Professional Documents
Culture Documents
Carbohydratesingrainlegumeseeds 20003034246
Carbohydratesingrainlegumeseeds 20003034246
1
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:07 A3867: AMA: Hedley: First Proof:30-Oct-00
Color profile: Disabled
Composite Default screen
Edited by
C.L. Hedley
Department of Applied Genetics
John Innes Centre
Norwich
UK
Associate Editors
Jane Cunningham and Alan Jones
CABI Publishing
3
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:09 A3867: AMA: Hedley: First Proof:30-Oct-00
Color profile: Disabled
Composite Default screen
A catalogue record for this book is available from the British Library, London, UK.
4
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:10 A3867: AMA: Hedley: First Proof:30-Oct-00
Color profile: Disabled
Composite Default screen
Contents
Contents
Contents
Contributors xi
Preface xv
1 Introduction 1
Editor: Cliff Hedley
1.1 The Grain Legumes 1
1.2 Grain Legume Production 1
1.3 Grain Legume Consumption 7
1.4 Grain Legume Carbohydrates 11
2 Carbohydrate Chemistry 15
Editor: Pavel Kadlec
2.1 The Carbohydrates 15
2.1.1 Soluble carbohydrates 16
2.1.2 Polysaccharides 22
2.1.3 Other carbohydrate components 28
2.2 Chemical Analysis of the Carbohydrates 31
2.2.1 Soluble carbohydrates (monosaccharides, sucrose,
α-galactosides, cyclitols) 31
2.2.2 Polysaccharides 45
2.2.3 Other carbohydrate components 56
5
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:10 A3867: AMA: Hedley: First Proof:30-Oct-00 Contents
Color profile: Disabled
Composite Default screen
vi Contents
3 Nutrition 61
Editor: Halina Kozlowska
3.1 Introduction 61
3.2 The Content of Carbohydrates in Grain Legumes Utilized in
Europe 62
3.2.1 The content of carbohydrates in grain legumes used for
human nutrition 62
3.2.2 The content of carbohydrates in grain legumes used for
animal nutrition 67
3.3 Physiological Effect of Grain Legume Carbohydrates in Animal
Nutrition 69
3.3.1 Consumption of grain legume carbohydrates in feed 69
3.3.2 Effect of mono- and disaccharides in animal nutrition 71
3.3.3 Effect of oligosaccharides in animal nutrition 71
3.3.4 Effect of starch in animal nutrition 74
3.3.5 Effect of non-starch polysaccharides (NSP) in animal
nutrition 76
3.3.6 Effect of grain legume carbohydrates in ruminant nutrition 78
3.4 Physiological Effect of Grain Legume Carbohydrates in Human
Nutrition 79
3.4.1 Nutritional classification of grain legume carbohydrates 79
3.4.2 Consumption of grain legume carbohydrates in food 82
3.4.3 Physiological effect of available carbohydrates from grain
legumes 84
3.4.4 Physiological effect of unavailable carbohydrates from grain
legumes 85
4 Processing 89
Editor: Bálint Czukor
4.1 Native Starch 89
4.1.1 Isolation 89
4.1.2 Granular structure 93
4.1.3 Functional properties 98
4.2 Modified Starch 101
4.2.1 Physical methods 102
4.2.2 Chemical methods 104
4.2.3 Biotechnological methods 108
4.3 Food Application of Native and Modified Legume Starches 109
4.4 Effect of Processing on Starch and Other Carbohydrates in
Foods 110
4.4.1 Resistant starch formation 110
4.4.2 Content, composition and digestibility 112
4.5 Legume Seeds as a Source of Raw Materials 116
6
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:11 A3867: AMA: Hedley: First Proof:30-Oct-00 Contents
Color profile: Disabled
Composite Default screen
Contents vii
6 Biotechnology 145
Editor: Nickolay Kuchuk
6.1 Introduction 145
6.2 In vitro Cultures and Plant Regeneration of Grain Legumes 146
6.2.1 Introduction to in vitro culture 146
6.2.2 Plant regeneration systems 148
6.2.3 Pioneering studies on pea regeneration 149
6.2.4 Regeneration via somatic embryogenesis 150
6.2.5 Regeneration via organogenesis and multiple shoot
formation 151
6.2.6 Recent studies to produce more efficient, fast and reliable
systems for regeneration 153
6.2.7 Factors effecting regeneration 154
6.2.8 Advantages of the different developmental pathways for
in vitro manipulation 155
6.3 Isolated Protoplasts from Grain Legumes 156
6.3.1 Introduction to protoplast cultures 156
6.3.2 Protoplast cultures from leguminous species 157
6.3.3 Application of grain legumes protoplasts to the study of
carbohydrates 158
6.4 Somaclonal Variation in Grain Legumes 162
6.4.1 Introduction 162
6.4.2 Factors causing variation 163
6.4.3 Mechanisms of somaclonal variation 163
6.4.4 Potential and disadvantages of somaclonal variation 164
6.4.5 Variation in grain legumes at the cell and tissue culture
level in vitro 165
6.4.6 Variation in grain legumes at the whole plant level 172
7
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
14 November 2000 14:12:37 A3867: AMA: Hedley: First Proof:14-Nov-00 Contents
Color profile: Disabled
Composite Default screen
viii Contents
8
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:12 A3867: AMA: Hedley: First Proof:30-Oct-00 Contents
Color profile: Disabled
Composite Default screen
Contents ix
References 241
Index 315
9
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:12 A3867: AMA: Hedley: First Proof:30-Oct-00 Contents
Color profile: Disabled
Composite Default screen
Contributors
Contributors
Contributors
xi
11
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:12 A3867: AMA: Hedley: First Proof:30-Oct-00 Contributors
Color profile: Disabled
Composite Default screen
xii Contributors
12
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:13 A3867: AMA: Hedley: First Proof:30-Oct-00 Contributors
Color profile: Disabled
Composite Default screen
Contributors xiii
13
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:13 A3867: AMA: Hedley: First Proof:30-Oct-00 Contributors
Color profile: Disabled
Composite Default screen
xiv Contributors
14
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:13 A3867: AMA: Hedley: First Proof:30-Oct-00 Contributors
Color profile: Disabled
Composite Default screen
Preface
Preface
Preface
xv
15
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:14 A3867: AMA: Hedley: First Proof:30-Oct-00 Preface
Color profile: Disabled
Composite Default screen
xvi Preface
colleagues at the John Innes Centre, Jane Cunningham and Alan Jones.
Jane has been at the hub of the CABINET project, responsible for all
communication relating to meetings and to the book. Jane’s cheerful and
efficient way of administering the project is a major reason why it has been
so successful. Alan has been a constant source of support in the running of
the project and has taken on major responsibilities for editing the whole
book, including constructing and redrawing all of the tables and figures.
Cliff Hedley
16
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:14 A3867: AMA: Hedley: First Proof:30-Oct-00 Preface
Color profile: Disabled
Composite Default screen
1
C. Hedley
Introduction
Introduction 1
Editor: Cliff Hedley
There are two things which I am confident I can do very well: one is an
introduction to any literary work, stating what it is to contain, and how it
should be executed in the most perfect manner . . .
Boswell Life, vol. 1, p. 2 (1755)
Samuel Johnson (1709–1784), English poet, critic and lexicographer
17
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:15 A3867: AMA: Hedley: First Proof:30-Oct-00 1
2
Color profile: Disabled
Table 1.1. A list of the common grain legumes grown in the world and their uses.
Soybean Glycine max (L.) Economically and agriculturally the most important legume in the world, providing protein
Merrill and oil to the food and animal feed industry and the base ingredients for hundreds of
chemical products. Not agriculturally important in Europe. Also consumed as tofu and soy
sauce in Far Eastern cookery.
White (flowered) Lupinus albus L. Originating in the Mediterranean area and found widely in North America over 200 species
lupin are known. The name lupin (sometimes spelled as lupine) comes from the Latin for ‘wolf’
18
Yellow lupin Lupinus luteus L. and derives from the mistaken belief that the plant ‘wolfed’ minerals from the soil. The
C. Hedley
Lupinus mutabilis L. contrary is true, lupins aid soil fertility and are drought tolerant. Agriculturally they are grown
Sweet lupin Lupinus angustifolius L. for animal feed.
Chickpea Cicer arietinum L. Also called Bengal gram, boot, chana chola, chole, gram, hommes, pois chiche and
1
Asian origin. Desi seeds are about 120 g, wrinkled at the beak with brown, fawn, yellow,
orange, black or green colour, normally dehulled and split to obtain dhal. Kabuli seeds are
Common bean Phaseolus vulgaris L. Also called French, garden, haricot, kidney, pinto, navy (baked bean), black, pink, black eye,
cranberry, great northern or dry bean. Beans harvested green in pods or dry for human
consumption or used for canning or freezing. Originated in Central and South America.
Faba bean Vicia faba L. Also called broad, horse or tick bean. Harvested fresh for human consumption or used for
(field bean) canning or freezing. The seed is harvested dry for animal feed.
Lentil Lens culinaris One of the most ancient of cultivated foods. Of unknown origin, the lentil is widely grown
Medic. L. throughout Europe, Asia and North Africa but is little grown in the Western Hemisphere. So
called because of the lens shape of the seeds.
Cowpea Vigna unguiculata Also called Poona pea, yard-long bean, catjang. An important food source in Africa (Nigeria),
(L.) Walp parts of Asia and Mediterranean area. Typically a dhal-type paste is made from the soaked
19
dehulled seeds.
Introduction
Pea Pisum sativum L. One of the most ancient of cultivated foods. Thought to have originated in several centres;
Ethiopia, Asia Minor, Caucasus area and Afghanistan. Garden peas were used by Mendel
(1865) in his studies of inheritance. The white flowered or field pea is grown as a vining crop
1
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
3
4
Color profile: Disabled
Table 1.2. Total cultivation area (ha) and production (t) of grain legumes in the World and Europe (FAO, 2000).
20
Albania 68,328,459 68,322,546 68,327,710 68,324,683 68,332,300 68,330,000
C. Hedley
1
Czech Republic 68,351,428 68,104,697 68,359,395 68,133,391 68,352,697 68,106,602
21
Sweden 68,353,400 68,168,800 68,345,900 68,142,300 68,343,900 68,139,600
Switzerland 68,234,679 68,314,500 68,234,266 68,312,100 68,324,400 68,311,100
Introduction
6 C. Hedley
basis, the most important grain legume is dry bean, covering many varieties
of Phaseolus vulgaris and amounting to about 28 million hectares. The most
important countries for dry bean production are India and Brazil, with
Europe accounting for only about 2% of the world production.
In Europe, the area of grain legumes (excluding soybean) has
decreased from about 5 million hectares in 1997 to about 4 million in 1999
(Table 1.2). In 1999 the total production of grain legumes in Europe
was about 8 million t, the highest proportion being produced in France
(c. 32%), followed by Russia (c. 11%), Germany (c. 10%), the UK (c. 9%)
and the Ukraine (c. 7%; Table 1.2). The dominant legume within Europe
is pea, with a total growing area in 1997 of about 1 million hectares and a
production of about 4 million t, more than 70% of which was produced in
France.
Soybean is by far the most economically important legume in the world
and is cultivated on around 60 million hectares. The main producers of
soybean are the USA, Brazil, China and Argentina, which together account
for 82% of the total world area. The cultivation of this species in Europe is
rather small and is only about 1% of the world area (FAO, 1998). In the last
few years the area of soybean in Europe has decreased from about 1 million
hectares in 1989 to about 0.7 million hectares in 1995, with Italy, Romania
and France being the largest producers.
There are large differences across the world in the proportion of
cultivated land occupied by legumes (Table 1.3). In the USA, legumes
account for about 16% of the total arable land, the great majority of which
is due to soybean production. In Europe, this proportion is much lower,
amounting to about 7% in Portugal, 5% in Austria and Denmark, 4% in
France, Italy and the UK, about 3% in Hungary and 2% in the Czech
Republic. Legumes play an even less important role in the agriculture of
Ireland, Finland, Germany and Belgium.
In the world, the average seed yield of grain legumes has been at a
similar level for many years, amounting to about only 0.8 t ha−1 (Table 1.4).
On average, seed yields in Europe are about three times higher than this
world average and generally higher than in the USA. France has the highest
seed yields at about 4.7 t ha−1, with Ireland, Belgium, The Netherlands,
Denmark, the UK and Austria below this, but still at satisfactory levels
(3.3–4.7 t ha−1). In Europe, the lowest yields of legumes are found in
Portugal, Spain, Bulgaria and Romania. The high yields found in most
European countries can be attributed to more productive varieties being
grown within a more intensive agricultural system.
Also, there is variation between legume species for yield across the
world. The average yield of peas on a world scale is 1.7 t ha−1, which is half
the average yield achieved in Europe. The highest pea yields in Europe are
found in The Netherlands, Belgium, France and Ireland, with yields from
4.2 to 4.6 t ha−1 followed by Denmark, UK, Austria and Italy, with yields
from 3.2 to 3.8 t ha−1. The Czech Republic, Hungary and Poland have lower
22
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:18 A3867: AMA: Hedley: First Proof:30-Oct-00 1
Color profile: Disabled
Composite Default screen
Introduction 7
Table 1.3. The proportion of cultivated land used for legume production (%).
Europe
Austria 2.1 3.0 5.1
Belgium 0.8 – 0.8
Bulgaria 1.4 0.4 1.8
Czech Republic 2.2 0.1 2.3
Denmark 4.6 – 4.6
Finland 0.4 – 0.4
France 3.6 0.4 4.0
Germany 0.7 – 0.7
Greece 0.8 0.1 0.9
Hungary 2.1 0.5 2.6
Ireland < 0.1< – < 0.1<
Italy 1.1 2.5 3.6
Netherlands 1.2 – 1.2
Poland 1.7 – 1.7
Portugal 6.6 – 6.6
Romania 0.8 1.3 2.1
Spain 1.4 0.1 1.5
Sweden 1.1 – 1.1
UK 3.5 – 3.5
USA 0.4 15.5 15.9
World total 0.5 0.4 0.9
yields in the range 2.1–2.6 t ha−1, which is similar to the yields obtained in
the USA.
The average yield of dry beans is low throughout the world, the
highest yields being found in France, Greece and Italy (1.7–1.9 t ha−1),
while Portugal, which is the main producer of dry beans in Europe,
achieves very low yields.
Average world yields of soybeans are relatively high, at about 2 t ha−1, in
spite of the large area of this species under cultivation. The average yield
within Europe is about 2.4 t ha−1, with the main producer, Italy, and Greece
attaining more than 3 t ha−1. In Romania, where the acreage of soybean is
relatively high, yields of soybeans are about half those of Italy. By far the
major producer of soybeans in the world is the USA, with a total production
of about 60 million t, which is about half of the total world production.
23
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:18 A3867: AMA: Hedley: First Proof:30-Oct-00 1
Color profile: Disabled
Composite Default screen
8 C. Hedley
Table 1.4. Grain legume yields in the World and Europe (t ha−1) (Carrouee, 1995).
Table 1.5. Food and feed uses of grain legumes for 12 EU member states
(Carouee, 1995).
24
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:19 A3867: AMA: Hedley: First Proof:30-Oct-00 1
Color profile: Disabled
Composite Default screen
Introduction 9
Overall, however, five times more grain legume seed is used for animal
feed than for human food. Within the European Union (EU), the produc-
tion of grain legumes accounts for about 25% of the total protein required
for animal feed. The contribution of grain legumes to the total amount of
animal feed, however, is only about 8% (Table 1.6).
The pea is by far the most important grain legume in Europe, com-
prising about 77% of the total, followed by faba bean (c. 19%) and lupin
(c. 4%). Within the EU in 1996–1997, about 3.5 million t of dry peas
(c. 88%) and about 0.5 million t of faba beans were utilized in animal
feed stuff (Bourdillon, 1998), amounting to about 5% of the total raw
ingredients used by the EU compound feed industry (Table 1.7). In France
the proportion of grain legumes in the concentrate mixture is about
10% and in Belgium about 12%, while in Germany the proportion is only
about 3%. The major part is made up of cereals and other high-protein
components, in particular soybean (Pahl, 1998).
With the exception of Spain, which imports most of its grain legumes
from non-European countries (above 40%), most (greater than 80%) of
the grain legume seed consumed within Europe is European in origin –
Table 1.6. Grain legume share (as a percentage) in protein crops and protein
requirement for animal production within the EU (Carouee, 1995).
Table 1.7. Grain legumes used for animal feed in Europe (Gatel and Champ,
1998).
