. Strm, A. and Williams, M.A.K., Carbohydr. Res.,
2004, vol. 339, pp. 17111716.
61. Rolin, C, in Pectins and their manipulation, Seymour, G.B.
and Knox, J.P., Eds., Oxford: Blackwell Publ, 2002,
pp. 222239.
62. Leroux, J., Langendorff, V., Schick, G., Vaisnav, V., and
Mazoyer, J., Food Hydrocoll., 2003, vol. 17, pp. 455
462.
63. Oosterveld, A. and Beldman, G., Searle Van Leeu-
wen M.J.F., Voragen A.G.J, Carbohydr. Polym.,
2000, vol. 43, pp. 249256.
64. Ralet, M.-C., Andr-Leroux, G., Qumner, B., and
Thibault, J.-F., Phytochemistry, 2005, vol. 66,
pp. 28002814.
65. Sosnina, N.A., Mironov, V.F., Karaseva, A.N., Minza-
nova, S.T., Karlin, V.V., Enikeev, K.M., Konovalov, A.I.,
Lapin, A.A., Kononov, A.S., and Takunov, I.P., Khim.
Prir. Soedin., 2000, no. 1, pp. 3234.
66. Turokhodzhaev, M.T., Khodzhaeva, M.A., Ivanova, I.A.,
et al., Khim. Prir. Soedin., 2000, no. 5, pp. 570572.
67. Mkrtchyan, T.A., Slepyan, G.G., and Nikogosyan, G.A.,
Ukr. Biokhim. Zh., 1998, vol. 70, pp. 98105.
68. Shkodina, O.G., Zeltger, O.A., Selivanov, N.Y., and
Ignatov, V.V., Food Hydrocoll., 1998, vol. 12, pp. 313
316.
69. Ptichkina, N.M., Markina, O.A., and Rumyantseva, G.N.,
70. Otserov, E.N. and Kostin, V.I., Uglevody amaranta i
ikh prakticheskoe ispolzovanie (Amaranth Carbohy-
drates and Their Use in Practice), Ulyanovsk: Izd. Ins.
Khim. Komi Nauch. Tsentr. Ural. Otd. Ros. Akad.
Nauk, 2001.
71. Konovalov, A.I., Otserov, E.N., and Sosnina, N.A., RF
Patent No. 212366, 1998.
72. Singthong, J., Ningsanond, S., Cui, S.W., and Goff, H.D.,
73. Dourado, F., Madureira, P., Vidal, S., and Pellerin, P.,
74. Iglesias, M.T. and Lozano, J.E., J. Food Eng., 2004,
vol. 62, pp. 215223.
75. Sanavova, M.Kh. and Rakhimov, D.A., Chem. Nat.
Products, 2004, vol. 40, pp. 8384.
76. Obro, J., Harholt, J., Scheller, H.V., and Orla, C., Phy-
77. Bush, M.S., Marry, M., Huxham, M., Jarvis, M., and
Mc Cann, M.C., Planta, 2001, vol. 213, pp. 869880.
78. Willats, W.G.T., Mc Cartney L., and Steele-King, C.G.,
Planta, 2004, vol. 218, pp. 673681.
79. Vierhuis, E., Structural Characteristic of Polysaccha-
rides from Olive Fruit Cell Walls in Relation to Ripen-
ing and Processing, The Netherlands: Wageningen
Univ, 2002.
80. Vierhuis, E., Schols, H.A., Beldman, G., and Vor-
agen, A.G.J., Carbohydr. Res., 2000, vol. 43, pp. 1121.
81. Cardoso, S.M., Silva, A.M.S., and Coimbra, M.A., Car-
bohydr. Res., 2002, vol. 337, pp. 917924.
82. Voragen, A.G.J, Beldman, G, and Schols, H.A, in
Advances in Dietary Fibre Technology, Mc Cleary, B.V.
and Prosky, L., Eds., Oxford: Blackwell, 2001, pp. 379
389.
83. Huisman, M., Elucidation of the Chemical Fine Struc-
ture of Polysaccharides from Soybean and Maize Ker-
nel Cell Walls, The Netherlands: Wageningen Univ,
2000, p. 159.
84. Duan, J., Wang, X., Dong, Q., Fang, J., and Li, X., Car-
bohydr. Res., 2003, vol. 338, pp. 12911297.
85. Wang, X.-S., Dong, Q., Zuo, J.P., and Fang, J.-N., Car-
bohydr. Res., 2003, vol. 338, pp. 23932402.
86. Habibi, Y., Heyraud, A., Mahrouz, M., and Vignon, M.R.,
87. Habibi, Y., Mahrouz, M., Marais, M.-F., and Vignon, M.R.,
88. Habibi, Y., Mahrouz, M., and Vignon, M.R., Carbohydr.