25
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:20 A3867: AMA: Hedley: First Proof:30-Oct-00 1
Color profile: Disabled
Composite Default screen
10 C. Hedley
France, Russia, Germany, the UK and the Ukraine being the largest
producers (Gatel and Champ, 1998). In most EU countries grain legumes
are produced by farmers in quantities that are too small and with quality
that is too variable. For this reason, the animal feed industry usually prefers
to use protein crops from abroad (e.g. soybean meal), which are more
homogenous.
In 1996, the average consumption of grain legumes in the world was
6.36 kg per person, made up of 2.51 kg of dry bean, 0.61 kg of pea and
3.26 kg of other grain legumes (Table 1.8). Human consumption of grain
legumes in European countries is relatively low. According to FAO data
(1996) in central, northern and western continental Europe the consump-
tion of grain legumes in 1996 averaged 2.37 kg per person, with a range of
0.2–9.3 kg depending on the country (Table 1.8). In many regions of the
world, including Mediterranean countries, the Middle East, North America
and East Africa, the consumption of grain legumes is three to four times
higher (ranging from 7.0 to 10.9 kg per person).
There has been a decline in the consumption of grain legumes within
Europe for several decades to its present very low level. The recent develop-
ment of vegetarian habits mainly in northern Europe, however, has stopped
this trend. The consumption of legumes is not necessarily affected by eco-
nomic factors. For example, according to household budget surveys in the
Czech Republic (Stikova et al., 1997), poor families (with small children)
had a similar consumption of legumes to average families, and among
poor, retired persons it was only slightly higher. It is interesting (Stikova
et al., 1996), that the highest consumption of grain legumes was found in
Total Total
Region Peas Beans Other range mean
26
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:20 A3867: AMA: Hedley: First Proof:30-Oct-00 1
Color profile: Disabled
Composite Default screen
Introduction 11
the group of families with the highest income, which could be due to a
higher level of education.
27
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:21 A3867: AMA: Hedley: First Proof:30-Oct-00 1
Color profile: Disabled
Composite Default screen
12 C. Hedley
Table 1.9. Protein, oil and carbohydrate composition (% of seed) of grain legume
seeds.a
Nutritionally, they contain starch, which is the major energy source for
humans and animals, plus many compounds, including the RFO and those
in the ‘fibre’ group, that either enhance or reduce nutritional value, or
have a positive or negative effect on health. Carbohydrates are also very
important, however, to the growth and development of the seed, forming
the main structural elements and the main translocation and storage
compounds. The consequences to the seed and to nutritional uses must
28
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:21 A3867: AMA: Hedley: First Proof:30-Oct-00 1
Color profile: Disabled
Composite Default screen
Introduction 13
29
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:22 A3867: AMA: Hedley: First Proof:30-Oct-00 1
Color profile: Disabled
Composite Default screen
2
Carbohydrate
P. Kadlec et al.Chemistry
Carbohydrate Chemistry 2
Editor: Pavel Kadlec
Contributors: Charlotte Bjergegaard, Krzysztof
Gulewicz, Marcin Horbowicz, Alan Jones, Pavel
Kadlec, Pavel Kintia, Christo Kratchanov, Maria
Kratchanova, Grazyna Lewandowicz, Maria
Soral-Smietana, Hilmer Sorensen and Jan Urban
31
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:23 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
16 P. Kadlec et al.
32
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:27 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 17
a-Galactosides
Oligosaccharides (from Greek oligos, a few) are compounds that give
only monosaccharide units after complete hydrolysis. Depending on the
number of monosaccharide residues per mole, oligosaccharides are
classified as trisaccharides, tetrasaccharides and so forth.
The α-galactosyl derivatives of sucrose are the most common group of
α-galactosides found in the plant kingdom. They are the most abundant
soluble sugars in plants and rank only second to sucrose in importance.
The most ubiquitous group of galactosyl sucrose oligosaccharides are
the raffinose family of oligosaccharides (RFO), so named after the first
member of this homologous series of α-galactosides.
The RFO are α(1→6) galactosides linked to C-6 of the glucose moiety
of sucrose. They are low molecular weight non-reducing sugars that are
soluble in water and water–alcohol solutions (Arentoft and Sorensen, 1992;
Arentoft et al., 1993). In addition to raffinose, this group of α-galactosides
includes stachyose, verbascose, ajugose and unnamed longer-chain oligo-
saccharides up to nonasaccharide (Cerning-Beroard and Filiatre-Verel,
1976). Chemically, the RFO may be considered as derivatives of sucrose.
Their IUPAC (International Union of Pure and Applied Chemistry) names
are listed below.
• raffinose α-D-galactopyranosyl-(1→6)-α-D-glucopyranosyl-(1→2)
-β-D-fructofuranoside
• stachyose α-D-galactopyranosyl-(1→6)-α-D-galactopyranosyl-
(1→6)-α-D-glucopyranosyl-(1→2)-β-D-fructofuranoside
• verbascose α-D-galactopyranosyl-(1→6)-[α-D-galactopyranosyl-
(1→6)-]2-α-D-glucopyranosyl-(1→2)-β-D-fructofuranoside
• ajugose a-D-galactopyranosyl-(1→6)-[α-D-galactopyranosyl-
(1→6)-]3-α-D-glucopyranosyl-(1→2)-β-D-fructofuranoside
Structures of these oligosaccharides are shown in Fig. 2.2.
The α-galactosides are often considered to be antinutritional factors,
because they are not hydrolysed by mucosal enzymes in the small intestine
of monogastric animals and pass into the lower gut where they are
33
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:28 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
18 P. Kadlec et al.
fermented with the production of gas (Cristofaro et al., 1974; Saini and
Gladstones, 1986; Price et al., 1988). Conversely, their ingestion in the form
of pure compounds in the diet increases the bifidobacteria population in
the colon, which in turn contributes positively to human health in many
ways (Minami et al., 1983; Tomomatsu, 1994; see Chapter 2).
Cyclitols
There are nine isomers of inositol, the prevalent natural form is cis-1,2,3,5-
trans-4,6-cyclohexanehexol (trivial name myo-inositol; Greek mys, muscle).
Myo-inositol is widely distributed in plants and animals and is a growth
factor for animals and microorganisms.
There are three other forms of underivatized inositol present in seeds
of some legume species: D-chiro-inositol, muco-inositol, and scyllo-inositol.
These naturally occurring isomers are synthesized from myo-inositol by
epimerization (Loewus and Dickinson, 1982; Loewus, 1990) and have the
same molecular formula (C6H12O6), and formula weight (180.16; for
further information see Hudlicky and Cebulak, 1993).
Myo-inositol is the primary source for the biosynthesis of many naturally
occurring derivatives including methyl-cyclitols (Hoffmann-Ostenhof and
Pittner, 1982). In plants of the Leguminosae family, myo-inositol is converted
by a specific O-methyl transferase into D-ononitol (4-O-methyl-myo-inositol),
34
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:33 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 19
35
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:34 A3867: AMA: Hedley: First Proof:30-Oct-00 2
20
Table 2.1. Occurrence of cyclitols, methyl-cyclitols and galacto-cyclitols in legume seeds.
Color profile: Disabled
(D-pinitol)
(ciceritol)
(galactinol)
(D-ononitol)
di-galacto-chiro-
B1 (fagopyritol B1)
tri-galacto-pinitol
galacto-chiro-inositol
methyl-scyllo-inositol
galacto-ononitol
methyl-chiro-inositol
galacto-pinitol A
inositol (fagopyritol B2)
di-galacto-pinitol A
galacto-myo-inositol
di-galacto-inositol
(galacto-ciceritol)
methyl-myo-inositol
galacto-pinitol B
myo-inositol
D-chiro-inositol
Lupin Lupinus luteus L. ++ ++ ++ + + + ++ ++ ++ ++
Lupin Lupinus albus L. ++ + + + + ++ ++ ++
Soybean Glycine max [L.] Merrill + ++ + + ++ ++ ++
Pigeon pea Cajanus cajan [L.] Millsp. ++ ++ + ++ ++ +++ +++
36
Cowpea Vigna unguiculata [L.] Walp + + + + +
Mung bean Vigna radiata [L.] Wilczek ++ ++ ++ +++ + + + + + ++ + +
P. Kadlec et al.
2
Chickpea Cicer arietinum L. ++ ++ ++ +++ + + + +++ + ++
Lucerne Medicago sativa L. ++ + + ++ ++ + + ++ +++ ++ +
+, Value between 0.05 and 0.10% of dry weight; ++, value between 0.10 and 0.50% of dry weight; +++, value above 0.5% of dry weight.
An empty cell means the level is below the limit of detection, trace amounts or no data available.
The table is compilation of data from: Aman (1979), Ford (1982), Frias et al. (1996c), Górecki et al. (1996), Horbowicz and Obendorf
(1994), Horbowicz et al. (1995), Ueno et al. (1973), Yasui (1980), Yasui et al. (1985).
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 21
inositol; Fig. 2.5) is relatively common and has been fully identified
(Quemener and Brillouet, 1983; Bernabé et al., 1993). Ciceritol is present
in the seeds of chickpea, lupin, lentil, soybean, kidney bean and lucerne.
The chickpea seed (Cicer arietinum, from which ciceritol is named), also
contains galacto-ciceritol (O-α-D-galactopyranosyl-(1→6)-O-α-D-galacto-
pyranosyl-(1→6)-O-α-D-galactopyranosyl-(1→2)-4-O-methyl-D-chiro-inositol)
(Nicolas et al., 1984). Ciceritol is a digalactosidic derivative of galactopinitol
A. Mimositol (O-α-D-galactopyranosyl-(1→6)-O-α-D-galactopyranosyl-(1→6)-
O-α-D-galactopyranosyl-(1→2s)-3-O-methyl-D-chiro-inositol) an isomer of
galacto-ciceritol, is a digalactoside derivative of galactopinitol B, and has
been reported and isolated from seeds of the Brazilian legume tree, Mimosa
scabrella (Ganter et al., 1991).
37
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:44 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
22 P. Kadlec et al.
2.1.2 Polysaccharides
Starch
Starch is the major storage carbohydrate (polysaccharide) in higher plants.
It exists in the form of granules, which are deposited as a reserve or storage
carbohydrate in plant organs such as seeds, tubers and roots. Starch is
unique among carbohydrates because it occurs naturally as discrete
38
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:52 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 23
granules (see Chapter 4). Starch granules are relatively dense, insoluble
and hydrate only slightly in cold water. They are also unique because, in
general, they are composed of a mixture of two polymers, an essentially
linear polysaccharide, amylose, and a highly branched polysaccharide,
amylopectin (BeMiller and Whistler, 1996).
39
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:56:59 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
24 P. Kadlec et al.
Fig. 2.9. Linear structure of amylopectin showing a side chain branching point.
Fig. 2.10. Amylopectin – cluster model showing the organization of the different
chain types within the molecule (Manners, 1989).
40
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:03 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 25
Fibre fraction
The first definition of dietary fibre (DF) was ‘the skeletal remains of plant
cells that are resistant to hydrolysis by the enzymes of man’. This definition
includes a wide spectrum of compounds within the DF fraction (Trowell,
1972). DF was redefined later to include ‘plant polysaccharides and lignin
which are resistant to hydrolysis by the digestive enzymes of man’ (Trowell,
1976). This chemically more precise definition restricted DF to polysaccha-
rides and lignin, but on the other hand expanded the definition to include
compounds outside the plant cell wall.
For the purpose of this book, the chemical structure of DF is described
for the cellulosic and non-cellulosic polysaccharides, including hemicellu-
loses, pectins and some associated components. In addition, the complex
structure of lignin is discussed, whereas the proteins and various amphi-
philic compounds, which also are considered as important components of
the cell wall and, therefore, DF (Andersen et al., 1997; Bjergegaard et al.,
1997a,b), are not described. An excellent review of the definition of terms
used to describe DF is given by Hall (1989).
41
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:04 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
26 P. Kadlec et al.
42
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:05 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 27
43
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:05 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
28 P. Kadlec et al.
Lignin
The term lignin is used now to refer not to a single chemical compound,
but rather to a group of structurally related amorphous, high molecular
weight, aromatic polymer compounds. They typically consist of monomeric
units of oxygen derivatives of phenylpropane with different degrees of
methoxylation of the aromatic nucleus. Lignin substances have a complex
three-dimensional structure and are insoluble both in water and in organic
solvents. Lignin is one of the chief constituents of plant cell walls and
DF, performing the role of a cementing substance with regard to the
other biopolymers of the cell walls. Lignin accounts for about 25% of the
composition of wood and occupies the second place in occurrence of
organic substances in nature after cellulose. It has been established that
lignins are heterogeneous in terms of chemical structure and molecular
mass, the molecular heterogeneity depending both on the age and the kind
of plant. They are normally linked by covalent and hydrogen bonds to
carbohydrates.
Lignification serves two main functions. It cements and anchors the
cellulose microfibrils and other matrix polysaccharides (pectins, hemicellu-
loses) and because the lignin–polysaccharide complexes are hard, they
stiffen the walls, thus preventing biochemical degradation and physical
damage to the walls. These properties of lignified walls are important in the
DF context, because they minimize the bacterial degradation of the walls
in the human colon. The occurrence of lignin in legume seeds is shown in
Table 2.2.
Saponins
Saponins are naturally occurring glycosides widely distributed in plants,
including soybean and pea seeds. Each saponin consists of a sapogenin,
which constitutes the aglycon moiety of the molecule and a sugar. The
sapogenin may be a steroid or a triterpene (the later type being most com-
mon form of saponin found in cultivated crop plants) and the sugar moiety
may be glucose, galactose, a pentose or a methyl pentose. The name comes
from Saponaria, soapwort, the root of which has been used as a soap (sapo,
Latin for soap). All saponins foam strongly when shaken in water. They
44
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:06 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 29
45
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:06 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
30 P. Kadlec et al.
46
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:07 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 31
Standards
Accurate and precise chemical analysis demands a comparison with the
known purified compound. Fortunately some of these compounds are
available commercially at a high purification, around 99% in some cases.
Sugar standards readily available from the major suppliers of laboratory
chemicals include: D(−)fructose (cat. no. F2543), D(+)galactose (G6404),
D(+)glucose (G7528), maltose (M5885), sucrose (S7903), raffinose
(R0250), and stachyose (S4001) (Sigma-Aldrich, St Louis, Missouri, USA).
Verbascose (O-VER) is available from Megazyme International Ireland
Ltd. (www.megazyme.com). Phenyl α-D-glucoside (P6626), D(+) melezitose
(M5375) or D(+) lactose (L1768) are suitable for use as internal standards,
since they are unlikely ever to be found in legume seeds in any significant
quantity.
Some of the more ‘exotic’ carbohydrates cannot be purchased and will
have to be prepared by each laboratory from biological sources. Among the
commonly found cyclitols and methylcyclitols in legume seeds, only two are
commercially available: myo-inositol, and D-pinitol. The others described in
Section 2.1.3 are unavailable. Details of procedures for the isolation and
purification of cyclitols are published in Schweizer et al. (1978) and Binder
and Haddon (1984). Table 2.4 summarizes suitable plant sources for
extracting and isolating cyclitols and galactocyclitols.
SAMPLE PREPARATION For low molecular weight sugars, water is the optimal
extraction solvent. Unfortunately, it is also an excellent solvent for
interfering hydrophilic components such as polysaccharides, proteins, etc.
In addition, α-amylases and α-galactosides, present in the plant material,
may degrade starch and the raffinose oligosaccharides if not inactivated
during, or prior, to extraction. These problems are minimized by
extraction in aqueous alcohols. Alcohol type and concentration, extraction
temperature and procedure vary considerably among the methods
described: 80% ethanol or methanol (v/v) is most commonly used, but
there are indications that in some cases these solvents lead to incomplete
extraction. Increasing the alcohol concentration above 80% (v/v) has
been shown greatly to reduce the amount of RFO extracted from plant
material (Cegla and Bell, 1977; Shukla, 1996; Bach Knudsen and Li, 1991).
Furthermore, marginally higher extraction yields have been noted with
methanol compared with ethanol (Shukla, 1996). This is in contrast to
other studies showing no difference between 80% methanol and water in
47
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:08 A3867: AMA: Hedley: First Proof:30-Oct-00 2
32
Table 2.4. Sources of standard of cyclitols, methyl cyclitols and their galactosides.
Color profile: Disabled
48
2. Common bugle (Ajuga reptans) 2. Bachmann (1993)
3. Castor bean seeds (Riccinus communis) 3. Kuo (1992)
P. Kadlec et al.
2
2. Buckwheat (Fagopyrum esculentum) 2. Horbowicz et al. (1998)
3. Jojoba beans (Simmondsia chinensis) 3. Ogawa et al. (1997)
Carbohydrate Chemistry 33
the extraction of low molecular weight sugars (Li and Schuhmann, 1980;
Li et al., 1985). It has been found that water extraction at 60°C and boiling
in aqueous ethanol (80%, v/v) gave comparable results. Muzquiz et al.
(1992) used two methods for the extraction of carbohydrates. Firstly,
extraction in 60% methanol at boiling temperature under reflux for 2 h,
and secondly, homogenization with 70% methanol for 1 min at room tem-
perature. Both methods gave satisfactory recoveries, but higher amounts
were recovered using the second method (Table 2.5).
Kvasnidka et al. (1996) compared the extraction efficiency of the RFO
from different varieties of pea, using sonication (80% ethanol (v/v), for 30
min) and boiling (80% ethanol v/v, for 30, 60, 120 min under reflux) and
found that the latter technique was twice as efficient compared with
RFO Amount added (mg) Total amount found (mg) Recovery (%)
49
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:10 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
34 P. Kadlec et al.
sonication and that the optimum extraction time for the boiling method
was 60 min.
Johansen et al. (1996) studied the effect of extraction solvents and
temperature on the extraction yields of monosaccharides, sucrose and
RFO from toasted soybean meal, cottonseed meal, field peas and a feed
mixture. They found that extraction in 80% (v/v) alcohol was strongly
influenced by the extraction temperature and that the maximum
extraction was only achieved at the boiling point. Extraction in water
and 50% (v/v) methanol or ethanol was less heat sensitive and gave
comparable results. It has also been shown that aqueous ethanol 50%
(v/v) was as effective as 50% (v/v) methanol, whereas lower yields
were observed at higher alcohol percentages. There was no consistent
difference in the extraction yield when comparing reflux with constant
stirring and water bath with occasional mixing, for any of the extraction
solvents used.
Comparison of five methods for extraction of oligosaccharides from
soybean and cottonseeds has been described by Bach Knudsen and Li
(1991).
In light of the above results, it can be concluded that the extraction
procedure for the analysis of soluble carbohydrates is always going to
be a compromise between the optimal extraction of a group of different
compounds, the level of recovery and the possible interaction of other
non-carbohydrate components present in the seed.
50
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:10 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 35
51
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:11 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
36 P. Kadlec et al.
split 1 : 50. The detector gas was hydrogen at 30 ml min−1 and air at
300 ml min−1. It should be noted that these operating conditions can be
transferred to other systems and used as a starting point for optimizing the
GC conditions. J & W Scientific are now able to supply high temperature
versions (DB-1ht) of the column used above. The advantages of this new
technology is that high boiling point TMS derivatives (e.g. TMS-verbascose)
will elute faster and produce sharper peaks, when hydrogen is used as the
carrier gas and it is possible to complete a GC analysis in 20 min! A typical
chromatogram using the above system is shown in Fig. 2.11 (from D.A.
Jones, personal communication).
Many analyses of cyclitols, galactocyclitols and other carbohydrate
contents in seeds of several Leguminosae species (and other species) have
been performed using this procedure (Horbowicz and Obendorf, 1994;
Horbowicz et al., 1995; Górecki et al., 1996; Horbowicz et al., 1998).
52
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:16 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 37
53
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:17 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
38 P. Kadlec et al.
54
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:17 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 39
eluted with deionized water to remove anionic substances from the sample.
The eluant is filtered through a Uniflo membrane prior to HPLC analysis
(Kuo et al., 1988).
55
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:18 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
40 P. Kadlec et al.
56
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:19 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 41
57
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:19 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
42 P. Kadlec et al.
ADVANTAGES OF TLC METHOD TLC can be a useful method for the initial
and rapid screening of material for soluble carbohydrate content before a
more detailed study is undertaken, as demonstrated by Jones et al. (1999b).
58
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:20 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 43
59
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:20 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
44 P. Kadlec et al.
REDUCING SUGAR METHOD There are several well known tests which make
use of the reducing action of sugars in alkaline solutions in the presence of
certain metallic salts, e.g. copper, silver, mercury and bismuth. Those of
copper have been employed by far the most extensively in sugar analysis
(Pearson, 1976). The basic form of the reaction is:
RCHO + Ag2O → 2 Ag + RCOOH
RCHO + 2 CuO → Cu2O + RCOOH
Unfortunately, for quantitative work these reactions do not proceed
stoichiometrically. This is because of the ability for many sugars to
mutarotate. This causes the carbon chain eventually to break and the free
aldehyde and ketone groups are lost. Close control of the reaction condi-
tions is needed along with calibration using standards. Fehling originally
devised the most widely known test based on this method, in 1848.
60
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:21 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 45
2.2.2 Polysaccharides
Starch
Many different ways for the determination of starch have been described in
the literature. In principle, two groups of methods can be distinguished.
Firstly, there are the polarimetric methods in which the starch is quantified
as a dissolved and partly degraded polymer (determination according to
Ewers and the calcium chloride method of the AOAC, see below). In the
second group of methods, the starch is fully hydrolysed into glucose and
then quantified by measuring the glucose content.
61
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:21 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
46 P. Kadlec et al.
62
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:22 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 47
63
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:23 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
48 P. Kadlec et al.
Table 2.6. In vitro and in vivo quantification of resistant starch (RS) as a propor-
tion of the total starch (TS).
in vitro in vivo
Champ Englyst
Origin of the RS methoda methodb Ileostomyc Incubationd
Beans 17 – – 17
Raw green banana 61 71 68 84
Retrograded high amylose corn starch 30 34 – 49
aChamp (1992); bEnglyst et al. (1992); cGöteborg (1995); dFaisant et al. (1993,
1995).
easier to reproduce than the Englyst method. Thirdly, the Englyst method
may reflect better the in vivo physiology than the other methods.
With all of the methods for RS analysis there is a fundamental problem
related to the definition of RS. None of these methods, including the one
developed by Englyst et al., takes into account the whole amount of RS
defined as ‘starch and products of starch degradation not absorbed in
the small intestine of healthy individuals’, since low molecular weight
fragments, soluble in aqueous ethanol, are not determined.
MEGAZYME KIT This method determines the total starch in a sample using a
‘kit’ sold by Megazyme International Ireland Ltd, Co. Wicklow, Ireland
(www.megazyme.com). It is based on the principles described in AACC
method 76–12 and is listed as AACC method 76–13. For samples that do
not contain high levels of resistant starch (e.g. wheat flour), complete
solubilization of starch is achieved by cooking the sample in the presence
of thermostable α-amylase. Samples that contain high levels of RS (e.g.
high-amylose maize) are completely solubilized by pre-treatment with
DMSO at 100°C. Glucose produced by the enzymatic hydrolysis of the
solubilized starch is measured using glucose oxidase/peroxidase reagent.
Samples containing high levels of glucose or maltodextrins have to be
washed with aqueous ethanol before analysis.
Fibre fraction
Some of the earliest analytical methods of DF include crude fibre, acid
detergent fibre (ADF) and neutral detergent fibre (NDF) as the most
commonly used methods. The principle of the crude fibre method implies
boiling and extraction of the sample by dilute acid and dilute alkali, with
subsequent isolation of the insoluble residue by filtration (AOAC methods
920.86, 962.09). The crude fibre method essentially determines the
cellulose and lignin content, however, the recovery may vary markedly.
64
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:23 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 49
65
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:24 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
50 P. Kadlec et al.
systems may be used, e.g. HPLC (Quigley and Englyst, 1992) and HPCE
(Rassi and Mechref, 1996). Alternatively, colorimetry may be used for
determination of groups of pentoses, hexoses and uronic acids; however
this technique is seldom used as routine. Uronic acids also may be analysed
by decarboxylation (Theander et al., 1994). Lignin is not included in the
method, but can be determined separately. The NSP may be separated into
a soluble and insoluble part in some of the methods. Monosaccharide
losses during the polysaccharide acid catalysed hydrolysis step are a
problem of major concern in enzymatic–chromatographic methods. The
conditions in acid hydrolysis are actually a compromise between complete
liberation and destruction of monosaccharides. Various methods exist,
differing in type and concentration of acid as well as temperature and time
used for hydrolysis. An AOAC Official Method has now been accepted
(Method 994.13).
The analytical methods for determination of DF described above
give only limited information on the level and nature of individual DF
components. More specific studies were originally performed using
the classical fractionation schemes evolved in the 1930s (Southgate,
1995a).
The experimental conditions used in these fractionations were
extremely vigorous and generally caused some severe modifications of the
structures of the components. Modern techniques for fractionation are
generally more gentle, although the possibility still exists, that a certain
number of bonds must be broken in order to extract components from the
cell wall, leading to incorrect conclusions about the chemistry, especially of
cell wall polysaccharides. Moreover, the specific procedure for extraction
of a particular type of component may result in only partial recovery of the
expected polysaccharide.
The fractionation is performed on purified plant material, and not
directly on the fresh tissue. Methods for preparation of plant material prior
to fractionation are described in the following sections, examples of the
analytical approaches for determination of the subfractions: cellulose,
hemicellulose, pectins and lignins are given, extraction of minor DF
compounds such as, e.g., phenolics are described briefly, and more details
are given elsewhere (Andersen et al., 1997; Bjergegaard et al., 1997a,b).
66
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:24 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 51
67
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:25 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
52 P. Kadlec et al.
fast and cheap to use supercritical fluid techniques (super critical extrac-
tion (SCE)/super critical chromatography (SFC); Andersen et al., 1997;
Bjergegaard et al., 1997a,b).
Fig. 2.12. Sequential extraction of pectin material from purified cell walls.
68
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:31 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 53
69
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:32 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
54 P. Kadlec et al.
70
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:32 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 55
71
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:33 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
56 P. Kadlec et al.
procedure and this must be taken into consideration when the method is
evaluated.
Pectin
Common methods exist for the isolation of pectic substances from plant
materials, and the main steps are outlined below.
1. Grind the plant material in warm ethanol or acetone.
2. Wash with ethanol, to inactivate endogenous enzymes.
3. Wash with sodium deoxycholate (SDC) (or enzyme treatment), to
remove proteins.
4. Wash with phenol–acetic acid–water mixture, to remove lipids and
pigments.
5. Treat with aqueous 90% DMSO (or enzyme treatment) to remove
starch.
6. Wash with ethanol, to remove the other organic solvents.
72
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:34 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 57
DETERMINATION OF NEUTRAL SUGARS The most widely used method for sugar
determination involves acid hydrolysis of the sample with 2 M trifluoro-
acetic or 1 M sulphuric acid, or by Saeman hydrolysis, which uses 72%
sulphuric acid for solubilization and subsequent hydrolysis with 0.4 M
sulphuric acid (Selvendran et al., 1979). After hydrolysis, the sugars
released are reduced to corresponding alditols and then converted to
alditol acetates. These volatile derivatives can be reliably analysed by GC as
single peaks (Blakeney et al., 1983). Uronides are not determined by this
method, but may be assayed together with neutral sugars by making TMS
derivatives of the anomeric methyl glycosides obtained by methanolysis.
In order to resolve the many peaks obtained from all sets of isomers, it is
necessary to use capillary columns in the GC.
The monosaccharides are subsequently analysed by high performance
anion exchange chromatography without the need for derivatization. It is
also possible to determine galacturonic acid and neutral sugars by a combi-
nation of enzymic hydrolysis and methanolysis, followed by HPLC separa-
tion on a C18 column eluted by water (Quemener and Thibault, 1990).
The types of glycosidic linkages between sugar residues are in general
determined by methylation analysis. Glycosyl-linkage analysis of uronosyl
residues in polysaccharides is possible only after reduction to the
corresponding neutral sugar units.
Lignin
The methods of lignin determination are essentially the same or similar to
those used to analyse the fibre, hemicellulose and cellulose fractions in
legume seeds. Suitable methods are summarized below.
VAN SOEST METHOD The residue of ADF is added to 72% H2SO4 at 0–4°C
for 3 h with stirring. After hydrolysis, the residue is filtered, and washed
with hot water, then acetone and dried in a 100°C oven for 12 h, cooled in a
desiccator and weighed.
KLASON METHOD In this procedure lignin is the residue after all the
enzyme and chemical treatments of the Englyst DMSO method have been
completed.
GOERING AND VAN SOEST METHOD The residue from the ADF is treated with
potassium permanganate solution, containing trivalent iron and mono-
valent silver as catalysts. Deposited manganese and iron oxides are dissolved
with an alcoholic solution of oxalic and hydrochloric acids, leaving
73
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:34 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
58 P. Kadlec et al.
Saponins
74
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:35 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
Carbohydrate Chemistry 59
(76 : 4 v/v) for a distance of 5.5 cm. Plates are air-dried and visualized with
a p-anisaldehyde reagent.
HPLC ANALYSIS Aliquots (10 µl) of each extract can be injected directly
on to a 250 × 4.6 mm Spherisorb S5 ODS 2 column and eluted using an
acetonitrile : water : trifluoracetic acid mixture with the composition
changing from 80 : 20 : 0.1 v/v to 20 : 30 : 0.1 v/v over a time span of
25 minutes using an eluant flow rate of 1 ml min−1. Detection is by UV
absorption at 210 nm.
For further details of experimental systems, see Fenwick et al. (1991)
and Tsukamoto et al. (1993).
75
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:36 A3867: AMA: Hedley: First Proof:30-Oct-00 2
Color profile: Disabled
Composite Default screen
3
H. Kozlowska et al.
Nutrition
Nutrition 3
Editor: Halina Kozlowska
Contributors: Pilar Aranda, Jana Dostalova,
Juana Frias, Maria Lopez-Jurado, Halina
Kozlowska, Jan Pokorny, Gloria Urbano,
Concepcion Vidal-Valverde and Zenon
Zdyunczyk
3.1 Introduction
The carbohydrate fraction of grain legumes is a major source of food
and feed energy. The optimal composition of grain legume carbohydrates,
however, depends on a number of factors, for example, consumer demand
and the requirements for animal feedstuff.
The diets for intensively farmed animals are required to have a high
energy value. The desired direction for grain legume breeders and proces-
sors, therefore, is to decrease the content of those carbohydrates that have
low energy values, or, which act as antinutrients. In this context many
authors classify α-galactosides in grain legumes as antinutrients (Eskin et al.,
1980; Saini, 1989). It has been known for many centuries that peas and
beans, although nutritional wholesome foods, produce wind (Gerarde,
1633). To quote from the British Herbal (Hill, 1756) ‘The fruits of these sev-
eral kinds are all of the same quality, wholesome as food, but apt to breed
wind’. High quantities of α-galactosides and non-starch polysaccharides are
believed to cause flatulence and reduce the net energy of seeds (Fernandez
and Batterham, 1995). From the point of view of animal nutrition, low
bioavailability of starch is also a disadvantage of grain legumes.
77
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:37 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
62 H. Kozlowska et al.
The most common legumes for human consumption are bean (Phaseolus
vulgaris), lentil (Lens culinaris), pea (Pisum sativus), chickpea (Cicer arie-
tinum) and faba bean (Vicia faba). The carbohydrate fraction in the seeds of
these legumes is composed of soluble carbohydrates (mainly fructose,
sucrose and low molecular weight oligosaccharides such as ciceritol,
raffinose, stachyose and verbascose), starch and longer chain oligosaccha-
rides and polysaccharides constituting dietary fibre. Table 3.1 collates the
information found in the literature for mono- and disaccharides, low
molecular weight oligosaccharides (or α-galactosides), total soluble sugars
and starch from bean, pea, lentil, chickpea and faba bean.
78
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:37 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 63
Table 3.1. Soluble carbohydrate and starch content (average and range as % dry
matter) of some common grain legumes, used mainly for human consumption.
79
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:38 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
64 H. Kozlowska et al.
80
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:38 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 65
Ciceritol was not present in either faba bean or pea. The content of
raffinose in faba bean was lower than in bean and pea and the stachyose
content was the lowest amongst the grain legumes reported (Table 3.1).
The starch content of the six varieties of faba bean reported in the
literature ranged from 39.2 to 47.2%.
DF is defined physiologically as the total amount of polysaccharides
and lignin not digested by endogenous enzymes of the human gastro-
intestinal tract. Since the composition of DF is complex, different methods
have been used to quantify its content and constituents. The composition
of DF reported can vary widely, therefore, depending on the methodology
used. Taking this consideration into account, a large amount of data has
been recorded on the content and composition of DF in legumes used for
human consumption.