Polym., 2005, vol. 60, pp. 319329.
89. Matsuhiro, B., Lillo, L.E., Senz, C., Urza, C.C., and
Zrate, O., Carbohydr. Polym., 2006, vol. 63, pp. 263
267.
90. Yamada, H, in Bioactive Carbohydrate Polymers,
Paulsen, B.S., Ed., Dordrecht: Kluwer Acad. Publ.,
2000, pp. 1524.
91. Paulsen, B.S. and Barsett, H., Bioactive Pectic Sub-
stances, Berlin: Springer Verla, 2005, vol. 186, pp. 69
101.
92. Lee, J.-H., Shim, J.S., Lee, J.S., Kim, M.-K.,
Chung, M.-S., and Kim, K.H., Carbohydr. Res., 2006,
vol. 341, pp. 11541163.
93. Yamada, H, in Cell and Developmental Biology of Ara-
binogalactan Proteins, Nothnagel, R.D., Ed., Boston:
Kluwer Acad. Publ., 2000, pp. 221251.
94. Sakurai, M.H., Kiyohara, H., Matsumoto, T., Tsumu-
raya, Y., Hashimoto, Y., and Yamada, H., Carbohydr.
Res., 1998, vol. 311, pp. 219229.
95. Guo, Y.-J., Matsumoto, T., Kikuchi, Y., Ikejima, T.,
Wang, B.X., and Yamada, H., Immunopharmacology,
2000, vol. 49, pp. 307316.
96. Matsumoto, T., Guo, Y.-J., Ikejima, T., and Yamada, H.,
Immunol. Lett., 2003, vol. 89, pp. 111118.
97. Samuelson, A.B., Paulsen, B.S., Wold, J.K., Knutsen, S.H.,
and Yamada, H., Carbohydr. Polym., 1998, vol. 35, pp.
145153.
98. Michaelsen, T.E., Gilije, A., Samuelsen, A.B., Hogasen, K.,
and Paulsen, B.S., Scand. J. Immunol., 2000, vol. 52,
pp. 483489.
99. Hetland, G., Samuelson, A.B., Lovik, M., Paulsen, B.S.,
Aaberge, I.S., Groeng, E.-C., and Michaelsen, T.E.,
Scand. J. Immunol., 2000, vol. 52, pp. 348353.
100. Hetland, G., Curr. Med. Chem., 2003, vol. 2, pp. 135
141.
101. Habibi, Y. and Vignon, M.R., Carbohydr. Res., 2006,
vol. 340, pp. 14311436.
102. Aboughe-Angone, S., Nguema-Ona, E., Ghosh, P., Ler-
ouge, P., Ishii, T., Ray, B., and Driouich, A., Carbohydr.
Res., 2008, vol. 343, pp. 6772.
103. Egelund, J., Petersen, B.L., Motavia, M.S., Damager, I.,
Faik, A., Olgen, C.E., Ishii, T., Clausen, H., Ulvskov, P.,
and Geshi, N., The Plant Cell, 2006, vol. 18, pp. 2593
2607.
104. Ishii, T., Matsunaja, T., and Hayashi, N., Plant Physiol.,
2001, vol. 126, pp. 16981705.
105. Pagan, J. and Ibarz, A., J. Food Eng., 1999, vol. 39,
pp. 193201.
106. Pagan, J., Ibarz, A., Llorca, M., and Coll, L., J. Sci.
Food Agric., 1999, vol. 79, pp. 10381042.
107. Pagan, J., Ibarz, A., Llorca, M., Pagan, A., and Barbose,
Canovas G.V., Food Res. Internat., 2001, vol. 34,
pp. 605612.
108. Faravash, R.S. and Ashtiani, F.Z., Food Hydrocoll.,
2008, vol. 22, pp. 196202.
109. Perrone, P., Hewage, C.M., Thomson, A.R., Bailey, K.,
Sadler, I.H., and Fry, S.C., Phytochemistry, 2002,
vol. 60, pp. 6777.
110. Rosenbohm, C., Lundt, I., Christensen, T.M.I.E., and
Young, N.W.G., Carbohydr. Res., 2003, vol. 338,
pp. 637649.
111. Wehr, J.B., Menzies, N.W., and Blamey, F.P.C., Food
Hydrocoll., 2004, vol. 18, pp. 375378.
112. Fu, J.-T. and Rao, M.A., Food Hydrocoll., 2001, vol. 15,
pp. 93100.
113. Bdi, G.K., Turgeon, S.L., and Makhlouf, J., Food
Hydrocoll., 2008, vol. 22, pp. 836844.
114. MacDougal A.J., Brett G.M., Morris V.J., Rigby N.M.,
Ridout M.J., and Ring S.G., Carbohydr. Res., 2001,
vol. 335. pp. 115126.
115. Paradossi, G., Chiessi, E., and Malovikov, A., Biopoly-
mers, 1999, vol. 50, pp. 201209.