The DF content in legume seeds depends on many factors, including
the species and the variety (Table 3.2). The DF content of common beans
indicated by different authors (Naivikul and D’Appolonia, 1978; Fleming,
1980; Chen and Anderson, 1982; Fidanza et al., 1982; Reddy et al., 1984;
Garcia-Olmedo et al., 1985; Paul and Southgate, 1985; Souci et al., 1986;
Lintas et al., 1992; Mongeau and Brassard, 1994, 1995; Vidal-Valverde et al.,
1992c) ranged from 11.2 to 27.5%, the contribution of the soluble compo-
nent being the highest for the grain legumes (8.1–10%). The DF content in
bean, noted by different authors, showed considerable differences for the
same types of seeds (Table 3.3). The results presented refer to the raw seeds
of bean. During cooking, changes in the DF content and composition can
be observed. Acevedo et al. (1994) noted that the DF content in black beans
differed according to the type of processing; it reached 26.5% in cooked
seeds, 28.1% in blended seeds and 29% in fried seeds. The proportion of
soluble DF in cooked, blended and fried beans reached 31.7, 26.7 and
Table 3.2. Content of dietary fibre and its components (as % dry matter) of some
common grain legumes, used mainly for human consumption.
NDF, neutral detergent fibre; ADF, acid detergent fibre; TDF, total dietary fibre;
SDF, soluble dietary fibre; IDF, insoluble dietary fibre; NSP, non-starch polysaccha-
rides.
81
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:39 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
66 H. Kozlowska et al.
Table 3.3. Dietary fibre (DF) (as a percentage of dry matter, DM) content in beans
(Phaseolus vulgaris).
82
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:40 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 67
Grain legumes utilized in Europe for animal nutrition vary widely in terms
of the content and composition of sugars, ranging from about 5% in faba
bean to about 13% in yellow lupin (Table 3.4). Total soluble sugars contain
only a small amount of monosaccharides and from 1 to 3% of sucrose.
Soluble sugars consist mostly of α-galactosides. The lowest content of α-
galactosides was found in faba bean seeds (less than 2.5%), an intermediate
83
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:40 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
68 H. Kozlowska et al.
Total Soluble
α-galacto- carbohy-
Sucrose Raffinose Stachyose Verbascose sides drates Starch
level in pea (about 5%), and the highest level in lupin (up to 12%). The
seeds of white lupin are characterized by having very small amounts
of verbascose, their main sugar being stachyose. The highest amounts of
verbascose were found in the seeds of yellow and blue lupin, pea and
faba bean (25, 33 and 50%, respectively) as a proportion of the total
α-galactosides.
Starch is the main carbohydrate present in pea and faba bean, which
are used commonly for animal nutrition. Only in lupin seeds is the
starch content below 1% DM. Abreu and Bruno-Soares (1998) analysed the
chemical composition of nine legume species and found the following
starch contents: 45.3% for pea, 40.0% for faba bean, 0.8% for narrow leafed
lupin and 0.7% for yellow lupin. The average starch content in the seeds
of 36 different lines of feed peas analysed by Bastianelli et al. (1998) was
49.2%, while the average for six lines of wrinkled pea was only 29.4%.
Apart from soluble sugars and starch, grain legume seeds are an impor-
tant source of dietary fibre in the diets of animals. The data presented
in Table 3.5 indicate that legumes utilized in animal feed differ in the
content as well as the composition of DF. The highest amounts of DF occur
in the seeds of Lupinus angustifolius (39.2%) and the main NSP components
in this species are galactose and glucose. A lower proportion of DF is found
in the seeds of Lupinus luteus, where glucose is the main component.
84
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:41 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 69
Table 3.5. The content of dietary fibre (DF) and composition of non-starch
polysaccharides (NSP) in seeds of different legume species (all data as g kg−1 of dry
matter) (Gdala and Braczewska, 1997a; Gdala et al., 1997b,c).
85
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:42 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
70 H. Kozlowska et al.
Table 3.6. Content of α-galactosides (GAL) in diets for pigs with the share of
different components rich in crude protein (CP).
CP GAL A B A B
86
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:43 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 71
87
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:43 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
72 H. Kozlowska et al.
Table 3.7. Ileal digestibility of α-galactosides (GAL) in pigs fed on a diet, without
(−) or with (+) α-galactosidase supplementation.
α-Galactosidase supplementation
88
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:44 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 73
Table 3.8. Absorption of nutrients (mg rat−1 h−1) from perfusion fluid supple-
mented with fructo-oligosaccharides (FOS) or oligosaccharides (OS) from lupin
seeds, administered to rat small intestine (Zdu4czyk et al., 1999).
89
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:45 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
74 H. Kozlowska et al.
Starch was extracted from seeds according to the methods of Faulkes et al. (1989).
90
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:46 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 75
91
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:46 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
76 H. Kozlowska et al.
and 84.4% for fine (< 100 µm) and coarse (> 100 µm) particle fractions,
respectively (Carré et al., 1998).
The nutritional value of diets rich in pea and faba bean starch can be
improved through heating or autoclaving (Longstaff and McNab, 1987;
Conan and Carré, 1989). Pelleting has been shown to increase starch
digestibility from 84% to over 95% (Carré et al., 1991). Since starch
digestibility correlates well (r 2 = 0.80) with the true metabolizable energy
obtained from pea (Longstaff and MacNab, 1987), it is apparent that
pelleting will increase the metabolizable energy of diets (Table 3.11).
Pelleting is the usual process in the feed industry and is very useful for
poultry feed, because chickens have a reduced feed intake when their
diet is finely ground. Similarly, extrusion has a beneficial effect on the
nutritional value of diets containing grain legumes. Extrusion increases
the in vitro rate of hydrolysis of starch by pancreatic amylase, and stimulates
the activity of amylase, chymotrypsin and carboxypeptidase A in the
pancreatic tissue. As a consequence, the apparent digestibility of starch in
the ileum has been to increase from 94.4 to 99.1% (Bengala Freire et al.,
1991).
Table 3.11. Nutritional value of meal and pelleted diets with pea seeds
(Peyronnet et al., 1996).
92
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:48 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 77
legume seeds depends on the age of the pigs and on the species of legume
seed, and ranges from 12.1 to 39.9% (Table 3.12; Gdala et al., 1997a,b).
The differences in digestibility coefficients presented in Table 3.12
result from differences in the grain legume species for the level and compo-
sition of the NSP. In pea and faba bean seeds the monosaccharide residues
of glucose, arabinose and uronic acids dominate the NSP fraction (Gdala
and Buraczewska, 1997a). Galactose and glucose are the main components
of lupin seed NSP (in total 60–66%), while uronic acids, arabinose and
xylose have intermediate levels (Gdala and Buraczewska, 1997a). Most of the
arabinose in pea and faba bean is present as arabinose-containing pectin
substances in the cell walls of the cotyledons (Selvendran, 1984). Pectins are
more rapidly and extensively digested in the large intestine compared with
cellulose, arabinoxylans and xylan polysaccharides (Gdala et al., 1997b).
Pigs are better at utilizing the energy derived from seeds containing a
high level of NSP. Poultry are able to digest only the water-soluble fraction
of NSP, while the water-insoluble fraction remains virtually undigested
(Carré et al., 1998). The NSP digestibility for pea diets is about 5.9%
in adult cockerels and only 2.8% in chickens (Carré et al., 1995). The
apparent ileal digestibility of DF of milled, or crushed, dehulled peas by
cockerels and chickens was 15 and 8%, respectively (Daveby et al., 1998).
Generally, NSP are the main constituents of the DF fraction in grain
legumes and a high content of DF in diets has a negative effect on nutrient
digestibility in animals (Freire et al., 1997). The insoluble DF fraction
decreases intestinal transit time, increases faecal bulk, delays glucose
absorption and slows starch hydrolysis. The water-soluble fraction of DF
increases the viscosity of the digest in the small intestine, depressing nutri-
ent absorption (Low, 1985). Dietary fibre can be a negative factor, there-
fore, that dilutes the energy content and decreases the nutrient availability
for animals. The soluble DF usually comprises about one-third of the total
DF in cereals, whereas in pea it is about 25% and about 32% in lupin (Bach
Knudsen, 1997). It has been demonstrated that a 1% increase of crude fibre
NSP residues Peaa Faba beanb Yellow lupin Narrow leaf lupin
93
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:49 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
78 H. Kozlowska et al.
in the diet diminished the digestibility of gross energy by 1.3% and the
utilization of metabolizable energy by 0.9% in pigs (Just et al., 1983).
Many authors have reported that dehulling seeds results in a lower DF
content and a higher nutritional value. The crude protein content of
dehulled white lupin seeds has been shown to increase by 20% and the
crude fibre content to decrease by 67% (Flis et al., 1997). As a consequence
of changes in the proportion of nutrients, the nutritional value of dehulled
seeds of L. albus and L. angustifolius for pigs was increased by 5–10 and 25%,
respectively (Fernandez and Batterham, 1995; Flis et al., 1997). Dehulling
lupin seeds has been shown also to increase the digestibility of energy in
chickens by 18% (Brenes et al., 1993). In addition, enzyme supplements
added to legume seed diets for chickens also have a positive effect on bird
performance (Brenes et al., 1993). In the case of pigs, enzyme supple-
mentation of diets is less effective compared with poultry because of more
intensive fermentation of sugars in the hindgut (Bedford et al., 1992).
Table 3.13. Carbohydrate content, the digestibility of energy and gas production
for different legume seeds (Abreu and Bruno Soares, 1998).
94
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:50 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 79
95
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:51 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
80 H. Kozlowska et al.
96
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:51 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 81
Table 3.16. The content of resistant starch (RS) in legume products (g (100 g)−1 of
starch).
Methods of RS RS
Legume products determination content References
Raw pea starch Weight experiment on ratsa 1.0 Berggren et al. (1995)
Pre-cooked lentil Weight experiment on ratsa 11.0 Tovar et al. (1992)
Pre-cooked red bean Weight experiment on ratsa 8.0 Tovar et al. (1992)
Canned pea Colectomized rats 15.2 Hildebrandt and
Marlett (1991)
Canned peab Weight experiment on ratsa 27.8 Björck and Sijeström
and three methods in vitro (1992)
Boiled red lentil In vivo, in ileostomists 13.8 Steinhart et al. (1992)
In vivo, in ileostomists 13.6 Jenkins et al. (1987)
Boiled white beans In vivo, intubation experi- 16.5 Noah et al. (1998)
ments in healthy subjects
Autoclaved white In vivo, in ileostomists 5.7 Muir and O’Dea (1993)
beans
Autoclaved white In vivo, in ileostomists 20.9 Schweizer et al. (1990)
beans
Boiled lentil In vitro digestibility of 16.5 Cummings and Englyst
starch (1995)
Boiled red lentil In vitro based on chewing 23.1 Äkerberg et al. (1998)
Boiled white beans In vitro based on chewing 16.7 Äkerberg et al. (1998)
Autoclaved white In vitro based on chewing 13.8 Äkerberg et al. (1998)
beans
aRats treated with antibiotics to suppress hindgut fermentation.
bPea with low content of starch – 19% dry matter (DM).
97
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:52 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
82 H. Kozlowska et al.
Immature seeds, dry seeds and processed seeds are all used for food pur-
poses. Most grain legumes, after simple processing (sprouting, cooking),
are consumed as vegetables, salads, soups, mashed seeds and cooked seeds.
The annual consumption of grain legume carbohydrates can be estimated
from the consumption and chemical composition of legumes. Examples of
the average consumption of carbohydrates in grain legumes are presented
in Table 3.17.
From these data the average daily consumption in 1996, of
α-galactosides, digested starch, RS and DF can be calculated and shown
to be very low (0.26, 2.89, 0.51 and 1.33 g, respectively). A similar intake of
digested starch and resistant starch from dried legumes (2.3 and 0.5 g,
respectively) has been found in relation to the Italian population
(Brighenti et al., 1998). Also, the daily intake of starch and RS in fresh,
frozen and canned grain legumes was 1.8 and 0.6 g, respectively (Brighenti
et al., 1998).
The calculated amounts given in Table 3.17 do not take into account
the considerable losses that occur during food preparation. For physio-
logical reasons, decreasing the α-galactoside content during preparation
of the seeds for consumption is very important, as will be discussed later.
The intake of α-galactosides in legume dishes in the Czech Republic and
in neighbouring countries are presented in Table 3.18, after taking into
consideration the losses during the preparation of legume dishes.
Considering the amount of starch in pea seeds (c. 45%), the RS content
in one portion of pea soup can range from 1.3 to 1.9 g. In one portion
of mashed pea (75–105 g), the content of total starch and RS ranges from
98
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:53 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 83
Table 3.18. Estimated intake of α-galactosides (GAL) in legume dishes (g per dish)
(Pokorny and Dostalova, unpublished).
Dry legumes Original GAL Losses on Final intake of
Dish (g) (g) cooking (%) GAL (g)
99
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:54 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
84 H. Kozlowska et al.
Relatively few studies have tried to quantify the digestibility of starch from
legumes in the small intestine of humans (Wolever et al., 1986; Schweizer
et al., 1990; Bothman et al., 1995). Numerous studies conducted on mono-
gastric animals, however, have revealed that starch is easily digested in the
upper intestinal tract, however, its digestibility coefficient is lower than that
of cereal starch (see Section 3.3.4). This low digestibility of starch together
with the low content of mono- and disaccharides and high content of
unavailable carbohydrates (α-galactosides, RS and NSP), make legume
seeds desirable component of human diets.
It is apparent that the seeds from grain legumes are characterized by
their relatively low glycaemic index (Table 3.19), which on average is less
than half that of white and wholemeal bread. It is beneficial, therefore, to
include grain legumes in diets for people with insulin-dependent diabetes,
an illness that is often found in elderly inhabitants of industrialized
countries (Wolever and Brand-Miller, 1995).
Food type GI
Cereal products
Bread, white 69
Bread, wholemeal 72
Rice, white 66
Rice, brown 66
Spaghetti, white 50
Corn flakes 80
Root vegetables
Carrots 92
Potato, new 70
Potato, instant 80
Legumes
Beans, navy 31
Beans, kidney 29
Beans, soya 15
Peas 33
Lentils 29
100
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:54 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 85
When grain legumes are used in diets as substitutes for animal products
(in the case of vegetarians) they are believed to act in two ways. Firstly, to
decrease the consumption of saturated fats and secondly to increase
the content of unavailable carbohydrates in the diet, thus reducing the
incidence of digestive tract cancers. A positive association with protein
and fat (r = 0.60 and 0.62, respectively) and a weak negative association
with NSP (r = −0.23) have been shown in a study of food consumption in
12 countries (Cassidy et al., 1994). A much stronger (r = −0.70) inverse
correlation was found, however, between colorectal cancer incidence and
starch intake.
101
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:55 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
86 H. Kozlowska et al.
102
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:56 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
Nutrition 87
Values (with legumes) in bold type represent a significant difference (P = 0.95) from
the corresponding without legumes dish.
103
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:56 A3867: AMA: Hedley: First Proof:30-Oct-00 3
Color profile: Disabled
Composite Default screen
4
B. Czukor et al.
Processing
Processing 4
Editor: Bálint Czukor
Contributors: Tatiana Bogracheva, Zsuzsanna
Cserhalmi, Bálint Czukor, Jozef Fornal, Ildikó
Schuster-Gajzágó, Erzsébet T. Kovács, Grazyna
Lewandowicz and Maria Soral-Smietana
Starchy legume seeds are rich in protein, starch and dietetic fibre, all of
which are very valuable for food and non-food applications (Salunkhe et al.,
1989). For this reason the processing of legume seeds includes the separa-
tion and production of these components. Studies that have led to the
development of industrial processing of grain legume seeds have been
carried out mainly on pea (Pisum sativum) and faba bean (Vicia faba). The
processes that are generally used are dry processing (air classification)
and wet processing. In general, the dry processing procedure produces
protein-rich and starch-rich products, while the wet processing procedure
produces purified protein, starch and dietetic fibre fractions. There are
several advantages of the dry process: the construction of pilot plants is
relatively simple, the process does not produce waste water and changes in
the structure and functional properties of the components are minimized.
Seed processing, which includes water extraction, is more complicated but
the higher purity of the products produced allows a wider range of applica-
tions. In addition, the wet process is necessary for scientific research on
105
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:57 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
90 B. Czukor et al.
starches, which demands a very high purity of starch. The basic principles
of these two processes are described in more detail below.
Dry processing
It has been found that the air classification process can be carried out
more successfully with grain legumes where starch is the main storage
product rather than oil. Among the starchy grain legumes, air classification
has been carried out on pea (P. sativum), faba bean (V. faba), mung bean
(Vigna radiata), green lentil (Lens culinaris), navy bean (Phaseolus vulgaris),
baby lima bean (Phaseolus lunatus) and cowpea (Vigna unguiculata). For
detailed information, see Reichert and Youngs (1978), Bramsnaes et al.
(1979), Talyer et al. (1981), Reichert (1982), Sosulski et al. (1985), Clott
et al. (1986) and Sosulski and McCurdy (1987). The first stage of this
process includes fine milling of the seeds. Flours prepared from starch-rich
seeds contain two distinct populations of particles, which differ in both size
and density and can be separated in a current of air. The starches and
dietetic fibres are concentrated mostly in the light, fine fraction and the
proteins and lipids in the heavy, coarse one. Repeating the air classifying
process can increase the purity of the fractions, however it decreases the
recovery of the products. Starch fractions with protein impurities as low as
2.5% can be produced; however, the recovery of the starch fraction is
only about 40% (Reichert and Youngs, 1978). Following air classification of
pea meal it has been found that the protein-rich fraction contains mainly
storage proteins, while the starch-rich fraction contains other functional
proteins, which adhere to the surface of the starch granules. The process of
air classification is illustrated in Fig. 4.1, and the influence of the number of
steps on the purification and recovery of the starch fraction from peas in
Fig. 4.2. Detailed information on the processing of pea and faba bean by air
classification has been described in a number of reports (Vose et al., 1976;
Reichert and Youngs, 1978; Bramsnaes et al., 1979; Vose, 1980; Talyer et al.,
1981; Reichert, 1982; Tyler and Panchuk, 1982; Wright et al., 1984; Clott
et al., 1986; Clott and Walker, 1987; Uzzan, 1988; Salunkhe et al., 1989).
Wet processing
When legume seeds are processed for food applications the hulls are
removed because it has been reported that they can contain antinutritional
compounds that can be released during the extraction process (Sosulski
and McCurdy, 1987; Uzzan, 1988). The dehulled seeds are then pin-milled.
It has been found (Gueguen, 1983) that an average flour particle size of
100–150 µm is most suitable for further separation of the components.
The next step of the process is protein extraction, which is carried out at
alkaline pH.
For round-seeded peas, pH 9–10 is most commonly used for extraction,
while for wrinkle-seeded (those lines containing the r gene) peas it is
usually higher (Schoch and Maywald, 1968; Vose et al., 1976; Colonna et al.,
106
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:57:58 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 91
Fig. 4.2. The influence of the number of purification steps on the recovery of pea
starch. Protein content, n; yield of starch, l.
107
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:05 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
92 B. Czukor et al.
1981; Sosulski and McCurdy, 1987; van der Poel et al., 1989; Hoover and
Sosulski, 1991; Wiege et al., 1995; Salunkhe et al., 1989). In a more recently
developed method (Bogracheva et al., 1995; Davydova et al., 1995), this
procedure was carried out at a lower pH (8.5), which is much better for
food applications, because stronger alkaline conditions may result in the
appearance of antinutritional components in the protein products
(Gueguen, 1983; Salunkhe et al., 1989). In the case of the r wrinkled-seeded
peas, some applications require a better separation of starch from the other
components and this is achieved by high-pressure disintegration (Meuser
et al., 1995).
The protein extract, which also contains soluble carbohydrates and
emulsified lipids, is separated from the insoluble fraction and the proteins
are isolated from this extract by acid precipitation or by ultra filtration.
Protein fractions obtained from such procedures are commonly called
protein isolates. The wet protein isolates are then dried. In industry this is
usually carried out using spread dryers at temperatures of more than
100°C. The drying process is relatively quick and so the functional proper-
ties of the proteins are not affected significantly. Freeze-drying, however, is
more appropriate when proteins are isolated for scientific purposes.
The insoluble fraction, which is left after separation of the protein
extract, includes starch, cell wall material, insoluble proteins and the
remaining lipids. The basis for the further separation of these components
depends on differences in their swelling properties. Starch granules have
restricted swelling at room temperatures (Blanshard, 1987; Zobel and
Stephen, 1995), whereas the swelling capacity of cell wall material is much
greater. The swelling properties give rise to a difference in size between
the starch granules and the cell wall particles. The insoluble fraction is
dispersed in a large amount of water and screened through a series of sieves
with pore diameters between 30 and 300 µm (Schoch and Maywald, 1968;
Colonna et al., 1981). The liquid passing through the sieves (termed starch
milk) is mainly a dispersion of starch granules, while the material trapped
by the sieves contains mainly cell wall material. The smaller the diameter of
the sieve pores, the smaller the particles of cell walls that are separated from
the starch milk and the lower the level of impurities. Starch granules are
not uniform in size, the size distribution being dependent on the source of
the starch (Davydova et al., 1995). The minimum diameter of the sieve to be
used is determined by the size distribution of the starch granules. For exam-
ple, 50–60 µm sieves are usually used for producing starch from pea and
faba bean (Colonna et al., 1981). The starch produced by such a method
contains 0.04–0.40% protein and about 0.1–1.0% of lipid as impurities
(Elliasson, 1988; Davydova et al., 1995).
In industry the starches are dried by spread drying machines, which are
specially constructed for this purpose. Starches produced for scientific
purposes, however, usually have an additional wash with water, alkaline or
salt solutions, or with organic solvents such as ethanol or acetone, to reduce
108
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:06 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 93
the level of impurities (Schoch and Maywald, 1968; Davydova et al., 1995).
The starches are then often dried in the open air, in some cases following
a final wash with ethanol, or acetone, to speed up the drying process
(Davydova et al., 1995).
Although the methods described above result in satisfactorily purified
starches, it should be noted that the use of organic solvents might partially
disturb starch granular structure, which in turn may affect the properties. A
diagrammatic representation of wet seed processing is shown in Fig. 4.3.
109
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:11 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
94 B. Czukor et al.
Fig. 4.4. Starch granules from wild-type pea seeds: (A) Using normal light
microscopy. Opposite page: (B) viewed using polarized light, prior to gelatinization
in 0.6 M KCl solution; (C) viewed using polarized light, after heating to the melting
point for B-type crystallites (66°C) in 0.6 M KCl solution.
110
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:15 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 95
111
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:20 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
96 B. Czukor et al.
112
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:23 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 97
granules from a round seeded pea line have been shown to contain both
A- and B-types of polymorph, the B polymorphs being in the centre and the
A polymorphs at the periphery of each granule (Bogracheva et al., 1998).
Since B-type polymorphs melt at a lower temperature than A-type, it is
possible to degrade the B polymorphs from the centre of the granules,
leaving only the A polymorphs intact at the periphery. This process can be
carried out on a microscope stage and observed using polarized light
(Fig. 4.4B and C; Bogracheva et al., 1998). In addition, granules from this
material have 63% double helices (Bogracheva et al., 1998), 33% of which
are arranged in crystallites at a moisture content of about 20% (Cairns et al.,
1997; Bogracheva et al., 1998). A method for determining the polymorph
composition of C-type starch has been developed recently (Cairns et al.,
1997). This method is based on calculations from X-ray diffraction
patterns of the crystalline portions of the starch, using characteristic peaks
associated with either A- or B-type polymorphs (Davydova et al., 1995;
Cairns et al., 1997). Using this method the proportions of A and B
polymorphs in round-seeded peas have been found to be c. 56 and 44%,
respectively.
Analysis of starch from a range of pea mutants (see Chapter 6,
Breeding and Agronomy) has shown that it is possible to genetically
manipulate the physical structure of the starch granules (Table 4.l;
Bogracheva et al., 1995, 1997, 1999; Davydova et al., 1995) It was found that
genes that affect the supply of substrate during starch synthesis (rb, rug3
and rug4) affect the total crystallinity and possibly the content of the A
polymorphs in the granules. On the other hand, genes that directly
affect the synthesis of starch polymers (r, rug5 and lam) increase the B
polymorph content but have little effect on the total crystallinity of the
granules (Table 4.1)
Table 4.1. The characteristics of the granular structure of starch in pea mutants
(Bogracheva et al., 1999).
Amylose Crystallinity
contenta Tp ∆T ∆H
Genotype (% starch) total % %B %A B/A (°C) (°C) (J g−1)
113
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:24 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
98 B. Czukor et al.
114
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:24 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 99
115
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:31 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Fig. 4.8. RVA viscograms of starch suspensions. Starch from: (1) r mutant pea;
(2) maize; (3) rb mutant pea; (4) wild-type pea; (5) potato.
116
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:34 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 101
117
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:41 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Steaming
A simple method for the physical modification of starch within cells is to
treat the legume seed with saturated steam (Kozlowska et al., 1989). Short
periods of this treatment are used during the production of protein
concentrates and isolates from faba bean seeds to improve the sensory
properties of these preparations. The structural changes that take place
during this type of processing can alter the cellular arrangement of protein
and starch within the seed storage tissue (A and B). After this treatment
the isolated starch shows intense amylose leakage and marked granule
deformation during heating in water at high temperatures. The steaming
of seeds also causes changes in the pasting and gel-forming properties. The
reduced swelling of starch granules from steamed seeds gives rise to pastes
with a lower viscosity and a higher temperature is required for amylose
migration from the granule (Fig. 4.10). Starch isolated from untreated
seeds forms a more rigid and more elastic gel compared with starch from
steamed seeds. The internal changes within the granule do not markedly
influence their appearance when viewed in the scanning electron micro-
scope (Kaczymska et al., 1994). Only the denaturation of protein bodies is
visible, the starch granule surface remaining very smooth and unchanged.
The starch properties of pea starches can vary between varieties, the
starch pastes giving different rheological behaviour when heated and
cooled. Starch from pea seeds with hard to cook (HTC)-defect was charac-
terized by having an increased viscosity of paste during heating as well as a
higher value for the G′ modulus on cooling, indicating a high rigidity of
starch gels (Fornal et al., 1995).
118
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:45 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 103
Annealing
Another type of heat treatment used to modify starch is annealing (Hoover
and Manuel, 1994; Jacobs et al., 1995). This entails maintaining the starch
at temperatures lower than the melting temperature (50–75°C) in excess
water. This process results in a decrease in the potential and extent of
amylose leaching. It is accompanied by an increase in the gelatinization
transition temperature, the enthalpy of gelatinization and by a decrease in
the gelatinization temperature range. It has been suggested that these
effects are due to interactions between amylose and the outer branches of
amylopectin, to an increase in double helices and to closer packing of the
crystallities within the granule (Hoover and Manuel, 1994). Annealing also
results in an increase in α-amylase hydrolysis, which may be due to a realign-
ment of starch chains in the amorphous regions of the granules (Minagawa
et al., 1987).
Gamma irradiation
As well as being used in food production, gamma irradiation also can
modify the chemical and physico-chemical properties of starch. Irradiation
has been shown to induce carbonyl derivatives (Raffi et al., 1981a), the
development of acidity (Raffi et al., 1981b) and hydrogen peroxide (Raffi
et al., 1981c) in haricot beans. The effect on starches from maize, manioc,
wheat, potato and rice is to increase the reducing power, the water-soluble
dextrin content and the Brabender viscosity values (Raffi et al., 1981d).
Extrusion
A very promising method for physically modifying starch is extrusion
(Smietana et al., 1996) and high pressure treatment (Kudla and Tomasik,
1992). Extrusion gives products that are free from foreign substances,
that can be used for children and in the development of non-allergenic
formulas and functional foods. High pressure treatment results in some
re-polymerization of dextrin formed during compression and in the order-
ing of starch granule structure into more crystal-like matter. To date, both
of these methods have been used for modifying potato starch but, as yet,
have not been used to modify legume starches.
In the extrusion processing of cereal and legume seeds, as such or in
blends, the transformation of starch and protein determines the properties
of the final product (Schukla, 1996). Dietary fibre, although less affected
during extrusion, can also have a significant effect on textural properties.
The thermodynamic effects during extrusion break hydrogen bonds
in starches, gelatinizing, or even dextrinizing, them in the process. The
required energy input is often a function of starch granule size, the extent
and type of crystallinity and the purity of the starch extract. Low-protein
and high-amylose starches require high inputs of energy to undergo starch
gelatinization. In the case of proteins, the secondary and/or tertiary
119
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:46 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
The most popular methods for modifying starches are based on the use of
chemicals.
Chemical modification can be carried out on three starch states: (i)
in the solid state, where dry starch is moistened with chemicals in a water
solution, air-dried and finally roasted at a temperature of over 100°C; (ii) in
suspension, where the starch is dispersed in water, the chemical reaction is
then carried out in water medium, the suspension is then filtered, washed
and air-dried; (iii) as a paste, where the starch is gelatinized with chemicals
in small amounts of water, the paste is stirred and when the reaction is
completed, the starch is air-dried. The chemicals used for modification
react with the free 2, 3 and 6 hydroxyl groups of the glucose units within the
starch.
The most common chemical modification processes are: oxidation
using sodium hypochlorite, hydrogen peroxide, persulphates or potassium
permanganate; esterification using acetic anhydride, vinyl acetate, ortho-
phosphoric acid, sodium or potassium orthophosphate or sodium tripoly-
phosphate, sodium trimetaphosphate, phosphorus oxychloride or urea;
etherification using ethylene oxide, propylene oxide, monochloroacetic
acid or quarternary amines.
The most important chemically modified starches from an industrial
point of view are the starch-esters and starch-ethers.
To date, legume starches have not been modified on a commercial
scale for non-food applications. There have been laboratory investigations,
however, on chemically modifying legume starches and acetylated starch,
hydroxypropyl starch, cross-linked starch, cationic and grafted starches
have been produced.
Acetylation
The only major study on the acetylation of legume starches has been
carried out on a range of bean (P. vulgaris) varieties, using acetic anhydride
(Hoover and Sosulski, 1985a,b; Vasanthan et al., 1995). X-ray diffraction of
the modified starches indicated that the acetyl groups mainly entered the
amorphous regions of the starch granule. The result of this process was
starches with a decrease in hydrolysis, gelatinization temperatures, enthalpy
of gelatinization, syneresis and in the extent of the viscosity increase during
the holding period at 95°C in a viscosity analyses (Fig. 4.11 and Table 4.2).
There was also an increase in viscosity in the amylose exudation at 95°C. No
120
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:46 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 105
Table 4.2. Syneresis of native and acetylated starch gels after storage at two differ-
ent temperatures for 7 days (Hoover and Sosulki, 1985a,b).
Phosphorylation
Phosphorylation of faba bean starch, using a mixture of mono- and disod-
ium phosphates at 160°C, resulted in a significant increase of swelling
(Soral-Smietana, 1995). Faba bean mono-starch phosphates can bind
components from solution or suspension into a homogeneous mass, which
121
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:51 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Cross-linking
Cross-linking refined starches from lentil, faba bean and pea with phos-
phorus oxychloride decreases water binding capacity, swelling power,
α-amylase digestibility and viscosity at 95°C in the amylograph, but
increases the degree of set-back (Table 4.3; Hoover and Sosulski, 1986).
Cross-linking occurs mainly in the amorphous regions of the starch granule
and hinders amylose exudation. The stable hot paste viscosities of
cross-linked starches would be of value where low pH and high temperature
are employed, during pressure cooking or sterilization, while the low
degree of set-back of pea starch should improve the freeze–thaw stability
and textural quality of frozen foods.
Hydroxypropylation
This chemical modification is based mainly on the addition of propylene
oxide to starch moistened with water containing sodium sulphate, the
mixture being heated and stirred for 24 h at 40°C (Hoover et al., 1988; Kim
et al., 1992). The main effect of this modification on native starch granules
is to produce starch with higher molar substitution (0.12) and a higher
Starch source 1 2 3 4 5 6 7
Native
Lentil 3.4 6.4 10.0 17 21 28 28
Faba bean 2.1 5.8 9.9 16 20 26 27
Field pea 1.9 5.3 9.2 12 19 23 23
Native (gelatinized)
Lentil 42.8 57.8 61.8 66 72 79 79
Faba bean 32.8 51.8 60.8 65 70 72 72
Field pea 30.8 45.8 55.8 62 68 68 70
Cross-linked (ungelatinized)
Lentil 3.2 6.0 10.0 16 20 27 27
Faba bean 2.0 5.5 9.4 15 19 25 26
Field pea 1.8 5.0 8.8 12 18 22 22
Cross-linked (gelatinized)
Lentil 38.8 54.8 58.8 63 69 76 76
Faba bean 29.8 49.8 58.8 63 68 70 70
Field pea 27.8 42.8 52.8 58 64 66 68
122
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:51 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 107
Cationization
Water-miscible solvents such as ethanol, 2-propanol and methanol are used
as the reaction medium for cationization of pea starches (Kweon et al.,
1996). Cationization that results in a degree of substitution of 0.02–0.05
reduces the pasting and gelatinization temperatures, increases the peak
viscosity and set-back on cooling and eliminates synersis after storage at 4°
and −15°C (Yook et al., 1994).