116. Tolstoguzov, V.B., Food Hydrocoll., 2003, vol. 17,
pp. 123.
117. Donato, L., Garnier, C., Novales, B., and Doublier, J.-L.,
118. Donato, L, Garnier, C, Novales, B, and Doublier, J.-L,
in Food Colloids: Interaction, Microstructure and Pro-
cessing, Dickinson, E., Ed., Cambridge: The Royal Soc.
of Chem., 2004, pp. 4858.
119. Beaulieu, M., Turgeon, S., and Doublier, J.-L., Internat.
Dairy J., 2001, vol. 11, pp. 961967.
120. Beaulieu, M., Corredig, M., Turgeon, S., Wicker, L.,
and Doublier, J.-L., Food Hydrocoll., 2005, vol. 19,
pp. 803812.
121. Gancz, K., Alexander, M., and Corredig, M., J. Agric.
Food Chem., 2005, vol. 53, pp. 2236.
122. Gancz, K., Alexander, M., and Corredig, M., Food
Hydrocoll., 2006, vol. 20, pp. 293298.
123. Matia-Merino, L. and Singh, H., Food Hydrocoll.,
2007, vol. 21, pp. 765775.
124. Liu, J., Verespej, E., Corredig, M., and Alexander, M.,
125. Liu, L.S., Fishman, M.L., Hicks, K.B., and Kende, M.,
Biomaterials, 2005, vol. 26, pp. 59075916.
126. Kratchanova, M., Slavov, A., and Kratchanov, C., Food
Hydrocoll., 2004, vol. 18, pp. 677683.
127. Takeda, C., Takahashi, Y., Seto, I., Kawano, G.,
Takayama, K., Onishi, H., and Machida, Y., Chem.
Pharm. Bull., 2007, vol. 55, pp. 11641168.
128. Marudova, M. and Jilov, N., Food Eng., 2003, vol. 59,
pp. 177180.
129. Yoo, S.-H., Fishman, M.L., Hotchkiss, Jr.A.T., and Lee, H.G.,
130. Lfgren, C. and Hermansson, A.-M., Food Hydrocoll.,
2007, vol. 21, pp. 480486.
131. Racape, E., Thibault, J.-F., Reitsma, J.C.F., and Pilnik, W.,
Biopolymers, 1989, vol. 28, pp. 14351448.
132. Yoo, S.-H., Fishman, M.L., Savary, B.J., and Hotch-
kiss, A.T., J. Agric. Food Chem., 2003, vol. 51,
pp. 74107417.
133. Lfgren, C., Walkenstrm, P., and Hermansson, A.-M.,
Biomacromolecules, 2002, vol. 3, pp. 11441153.
134. Lfgren, C and Hermansson, A.-M, in Gums and Stabilisers
for the Food Industry, Williams, P.A. and Phillips, G.O.,
Eds., Cambridge: The Royal Soc. of Chem., 2004, vol. 12,
pp. 153159.
135. Oakenfull, D.G, in Gums and Stabilisers for the Food
Industry, Williams, P.A. and Phillips, G.O., Eds., Cam-
bridge: The Royal Soc. of Chem., 2000, vol. 10, pp.
277284.
136. Hansson, A., Leufvn, A., Pehrson, K., and Stenlf, B.,
J. Agric. Food Chem., 2002, vol. 50, pp. 38033809.
137. Hansson, A., Leufvn, A., and van Ruth, S.M., J. Agric.
Food Chem., 2003, vol. 51, pp. 20002005.
138. Monge, M.E., Negri, R.M., Giacomazza, D., and
Bulone, D., Food Hydrocoll., 2008, vol. 22, pp. 916
924.
139. Monge, M.E., Bulone, D., Giacomazza, D., Negri, R.M.,
and Bernik, D.L., Combinator.. Chem. High Throughput
Screen., 2004, vol. 7, pp. 337344.
140. Monge, M.E., Bulone, D., Giacomazza, D., Bernik, D.L.,
and Negri, R.M., Sensors and Actuators B-Chemicals,
2004, vol. 101, pp. 2838.
141. Elbert, D.L., Herbert, C.B., and Hubbell, J.A., Lang-
muir, 1999, vol. 15, pp. 53555362.
142. Richert, L., Lavalle, P., Vautier, D., Senger, B., Stoltz, J.F.,
Shaaf, P., Voegel, J.C., and Picart, C., Biomacromolecules,
2002, vol. 3, pp. 11701178.
143. Marudova, M., MacDougal A.J., Ring S.G, Carbohydr.
Res., 2004, vol. 339, pp. 209216.
144. Marudova, M., MacDougal A.J., Ring S.G, Carbohydr.