The principal effects of cationization are to promote rapid granule
dispersion at low pasting temperatures and to give a molecular dispersion
of amylose and amylopectin on heating to 95°C. On cooling, the gel
structures are firm and the cationic groups control the realignment of
starch during low-temperature storage.
Investigations on grafting green gram, pigeon pea and garden pea
starches showed that the graft yield of the reaction is less than for cereal
and root starches (Patel et al., 1986). The graft yield in a gelatinized system,
however, is almost independent of starch source. The use of starch graft
copolymers as absorbents depends on their competitiveness on price com-
pared with full synthetic absorbents produced from partially cross-linked
acryl copolymers. (NB. Graft copolymers are specific copolymers, that are
obtained in reactions between macromolecular substances and a substance
of low molecular weight.)
123
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:54 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Many modified starches used in food products are the result of germinating
or fermenting legume seed in vivo, or hydrolysing starches in vitro, using
amylolytic enzymes (Bhat et al., 1983; Revilla et al., 1986a,b; Rodriguez et al.,
1988; Abia et al., 1993; Frias et al., 1998).
Hydrolysis
The first step in enzymic hydrolysis is ‘endocorrosion’, which begins at
the centre of the starch granule. The granule then degrades sequentially
resulting in compartmentalization and subsequent fragmentation. The
surface becomes porous and the granule is then divided into numerous
polyhedral forms of various size (0.4–10 µm). This degradation process has
been reported for starch from lentil (Revilla and Tarango, 1986a,b) and
chickpea (Rodriguez et al., 1988). In general, the susceptibility of starch
granules to modification by amylases depends on the physical structure of
the granules, the amylose content, the degree of polymerization and on the
presence of non-reducing ends on the granule surface (Bhat et al., 1983;
Madhusudhan and Taranathan, 1995).
Germination
Germination induces the release of hydrolytic enzymes, which produce
changes in the physical properties and functionality of seed components.
Starch extracted from faba bean, chickpea and kidney bean before and
after germination was significantly more digestible than those from
ungerminated samples (Table 4.4; Shekib, 1994). Cooking the isolated
starches from both the ungerminated and germinated samples further
increased their digestibility.
Isolated starch
A 33.2 32.3 39.5
B 65.1 62.1 67.2
C 47.2 44.7 49.4
D 84.3 82.1 90.3
Cooked corn starch – – – 90.3
124
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:54 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 109
Fermentation
Fermentation can be taken into consideration as a possible method for
modifying starch properties for food use, for example puddings. This
process has only limited effects on granule swelling and no apparent
effect in solubility. Marked changes, however, were found in the apparent
viscosity of cold pastes and in the intrinsic viscosity (Abia et al., 1993; Nche
et al., 1994; Yadav and Khetarpaul, 1994; Matthews, 1999).
Yadav and Khetarpaul (1994) produced wadi from black-gram dhal
(Phaseolus mungo) and examined the in vitro digestibility of starch, phytic
acid and polyphenols (Table 4.5). When black-gram dhal wadies were
fermented at 25, 30 or 35°C for 12 and 18 h, the improvement in starch
digestibility ranged from 57% to more than 88% over the control value.
Table 4.5. Effect of temperature and fermentation time on in vitro starch and pro-
tein digestibility of wadies prepared from black-gram dhal (Yadav and Khetarpaul,
1994).
125
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:55 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
The most common form of resistant starch (RS) in the diet and the most
important from a technological point of view is retrograded starch (RS III),
126
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:56 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 111
127
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:56 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Soaking
Grain legumes are rarely eaten in a raw state and are usually cooked or
processed first.
Perhaps the simplest method for processing legume seeds is to soak
them in water. This process can reduce the level of reducing and non-
reducing sugars 16–40% (Jood et al., 1988). There is an increase, however,
in in vitro starch digestibility of 17–23% after a 12-h soaking (Bishnoi
and Khetarpaul, 1993). This enhancement of starch digestibility may be
attributed to the loss of antinutritional factors such as phytic acid and
polyphenols, which inhibit the activity of α-amylase (Deshpande and
Cheryan, 1984). Conversley, it has been suggested that prolonged soaking
of intact peas may allow the mobilization of phenolics, which are known
to interfere with starch digestion from the seed coat to the cotyledons
(Deshpande and Salunkhe, 1982).
Soaking in water and NaHCO3 solution also significantly reduces
the levels of stachyose, verbascose and raffinose. The reduction is usually
higher in NaHCO3 solutions than in water and can account for 46–100% of
the α-galactoside content (Jood et al., 1985; Vijayakumari et al., 1996). Only
1–10% of these losses can be explained by leaching into the soaking
solution, the remainder being due to hydrolysis by α-galactosidase released
by the imbibed seed (Vidal-Valverde et al., 1992a).
In general, soaking is not used by itself, but in combination with
germination, cooking or autoclaving.
128
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:57 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 113
decrease of sucrose after 90 min, while the level of raffinose, stachyose and
verbascose increased (Revilleza et al., 1990). This increase could be attrib-
uted to hydrolysis of oligosaccharides bound to proteins or other macro-
molecules, or to the hydrolysis of high molecular weight polysaccharides.
Steam-heated legumes are rich in resistant starch, while high pressure
steaming and dry pressure cooking decreases the total and the available
starch, and stabilizes or decreases the resistant starch level (Tovar and
Melito, 1996; Periago et al., 1997). There is an increase in the total non-
starch polysaccharide content during cooking, the content of soluble non-
starch polysaccharides generally increasing and the insoluble non-starch
polysaccharide content decreasing (Periago et al., 1997).
Extrusion
Depending on the extrusion conditions (temperature, moisture content,
screw speed), the loss of total sugar and α-galactoside content can be about
30% and 50–60%, respectively, while the starch digestibility can increase
significantly (Tuan and Phillips, 1991; Borejszo and Khan, 1992). Extrusion
cooking marginally decreases the total content of dietary fibre of peas at
168°C, but the content of resistant starch increases significantly from about
1.5% to about 3.3% of the total starch (Berghofer and Horn, 1994).
Dry roasting
At high temperatures, dry roasting results in the complete reduction of the
oligosaccharide content in the hyacinth bean after 2 min, although the
levels of sucrose and the oligosaccharides were higher after a roasting
time of less than 0.5 min (Revilleza et al., 1990). The effect after 2 min was
probably due to a non-enzymatic browning reaction, oxidation of sugars or
to pyrolysis (Fig. 4.13). Contrary to the effect of other processes, frying and
roasting considerably reduces starch digestibility of legumes (Kelkar et al.,
1996). Frying significantly reduces the sucrose content of legume seeds,
probably because of the composition of the frying medium (Jood et al.,
1985).
Freeze-drying
There is no apparent effect of freeze-drying on the sugar, starch and pectin
contents of green beans (Oruna-Concha et al., 1996), although there was a
slight decrease in the digestibility of starch in the tepary bean (Abbas et al.,
1987).
Microwaving
Repeated microwave treatments decrease the total dietary fibre content of
green bean, primarily because of losses in the soluble dietary fibre content
(Svanberg et al., 1997).
129
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:58:57 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Fig. 4.13. The effect of dry roasting on the soluble sugars of hyacinth bean
(Lablab purpureus; Revilleza et al., 1990).
Germination
Germination is one of the best known methods for decreasing the
α-galactoside content and increasing the starch digestibility of legumes.
During germination there are significant decreases in the total soluble
carbohydrate content and in the total starch content, and increases in
the reducing and non-reducing sugar content and in the in vitro starch
digestibility. In addition, this process totally eliminates the α-galactosides
(raffinose, stachyose, verbascose) (Jood et al., 1988; Revilleza et al., 1990;
Trugo et al., 1990; Vidal-Valverde et al., 1992a; Vidal-Valverde and Frias, 1992;
Bishnoi and Khetarpaul, 1993; Shekib, 1994; Urooj and Puttaraj, 1994;
Urbano et al., 1995; Kelkar et al., 1996). The extent of the changes is mainly
determined by the germination conditions (Nnanna and Philips, 1988).
Germination significantly increases the resistant starch content
resulting in an increase in the total dietary fibre content, the soluble
fraction decreasing and the insoluble fraction increasing significantly
(Veena et al., 1995).
Fermentation
Natural fermentation enhances protein digestibility and eliminates par-
tially or completely the antinutritional factors. The effect of fermentation
on starch digestibility has been studied in Bengal gram, cowpea and green
gram (Urooj and Puttaray, 1994). The seeds were soaked in water for 3 h,
drained, ground and allowed to ferment for 4 h at room temperature. After
fermentation the meal was steam-cooked for 10 min. The fermentation
treatment reduced the total soluble carbohydrate, the starch and the
130
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:01 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
Processing 115
131
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:02 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
132
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:03 A3867: AMA: Hedley: First Proof:30-Oct-00 4
Color profile: Disabled
Composite Default screen
5
Seed
J. Górecki
Physiology
et al. and Biochemistry
The legume seed can be separated into three major tissues – testa, endo-
sperm and embryo – which in turn can be further divided into the two
cotyledons and the embryonic axis (Fig. 5.1). The testa (Latin for ‘shell’),
seed coat or hull is a maternal tissue that surrounds the embryo and
attaches the seed to the pea pod via the stalk-like funicle. During early
development, the testa, acts as a nurturing tissue, distributing nutrients
from the mother plant to the developing embryo. Early in development this
is by diffusion via the endosperm and later on by direct contact with the
embryo. Sucrose, the main form of imported carbohydrate in legume seeds
(Pate et al., 1974; Fellows et al., 1978; Patrick and McDonald, 1980; Schmitt
et al., 1984), is unloaded from the vascular bundles within the testa. It
is then thought to be transported symplastically through the seed coat
(Patrick and Offler, 1995). Its import into the embryo is regulated by
133
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:04 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
134
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:06 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
135
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:07 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
Table 5.1. Stage of development descriptions for soybean (adapted from Fehr
et al., 1971, and Lowell and Kuo, 1989).
136
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:07 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
137
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:08 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
et al., 1995a,b; Hedley et al., 1996; Lloyd et al., 1996a,b; Casey et al., 1998;
Craig et al., 1998; Harrison et al., 1998; Craig et al., 1999). Many of these
more recent studies have been able to utilize the range of starch mutants
described in Chapter 7. They have also provided information for studying
the growth and development of seeds from other grain legume species, in
particular lentil (Bakhsh et al., 1991, 1992; Frias et al., 1994b, 1996d).
Table 5.2. Soluble carbohydrates in the axes and cotyledons of different grain
legume species (mg g−1 dry matter).
(a) Axes
Soybeanb 111.3 26.3 131.4 trace 1.7 288.6
Peac 69.7 19.6 65.5 45.6 5.7 200.5
Yellow lupina 20.3 23.9 199.6 59.5 5.8 248.0
Faba beanb 57.0 15.0 65.3 99.4 0.3 243.6
Lentilb 39.2 5.4 92.4 33.7 10.8 236.5
Pigeon peab 61.5 13.4 35.7 90.1 4.8 237.1
Cowpeab 45.3 12.2 110.0 11.2 0.9 184.7
Chickpeab 36.1 22.9 35.1 1.8 22.2 171.5
(b) Cotyledons
Soybeanb 76.1 11.8 43.5 trace 2.6 141.0
Peac 26.6 4.9 13.7 34.8 2.4 80.1
Yellow lupina 7.7 8.4 48.3 23.6 4.3 110.0
Faba beanb 19.3 3.4 11.1 52.5 0.0 88.4
Lentilb 11.5 2.4 25.4 18.0 33.0 85.2
Pigeon peab 18.8 9.4 16.5 34.6 11.4 95.4
Cowpeab 15.2 3.3 55.2 10.8 2.3 87.7
Chickpeab 20.6 4.3 26.7 1.0 34.5 94.2
aGórecki et al. (1997).
bHorbowicz and Obendorf (1994).
cLahuta (unpublished).
138
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:08 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Table 5.3. Distribution of soluble carbohydrates in dry seeds in various legumes (mg g−1 defatted meal) (Kuo et al., 1988).
Pea cv. Little Marvel 62.3 11.6 32.3 19.1 63.0 125.3
Soybean cv. Williams 82 72.7 12.6 43.4 trace 56.0 120.2
cv. Amsoy 71 64.2 11.6 41.0 trace 52.6 125.3
Cowpea 25.9 3.7 46.4 3.6 53.7 79.6
Mung bean cv. Berken 13.9 3.9 16.7 26.6 47.2 61.1
Lima bean cv. Fordhook 36.0 6.9 30.3 trace 37.2 73.2
139
Common bean cv. Top Crop 19.4 2.5 34.3 trace 36.8 56.2
cv. Blue Lake 29.0 2.2 28.1 trace 30.3 59.3
red kidney bean 21.5 3.1 31.6 trace 34.7 56.2
Pole bean cv. Kentucky 26.7 4.3 26.2 trace 30.5 57.2
5
Groundnut cv. Florunner 81.0 3.3 9.9 trace 13.2 94.2
Monosaccharides
The monosaccharides found in legume seeds are often involved as transi-
tory intermediates in the synthesis of higher polymers of carbohydrates.
They are also often found in the phosphorylated form. Free mono-
saccharides are most readily detected during early seed fill and are rapidly
utilized as development continues. Often monosaccharides, such as
fructose, glucose and galactose, are found only in trace amounts in mature
seeds (Horbowicz et al., 1995; Sun and Leopold, 1995). In soybean, how-
ever, the amounts can be relatively high early in development (Yazdi-
Samadi et al., 1977), while in mature lucerne seeds no monosaccharides
have been detected (Horbowicz et al., 1995). It has been suggested that it is
advantageous to the plant if the concentrations of the monosaccharides are
reduced. This is because the reducing nature of these compounds has been
implicated in the Maillard reaction, which causes oxidative stress through
the formation of free radicals, particularly after the seed has germinated
(Sun and Leopold, 1995). Once a seed has germinated the amounts of
galactose still remain almost undetectable, despite the removal of the
galactose moieties from oligosaccharides. This is due to the high levels of
galactokinase converting any free galactose to a safer phosphorylated form
during germination (Dey, 1985).
Disaccharides
The most abundant disaccharide found in seeds is sucrose, which is
the principal translocated photoassimilate (Lin et al., 1984; Patrick and
McDonald, 1980; Lichner and Spanswick, 1981; Schmitt et al., 1984;
reviewed in Patrick and Offler, 1995). Early in the development of pea
seeds it can reach high levels. As the seed develops, however, the sucrose
content falls as it is utilized for dry matter accumulation, so that by the time
of maximum fresh weight of a pea seed it consists of around 8%, or 20 mg
per whole seed (Harrison, 1996).
Oligosaccharides
The most commonly occurring oligosaccharides found in plants are those
based upon α-galactosyl derivatives of sucrose (reviewed in Dey, 1990).
These compounds are almost ubiquitous throughout the plant kingdom
and rank second only to sucrose as the most abundant soluble carbo-
hydrates. These oligosaccharides can comprise from 2 to 13% of legume
seed dry weight and they are believed to play an important role in seed
storability and in protecting the seed from desiccation stress (Obendorf,
1997; Sun, 1997).
The galactosyl moiety of these compounds can be linked to either the
fructose or the glucose moieties of sucrose. Depending upon the bonding
of the molecules to each other, they will form different families of oligo-
saccharides (Table 5.4).
140
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:10 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
141
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:11 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
Fig. 5.2. Biochemical pathway for some of the major α-galactosides and cyclitols.
142
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:18 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
143
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:19 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
The role of starch in the plant has been reviewed by Morrison and Karkalas
(1990) and in seeds by Sivak and Preiss (1995). The role of starch in pea
seeds has been reviewed by Smith and Denyer (1992) and the biosynthesis
144
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:23 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
Fig. 5.4. Proposed pathways of synthesis of the main cyclitols and methyl-
cyclitols present in legume seeds.
145
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:30 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
have very little internal membrane (Journet and Douce, 1985). In contrast,
the starch-storing plastids of developing pea embryos (and probably other
grain legumes, which accumulate starch, as a main storage product) are
derived directly from chloroplasts and retain chloroplast-like characteris-
tics throughout their development. Developing pea embryos contain
chloroplasts (located primarily on the outer edge of cotyledons), which
store little or no starch (Smith et al., 1990).
The starch accumulation in developing faba bean seeds has been
reviewed by Weber et al. (1995b, 1997). During the cell-division phase of the
faba bean embryo (15–18 days after flowering; DAF) starch is mainly found
in two layers of the seed coat, in the hypodermal and chlorenchymal cells
and in the outer parenchymal cells. When storage-product synthesis begins
in the cotyledon (about 25 DAF), starch is deposited as single granules in
the cells of the adaxial region of cotyledons. Later, starch deposition
spreads from the adaxial cells to the periphery. During this process the
quantity and size of the starch granules increases in the cells of the adaxial
region and new granules appear in the abaxial cells. During the cell-elonga-
tion phase (about 30–35 DAF), starch is also present in the axis cells. At no
developmental stage is starch found in the palisade cell layers of the seed
coat, the peripheral cells of the abaxial zone, or in the provascular and
calyprogenous cells.
Developing seeds of pea, faba bean and common bean accumulate
starch up to full seed maturity (Meredith et al., 1988; Lahuta et al., 1995).
In seeds, which accumulate oil as reserve material (e.g. soybean), starch
synthesis occurs in the cotyledon growth phase (about 35–40 DAF) and
decreases to trace amounts during seed maturation (Monma et al., 1991).
146
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:31 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
Zygotic seeds
Seeds of most plant species exhibit the ability to withstand desiccation, in
many cases achieving water contents of less than 5–10% on a fresh weight
basis. These seeds are termed orthodox and can be stored for many years
under dry conditions. Orthodox seeds, which include those from legumes,
are not capable of withstanding desiccation at early stages of development.
The ability to tolerate desiccation is acquired during later stages of seed
development and is lost after germination. It is believed that the acquisition
of desiccation tolerance is developmentally controlled (Galau et al., 1991;
Bewley and Oliver, 1992; Kermode, 1997). Galau and co-workers (1991)
suggest that desiccation is acquired before maturation drying at the ‘post
abscission stage’, when the vascular connection between the seed coat and
the parent plant is lost.
Acquisition of desiccation tolerance in maturing seeds involves several
components including the accumulation of a special group of proteins
(Blackman et al., 1991; Galau et al., 1991; Bewley and Black, 1994; Vertucci
and Farrant, 1995), non-reducing sugars and/or galactosyl cyclitols (Koster
and Leopold, 1988; Horbowicz and Obendorf, 1994; Obendorf, 1997), free
radical scavenging systems (Senaratna et al., 1985; Koster and Leopold,
1988; Lowell and Kuo, 1989; Hendry, 1993; Leprince et al., 1993; Finch-
Savage et al., 1994) and abscisic acid (Bartels et al., 1988; Anandarajah
and McKersie, 1990; Blackman et al., 1991; Vertucci and Farrant, 1995).
The role of sugars and proteins in seed desiccation tolerance has been
extensively studied. With regard to proteins, in desiccation tolerant seed a
147
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:32 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
148
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:32 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
Fig. 5.5. Contents of soluble sugars, cyclitols and galactosyl cyclitols in the axis of
maturing yellow lupin seeds (Górecki et al., 1997).
149
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:39 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
seeds coincides with an increase in the RFO and the acquisition of desicca-
tion tolerance, while in pea only the RFO accumulate (Fig. 5.6; Górecki
et al., 1997). Galactopinitol A, galactopinitol B and fagopyritol B1 accumu-
late in the embryonic axis tissues of developing soybean seeds in association
with desiccation tolerance and in parallel with stachyose accumulation
(Obendorf, 1997; Obendorf et al., 1998). Immature soybean seeds accumu-
late galactopinitols during slow drying (Obendorf et al., 1996). Galacto-
pinitol A, galactopinitol B and fagopyritol B1 accumulate in parallel with
stachyose in axis and cotyledon tissues during in vitro growth of embryos.
Evidence, therefore, supports the suggestion that galactosyl cyclitols in
seeds may enhance the physiological role of α-galactosides.
In some seeds that have very low RFO levels, galactosyl cyclitols may
replace the role of the α-galactosides in the acquisition of desiccation
tolerance. Desiccation-tolerant buckwheat seeds, for example, accumulate
150
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:46 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
Somatic embryos
Oligosaccharides and galactosyl cyclitols seem to play an important role in
the acquisition of desiccation tolerance of somatic embryos. These are used
as synthetic or artificial seeds for the propagation of high-value plants, or as
a plant breeding tool in the development of new cultivars (see Chapter 6).
Somatic embryos have been obtained for more than 150 species of
important agricultural crops including legumes, cereals and grasses, and
are genetically identical to the donor plant. Lucerne somatic embryos
deposit storage proteins and carbohydrates and acquire desiccation toler-
ance (Lai and McKersie, 1994). Morphologically, however, lucerne somatic
embryos do not have fully developed cotyledons and lack an endosperm
and testa. Also, instead of galactomannan being the endospermic carbohy-
drate reserve, somatic embryos contain starch, sucrose and raffinose (Lai
and McKersie, 1994; Horbowicz et al., 1995).
Unlike zygotic seeds, somatic embryos have elevated levels of sucrose
and do not accumulate D-pinitol or its galactosyl derivatives (galactopinitol
A, galactopinitol B, ciceritol or trigalactopinitol A). Lower levels of
stachyose accumulate during the maturation of somatic embryos. During
desiccation, however, stachyose increases in somatic embryos to levels
similar to those found in mature seeds. The decrease in sucrose concentra-
tion and the increase in stachyose during drying results in a decline in
the sucrose : oligosaccharide ratio. Also, reducing sugars decrease during
desiccation of somatic embryos (Horbowicz et al., 1995). Except for the lack
of pinitol and galactosyl pinitols, changes in soluble carbohydrates during
the maturation and desiccation of lucerne somatic embryos are similar to
zygotic seeds and are associated with desiccation tolerance.
151
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:46 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
152
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:47 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
plant maturation to the content and composition of the RFO and other
soluble carbohydrates in seeds. It has been shown that white lupin seeds
matured at 28°C accumulate only 53–70% as much dry matter as seeds
matured at 13°C (Górecki et al., 1996). These changes were accompanied
by only minor changes in the RFO. Pinitol and the galactose-containing
pinitols, however, were more than doubled by seed maturation at 28°C, but,
collectively, these compounds make up less than 10% of the total soluble
carbohydrates. The effect of maturation temperature on the composition
of soluble carbohydrates in yellow lupin seeds has also been studied
(Górecki et al., 1996). In this species, seeds matured at 18°C had more
than twice the amount of stachyose and verbascose compared with seeds
matured at 25°C. From these limited experiments it can be suggested that
the RFO and galactosyl cyclitols may play some role in temperature stress
response on maturing seeds.
In general, seeds attain their maximum viability and vigour after the final
stage of maturation and then, during storage, they gradually deteriorate
until death. The decline of seed quality during storage is expressed firstly as
a decrease in the growth rate of germinating embryonic axes (vigour) and
secondly as a loss of the ability to germinate (viability). Seed quality loss
during storage is associated with increased membrane permeability and
many distinct biochemical changes. These include lipid peroxidation,
chromosome aberration and damage to DNA, changes in RNA and protein
synthesis, reduction in respiration and changes in enzymes and reserve
substances (Bewley and Black, 1985).
It has been observed that the content of soluble carbohydrates declines
with increased storage duration (Taufel et al., 1960; Yaklich, 1985; Petruzelli
and Tarano, 1989; Kataki et al., 1997; Zalewski and Lahuta, 1998). Similarly,
there is a positive correlation between the decline in RFO content and the
reduction of seed longevity (Bernal-Lugo and Leopold, 1992). The deple-
tion of soluble sugars may result in the limited availability of respiratory
substates for germination (Edje and Burris, 1970). Other possibilities are
that a depletion in the oligosaccharide content may reduce the protective
effects of sugars on the structural integrity of membranes, or may reduce
the ability of the seed to maintain a glassy state, resulting in a non-
crystalline liquid state of high viscosity (Bruni and Leopold, 1991).
In dry legume seeds, the soluble carbohydrates comprise mainly
sucrose, together with different quantities of oligosaccharides, in particular
raffinose, stachyose and verbascose. Sucrose is exceptionally effective in
protecting membrane integrity in dry systems (Crowe and Crowe, 1986) as
well as being one of the best vitrifying sugars (Green and Angell, 1986). As
mentioned above (see Section 5.3.3), raffinose and other oligosaccharides
153
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:47 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
Degradation of starch
Reserve starch is deposited in amyloplasts within the embryonic axis and
cotyledon cells. During seed maturation the membranes of the amyloplasts
appear to disintegrate, exposing the starch granules directly to the cyto-
plasm of the cells (Harris, 1976; Halmer, 1985). Starch breakdown in
legume cotyledons commences shortly after imbibition, but the rate of
hydrolysis differs between species and varieties. The spatial pattern of
starch degradation within tissues also varies (Ziegler, 1995). In Phaseolus
cotyledons this process progresses from the central region, in peas from
the outer cotyledon face inwards and in mung bean from the inner face
outwards (Bewley and Black, 1978). Surface pores on the starch granules
may facilitate the selective penetration of degrading enzymes, since
granules appear to be broken down primarily from within (Harris, 1976).
Since starch granules are effectively insoluble, breakdown occurs in three
phases (Preiss and Levi, 1980). Firstly, the granules are reduced to large
154
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:48 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
155
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:48 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
156
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:49 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
Mature seeds usually contain isoforms of this enzyme that differ in activity
and molecular mass (Pridham and Dey, 1974).
In developing seeds, the activity of α-D-galactosidase increases during
the period of intensive RFO synthesis, reaching its highest level at full
maturity. This increase may result from the structural transformation of
isoenzymes, which leads to changes in their specific activities (Pridham and
Dey, 1974). In the maturing embryo, the synthesis of the oligosaccharides
and α-D-galactosidase probably occur in different cellular compartments.
In pea cotyledons, α-D-galactosidase has been localized in cell vacuoles that
are storing the lectin precursors (Harley and Beavers, 1989). In the cells of
soybean cotyledons, α-D-galactosidase occurs in cisterns of the Golgi
apparatus and it may be deposited in protein bodies (Hermann and
Shannon, 1985). A similar observation has been made in faba bean (Datta
et al., 1985). A direct role has been established for α-D-galactosidase and
α-D-mannosidase in the hydrolysis of glycoproteins and storage galactosides
in the cotyledons of germinating narrow-leafed lupin (Plant, 1984). In yel-
low lupin seeds the activity of α-D-galactosidase increases only at the begin-
ning of germination (Login et al., 1995), when the accumulated RFO and
galactosyl cyclitols undergo complete decomposition (Górecki et al., 1997).
While the role of α-D-galactosidase in the hydrolysis of saccharides and
glycoproteins in germinating seeds is understandable, the role, if any, of
this enzyme in non-germinating (stored) seeds remains to be elucidated.
The seeds of various species appear to use the RFO as part of their storage
material and it has been observed that the oligosaccharide content
decreases with increased storage duration (Taufel et al., 1960; Yaklich, 1985;
Bernal-Lugo and Leopold, 1992; Horbowicz and Obendorf, 1994).
Similarly, there is a positive correlation between the decline in RFO
content and the depression of seed longevity.
157
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:50 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
by Górecki et al. (1997) and Górecki and Obendorf (1997). In these studies
it was found that radicle tissues are often more sensitive to desiccation than
hypocotyls. Pea root tissues lost desiccation tolerance during the first 36 h
of germination, while 80% of epicotyls survived slow drying treatment
and 40% survived fast drying treatment of seedlings, for up to 96 h after
imbibition (Fig. 5.7). During desiccation, sucrose levels increased five-
to tenfold in root and hypocotyl tissues and even more in epicotyls.
Glucose and fructose increased during germination and remained elevated
after drying. These changes in saccharides reflected the mobilization of
Fig. 5.7. Pea seedling germination (A) and length (B) of axis, radicle and epicotyl
as a function of hours after imbibition. Desiccation tolerance (C, E) and length
(D, F) of radicle and epicotyl at 6 days after rehydration of (C, E) fast-dried or (D, F)
slow-dried seedlings as a function of seedlings as a function of seedling age when
dried (Górecki and Obendorf, 1997).
158
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:57 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
159
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:57 A3867: AMA: Hedley: First Proof:30-Oct-00 5
Color profile: Disabled
Composite Default screen
6
N. Kuchuk et al.
Biotechnology
Biotechnology 6
Editor: Nickolay Kuchuk
Contributors: Miroslav Griga, Georgina
Kosturkova, Nickolay Kuchuk and Mladenka
Ilieva-Stoilova
And he gave it for his opinion, that whoever could make two ears of corn
or two blades of grass to grow upon a spot of ground where only one
grew before, would deserve better of mankind, and do more essential
service to his country than the whole race of politicians put together.
Gulliver’s Travels, ‘A Voyage to Brobdingnag’, ch. 7 (1726)
Jonathan Swift (1667–1745), Anglo-Irish poet and satirist
6.1 Introduction
This chapter concentrates on the biotechnological techniques developed
in the fields of plant cell tissue culture and genetic engineering.
The cultivation methods for explants and single cells (protoplasts) and
for plant regeneration in vitro are described as basic approaches that allow
valuable genotypes to be propagated as well as to produce fertile plants
from somatic cells. In vitro somaclonal variation and cell selection is
considered as a new source of diversity for plant breeding. Methods for
genetic transformation and for the production of transgenic grain legumes
are summarized to give an idea about ‘the state of art’ of this technology.
Hopefully, this information will promote a better understanding of the
current opportunities and future prospects of plant biotechnology, as well
as the possibilities for the future manipulation of carbohydrate metabolism,
content and composition in grain legume seeds.
161
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:58 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
The ability of plant cells to express their entire genetic information and
regenerate whole plants (i.e. totipotence), is the basis of their development
in controlled in vitro conditions and to the establishment of biotechnologi-
cal techniques and methods for genetic manipulation.
In vitro plant cells from excised tissues (explants) might undergo de-
differentiation and re-differentiation following several developmental path-
ways, depending on the culture environment. Unorganized cell division
and growth can be stimulated, leading to the formation of callus and
suspension cultures. Morphogenesis can be induced in meristem, callus,
suspension and protoplast cultures, leading to the formation of somatic
embryos, organs (shoots and roots) and a whole plant.
Undifferentiated cell growth in callus or suspension cultures can be
adequate for the purposes of some biotechnology processes (like secondary
metabolite production, and some investigations in physiology, genetics,
cytology, biochemistry, etc.; see Section 6.6). Regeneration of plants, how-
ever, is essential for the application of recent advances in biotechnology,
especially those concerning genetic engineering for plant improvement.
Realization of regeneration capacity in vitro depends on knowledge of
the requirements for stimulation of the morphogenic response (Halperin,
1986). Unfortunately, information on genetic, epigenetic and physiological
status of the explant is still limited and in practice the general approach is
to find out the most appropriate chemical or physical stimuli to provoke
totipotency of the cell. Up to now, this process has been mainly empirical
and the formulation of strict rules and general protocols has not been
possible.
The establishment of in vitro cultures and the induction of morpho-
genesis in grain legumes has proved more difficult in comparison with
most Solanaceae and Brassicaceae species. For a long time recalcitrance
in regeneration has been the largest obstacle for genetic manipulation.
Several important observations have led to the development of efficient
regeneration systems. These have focused on the role of the genotype,
the explant, the application of relatively high auxin concentration for
induction of somatic embryogenesis and the use of powerful cytokinins
for multiple shoot proliferation. Definite success has been achieved in the
regeneration of soybean and pea and this success has been immediately
applied to genetic manipulation (Barwale and Widholm, 1990; Griga and
Novák, 1990; Christou, 1992; see Sections 6.2 and 6.4). The development
of efficient in vitro methods and plant regeneration protocols for other
large-seeded legumes has made significant progress in the last two decades
and organogenesis/somatic embryogenesis has been reported at least
162
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:58 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 147
Species References
163
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 14:59:59 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Species References
C. arietinum Altaf and Ahmad (1990); Malik and Saxena (1992b); Brandt
and Hess (1994); Polisetty et al. (1997); Santangelo et al.
(1997)
G. max Parrott et al. (1992); Sharma and Kothari (1994); Kaneda
et al. (1997)
L. culinaris Malik and Saxena (1992b); Polanco and Ruiz (1997)
L. albus Sator (1990); Rybczynski and Podyma (1993b)
P. vulgaris Mohamed et al. (1992); Herselman and Mienie (1995)
P. sativum Jackson and Hobbs (1990); Kosturkova et al. (1997)
V. faba Mohamed et al. (1992)
164
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:00 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 149
In the early 1970s, Gamborg et al. (1974) and Kartha et al. (1974) first
reported shoot regeneration from callus tissue and from apical meristem,
respectively. Initial attempts to induce organogenesis in callus cultures
studied the response of different plant tissues, genotypes and media com-
position. Gamborg et al. (1974) first induced shoot formation in callus from
macerated apical meristems grown on media containing 1 µM naphthalene
acetic acid (NAA) and 0.2–5.0 µM 6-benzyl amino purine (BA), the latter
being important for vigorous shoot formation. Shoots were formed de novo
exogenously on the callus. While most of the calli produced one or more
shoots, root formation was poor and did not occur regularly.