Res., 2004, vol. 339, pp. 19331939.
145. Marudova, M., Lang, S., Brownsey, G.J., and Ring, S.G.,
146. Bedouet, L., Denys, E., Courtois, B., and Courtois, J.,
Carbohydr. Polym., 2006, vol. 65, pp. 165173.
147. Gorshkova, T.A., Chemikosova, S.B., Salnikov, V.V., Pav-
lencheva, N.V., Gurjanov, O.P., and Stolle-Smits, T.,
Industr. Crops and Products, 2004, vol. 19, pp. 217224.
148. Yashoda, H.M., Prabha, T.N., and Tharanathan, R.N.,
149. Orla, C., Seymour, G.B., Willats, W.G.T., van Ale-
been, G.-J.W.M., Schols, H.A., and Knox, J.P., Plant
Physiol., 2001, vol. 126, pp. 210221.
284
OVODOV
150. Orla, C., Huisman, M.M.N., Willats, W.G.T., Hux-
man, I.M., Jarvis, M.C., Dover, C.J., Thomson, A.J.,
and Knox, J.P., Planta, 2002, vol. 215, pp. 440447.
151. Regwell, R.J., MacRae E.A., Hallet I., Fisher M., Perry J.,
Harker R, Planta, 1997, vol. 203, pp. 162173.
152. Majumder, K. and Mazumdar, B.C., Sci. Hort., 2002,
vol. 96, pp. 91101.
153. Majumder, K. and Mazumdar, B.C., Indian J. Plant
Physiol., 1998, vol. 3, pp. 4245.
154. Potievskii, E.G. and Novikov, A.I., Meditsinskie aspe-
kty primeneniya pektina (Medical Aspects of Pectin
Use), Moscow: Med. Kniga, 2002.
155. Popov, S.V., Vzaimodeistvie fagotsitov mlekopitayush-
chikh s polisakharidami rastenii (Interaction of Mam-
malian Phagocytes with Plant Polysaccharides), Syk-
tyvkar: Izd. Komi Nauch. Tsentr. Ural. Otd. Ros. Akad.
Nauk, 2002.
156. Khasina, E.I., Sgrebneva, M.N., Ovodova, R.G.,
Golovchenko, V.V., and Ovodov, Yu.S., Dokl. Akad.
Nauk, 2003, vol. 390, pp. 413415.
157. Krylova, S.G., Emova, L.A., Zueva, E.P., Khotim-
chenko, M.Yu., Amosova, E.N., Razina, T.G., Lopatina,
K.A., and Khotimchenko, Yu.S., Byull. Eksp. Biol. Med.,
2008, vol. 145, pp. 678681.
158. Olano-Martin, E., Rimbach, G.H., Gibson, G.R., and
Rastall, R.A., Anticancer Res., 2003, vol. 23, pp. 341
346.
159. Nangia-Makker, P., Conclin, I., Hogan, V., and Raz, A.,
J. Natl. Cancer Inst., 2002, vol. 94, pp. 18541862.
160. Eliaz, I., Clin. Pract. Altern. Med., 2002, vol. 2,
pp. 177179.
161. Guess, B.W., Scholz, M.C., Strum, S.B., Lam, R.Y.,
Johnson, H.J., and Jennrich, R.I., Prostate Cancer and
Prostate Dis., 2003, vol. 6, pp. 301304.
162. Sriamornsak, P., Eur. J. Pharm. Sci., 1999, vol. 8,
pp. 221227.
163. Sriamornsak, P., Sungthongjeen, S., and Puttipipatkha-
chorn, S., Carbohydr. Polym., 2007, vol. 67, pp. 436
445.
164. Vandamme, T.F., Lenoury, A., Charrueay, C., and Cha-
rimeil, J.-C., Carbohydr. Polym., 2002, vol. 48, pp.
219231.
165. Liu, L.S., Fishman, M.L., and Hicks, K.B., Cellulose,
2007, vol. 14, pp. 1524.
166. Liu, L.S., Fishman, M.L., Kost, J., and Hicks, K.B.,
167. Schmidgall, J. and Hensel, A., Internat. J. Biol. Macro-
mol., 2002, vol. 30, pp. 217225.
168. Liu, L.S., Fishman, M.L., Hicks, K.B., and Kende, M.,

Current Views On Pectin Substances: Yu. S. Ovodov

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Current Views On Pectin Substances: Yu. S. Ovodov

Uploaded by

Copyright:

Available Formats

ISSN 1068-1620, Russian Journal of Bioorganic Chemistry, 2009, Vol. 35, No. 3, pp. 269284. Pleiades Publishing, Ltd.

You might also like