Malmberg (1979) used epicotyl sections to obtain callus on MS
media (Murashige and Skoog, 1962) supplemented with BA and NAA. He
observed that organogenic ability was genotype dependent (six out of 16
lines responded) and decreased with prolonged culture. Root formation
was induced by NAA but, as previously reported, it was not satisfactory.
Atanassov and Mehandjiev (1979) observed that the developmental
stage of the explant influenced the efficiency of callogenesis, the organo-
genic response being observed only for 20–22-day-old embryos. Bud
formation was stimulated by 0.5 mg l−1 BA and could be maintained
for eight subcultures, but rooting of the regenerants was problematic.
Cytological analysis showed that 13% of the newly formed shoots had
mixoploid cells at the vegetative apex.
An effective system of de novo regeneration from callus derived from
immature leaflets (0.9–1.8 mm) was developed by Mroginski and Kartha
(1981). The combination of BAP (10 µM) and NAA (10 µM) was the best
for callus induction and subsequent shoot regeneration.
Hussey and Gunn (1984) succeeded in obtaining vigorously grown
callus with superficial meristems, using plumules that continuously regen-
erated shoots over a period of nearly 3 years. Callogenesis was induced on
MS medium supplemented with 1 mg l−1 BA and 4–8 mg l−1 indole-3-acetic
acid (IAA), and was maintained by reducing indole-3-butyric acid (IBA) to
0.25 mg l−1. The in vitro response differed between genotypes and only two
(cvs. Puget and Upton) out of five varieties maintained regenerative callus.
During the first year of callus growth the plants were diploid and mostly
morphologically normal. Many of the shoots regenerated after 2 years,
however, showed considerable morphological variation and difficulties in
rooting.
Natali and Cavalini (1987a,b, 1989) examined some factors affecting
rhizogenesis in cultures obtained from macerated vegetative apices or
immature embryos. The highest frequency of rooting was achieved at
half strength MS medium supplemented with 2 mg l−1 IBA. Difficulties
in rooting seemed to affect only shoots regenerated from callus, but not
formed from meristematic tissues. Grafting of regenerated plantlets on
165
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:00 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
166
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:01 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 151
167
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:01 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
168
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:02 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 153
medium containing 5–50 µM TDZ. When these seedlings were exposed for
a prolonged time (3–4 weeks) to the same medium, numerous shoots
emerged de novo from the base, or from the upper part, of multiple shoots.
Bohmer et al. (1995) developed a protocol for high frequency shoot
induction and plant regeneration from protoplast derived pea callus, using
TDZ as the key factor in the system.
6.2.6 Recent studies to produce more efficient, fast and reliable systems
for regeneration
169
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:02 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Explant
Among the various tissues used as initial material for in vitro cultures it
seems that whole or parts of (cotyledons or embryonic axes) immature
zygotic embryos are preferable as embryogenic and organogenic explants.
Cotyledonary nodes from seedlings and meristematic tissues are suitable
material for the induction of adventitious bud formation and micro-
propagation, while young leaflets, epicotyl and hypocotyl have been used
less. The developmental stage of the explants is very important as it can
170
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:03 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 155
Growth regulators
Auxins and cytokinins, or substances with a similar structure generally
regulate in vitro development and plant regeneration. Choosing the right
growth regulators and the correct concentration seems to depend on the
explant, its developmental stage and on the genotype, leading to a variety
of regeneration protocols. Some general observations, however, can be
made. The presence of a high concentration of auxin is essential for
somatic embryogenesis, but the type of auxin can differ: 2,4-D and picloram
being cited as superior; NAA is less efficient for embryo induction, but
is necessary for embryo conversion; NAA and IAA, depending on the con-
centration, can induce callogenesis or rhizogenesis. BA alone, or in combi-
nation with an auxin, has been the most commonly used cytokinin for
induction of organogenesis and shoot proliferation. Zeatin is very efficient
in shoot induction, but TDZ seems to be superior as well as being efficient
in the embryo-conversion process (Griga, 1998).
Genotype
Recalcitrance in grain legumes could be the result of a long history of
inbreeding and selection, leading to a reduction in genetic variability.
Screening a large number of genotypes could help to discover those with
a better response to organogenesis and/or embryogenesis. A correlation
between embryogenic and organogenic capacity in different responding
cultivars, however, is not always observed. With regard to rooting frequency,
the data are also contradictory. There are reports, however, that these
processes may be under genetic control (Althers et al., 1993; Bencheikh and
Gallais, 1996).
171
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:03 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Definition
The term protoplast refers to all components of a plant cell excluding the
cell wall. The plant cell wall consists of three primary components, cellulose
(25–50%), hemicellulose (average 50%) and pectin substances (about
5%). Cocking (1960) first used hydrolytic enzymes for digesting the cell
wall of tomato root tips to release plant protoplasts. This method allows the
quick isolation of an indefinite number of uniform plant protoplasts from
any type of plant tissue from any plant species.
172
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:04 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 157
Interest towards protoplasts from leguminous species dates from the early
1970s. Most investigations were carried out on soybean and pea, which are
the most important grain legume species. Some of the achievements
are presented in Table 6.2. Grain legumes proved to be recalcitrant,
however, which has made success in regenerating plants from protoplasts
difficult. Different sources for producing protoplasts (cell suspension,
leaf mesophyll, hypocotyl, epicotyl, etc.) and various culture conditions
Pea
Cell division Landgren (1976); Jia (1982)
Callogenesis Constabel et al. (1973); Gamborg et al. (1975); von Arnold
and Eriksson (1976): Kuchuk (1989)
Embryogenesis, Puonti-Kaerlas and Eriksson (1988); Lehminger-Mertens and
organogenesis Jacobsen (1989a); Ochatt et al. (1998)
Plant regeneration Lehminger-Mertens and Jacobsen (1989b); Boehmer et al.
(1995); Sanago et al. (1996), Ochatt et al. (1998)
Soybean
Cell division Kao et al. (1970); Gamborg et al. (1983); Lu et al. (1983);
Tricoli et al. (1986); Hammat and Davey (1988)
Callogenesis Zieg and Outka (1980); Xu et al. (1982); Oelck et al. (1983);
Kuchuck (1989)
Plant regeneration Myers et al. (1989); Guo (1991); Dhir et al. (1991, 1992); Lu
et al. (1993)
173
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:05 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Plant protoplasts lack cell walls and are, therefore, a good experimental sys-
tem for various types of study, such as genetics (using somatic hybridization
and genetic transformation), physiology, cytology, biochemistry and other
fields of biological science. In particular, they are an excellent experi-
mental system for basic studies of cell wall regeneration, cell division,
membrane fusion, membrane transport, virology, endocytosis and transfer
of organelles (Fowke and Constabel, 1985; Fowke and Wang, 1992). The
sections below cover the role of carbohydrates in different cell processes
of grain legume protoplasts and present possibilities on how protoplast
systems may be exploited as tools for carbohydrate research.
174
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:05 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 159
175
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:06 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
176
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:06 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 161
seedlings (leaf, shoot tip, epicotyl, hypocotyl and root tip) and observed the
accumulation of starch after 1 week. Starch accumulation was not observed
in protoplasts derived from shoot tips or from the first lateral shoots
originating from the cultured embryonic axis minus the cotyledons. In
these cultures, particularly homogenous meristematic cells and sustained
protoplast division was achieved.
In contrast to previous reports, Gram et al. (1996), studying starch accu-
mulation in relation to the frequency of cell division and regeneration in
pea protoplasts, suggested that starch accumulation precedes the division
of pea protoplasts. They found that the starch content increased rapidly
during the first 3 days of culture, prior to the onset of division, resulting in a
4.2-fold increase in the intracellular starch area and a threefold increase
(from 27 to 80%) in the number of protoplasts containing starch. Mitosis
was observed after the fourth day and the number of protoplasts under-
going division increased in a stepwise manner, preceded by further starch
accumulation. Since dividing protoplasts were initially 33–66% smaller and
contained 8–42% less starch than non-dividing protoplasts, the dividing
protoplasts contained relatively more starch (6–12%) than non-dividing
protoplasts on a per unit volume basis. Interestingly, the starch level
reached before the onset of the first mitosis was comparable to the level
found in actively dividing micro-calli, suggesting a requirement of certain
levels of starch accumulation for the induction of pea protoplast division.
It can be suggested that there may be an optimum starch content for
protoplast division and that levels below or above this threshold may be
inhibitory for mitosis.
177
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:07 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
There are two basic applications of cultured plant cells that exploit their
totipotence (ability to express the entire genetic information and regener-
ate plants). Firstly, to clone the cells and produce plants that are identical
and, secondly, to change/manipulate the genome and create novel types of
plants. The former application is to maintain valuable genotypic traits and
the latter application is to improve or obtain new characters. Using somatic
cell techniques, plants can be genetically manipulated using somaclonal
variation, in vitro mutagenesis and somatic hybridization/cybridization and
transformation by the introduction of foreign genes. Changes in genomes
generally appear at a low frequency and a large population of an organism,
therefore, is necessary for manipulation. In this respect, in vitro culture
(callus culture, cell suspension or isolated protoplasts) has the advantage of
a very large number of individuals in a small space, compared with the large
growing area and intensive labour required to treat equivalent numbers of
plants.
During the early stages of the development of tissue culture methods,
in vitro cultured cells were believed to be uniform and similar to the initial
material. Regenerants obtained through embryogenesis or organogenesis
in vitro as a result of asexual reproduction, therefore, should be pheno-
typically and genotypically identical to the donor plant. In the early 1970s,
however, there were reports of morphological and cytological changes in
tobacco plants grown from in vitro cultures (Zagorska et al., 1974). Later,
there were similar reports of variability in tissue culture-derived plants. This
phenomenon was given the name of ‘somaclonal variation’ by Larkin and
Scowcroft (1981) and is generally found in plants regenerated from callus
tissue, cell suspensions or isolated protoplasts and, to a lesser extent, in
meristem or shoot tip cultures.
178
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:07 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 163
The isolation of cells from the integrity of the whole plant and the
reorganization of genome function during the establishment of in vitro
culture often cause fundamental destabilization of the genetic and
epigenetic status. The content and composition of culture media affect
chromosomal and cell cycle instability (Gould, 1986). For example, plant
growth regulators (auxins and cytokinins), that are essential for the de-
differentiation and re-differentiation of the cells in vitro, often act as agents
for genomic changes. Also, inorganic (e.g. phosphates and nitrates) and
organic (e.g. carbohydrates) compounds within the medium contribute
to cell cycle abnormalities. Even physical factors, such as temperature, the
light regime and the viscosity and osmolarity of culture media, are known to
affect the cell division cycle of plant cells in culture. The culture phase and
the rate of subculture are also of importance, since prolonged cultivation
can cause more changes in the nuclear and cytoplasmic genome. Selection
of one cell type, however, can occur during subculture, resulting in less
diversity being observed in such cultures when they are maintained for a
prolonged period (Zagorska, 1995). Pre-existing genetic differences, like
the ploidy level of the initial material, the explant origin and the geno-
type can be another source of variation. The regeneration pathway, via
organogenesis or embryogenesis, is another factor causing or eliminating
the appearance of somaclones. Somatic embryogenesis, as a mechanism of
plant formation from a single cell, was postulated to give ‘free of variability’
regenerants (Vasil, 1986). Nevertheless, recent evidence is presented
below on variant plants that have been regenerated via somatic
embryogenesis.
179
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:08 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
180
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:08 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 165
6.4.5 Variation in grain legumes at the cell and tissue culture level in vitro
Before the term somaclonal variation was introduced into the literature
(Larkin and Scowcroft, 1981), a relatively large amount of evidence was
available that plant tissues in culture exhibit a broad spectrum of cytologi-
cal and karyological changes and abnormalities. This fact also included
grain legume species. Since the main aim of this part of the chapter is to
review heritable somaclonal variation at the plant level, typical examples of
variation on cell and tissue culture level in vitro are only mentioned briefly
(Table 6.3). Within this table, various altered cell and callus lines
are characterized; however, with the absence of plant regeneration, the
heritable nature of these changes is subject to speculation.
181
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:09 A3867: AMA: Hedley: First Proof:30-Oct-00 6
166
Color profile: Disabled
Arachis hypogaea Anther-derived callus Chromosome number Mixoploid callus (haploid to Bajaj et al. (1981)
(peanut, groundnut) octoploid cells)
Cajanus cajan Anther-derived callus Chromosome number Haploid, diploid and Bajaj et al. (1980)
aneuploid cells
Cicer arietinum Callus, cell suspension Salt tolerance (NaCl) NaCl-tolerant cell lines Gosal and Bajaj (1984)
Callus Salt tolerance (NaCl) NaCl-tolerant cell lines Pandey and Ganapathy (1984)
Anther-derived callus Chromosome number Haploid to octoploid cells Bajaj and Gosal (1987); Gosal
and Bajaj (1988)
182
Glycine max Cell suspension Resistance (reaction) to Accumulation of a Ebel et al. (1976)
elicitor from Phytopthora phytoalexin glyceolin in
N. Kuchuk et al.
6
Callus; excised Resistance (reaction) to Bioassay for screening Gray et al. (1986); Willmot
cotyledons soybean brown stem rot resistant mutants in culture; et al. (1989)
Phaseolus vulgaris Excised roots, callus, Resistance to Pseudomonas Differential growth inhibition Bajaj and Saettler (1968,
183
sclerotiorum culture filtrate screening of partial
physiological resistance
Biotechnology
6
tip-derived resistance levels of free
Callus Proline; elevated tolerance to
Pisum sativum Callus Chromosome number; Polyploid cells (3n, 4n and Kallak and Yarvekylg (1968,
chromosome aberrations; more); multinuclear cells; 1971, 1976, 1977a,b)
mitotic abnormalities karyological abnormalities
Callus Chromosome number Polyploid (up to 12n) and Frolova and Shamina (1974)
aneuploid cells
Immature Chromosome number; Polyploid cells (up to 8n and Mikhailov and Bessonova
cotyledon-derived chromosome aberrations more); chromosomal (1975)
callus aberrations
Pollen callus Chromosome number Haploid and mixoploid Gupta (1975)
(mainly tetraploid) cell
populations
Cell suspension Chromosome number Multinuclear cells, aneuploidy Ghosh and Sharma (1979)
184
Callus culture Chromosome number Highly polyploid cells (32n or Knosche and Gunther (1980);
more) Knosche (1981)
N. Kuchuk et al.
Callus culture Tolerance to the herbicides Calli with improved tolerance Jakel et al. (1990)
Propham and Probanil to herbicides
(O-isopropyl-3-chlor-
6
Callus, cell suspension Salt tolerance (NaCl) NaCl-tolerant cell lines Gosal and Bajaj (1984)
Callus Chromosome number, Haploid, diploid and Kunakh et al. (1984)
185
(ploidy level measured polyploid cells (more than
cytophotometrically as 16C)
Biotechnology
C-value)
Callus Chromosome number Polyploid and eneuploid cells Ogura (1982)
Callus culture Tolerance to the herbicides Calli with improved tolerance Jakel et al. (1990)
6
(O-isopropyl-3-chlor-
phenylcarbamate)
callus and cell cultures has been published by Griga et al. (1986). Variation
in ploidy level and other chromosomal changes and aberrations in tissue
and cell cultures of grain legume species, other than pea or faba bean, are
less frequently documented in the literature (Table 6.3).
The inability to regenerate plants from cytologically altered cells and
tissues in many of the above cited reports, may be due to the selective
advantage of normal (diploid) cells in the process of organ formation
(diplontic selection). The inability of plant regeneration from cytologically
altered tissues determines that these changes cannot be practically
exploited at present and may only serve as a model for theoretical studies.
The mechanisms of chromosome variation in plant tissue cultures have
been reviewed by Ogura (1990).
186
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:11 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 171
187
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:12 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
188
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:13 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 173
189
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:13 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Table 6.4. Somaclonal variation in grain legumes on the whole plant level (regenerants and their seed progenies).
174
Species Type of in vitro culture Plant regeneration via Trait Response (value/description) References
Color profile: Disabled
Glycine max Immature zygotic Somatic Qualitative and Albinotic chimaeras (R0), Barwale and Widholm
embryo-derived embryogenesis, quantitative chlorophyll deficiency, (1987)
callus organogenesis traits abnormal leaf morphology,
dwarf plant habit (R1–R3)
Cotyledonary node Proliferation of shoot Qualitative and Plant height, sterility Graybosch et al. (1987)
meristems and de quantitative (R0, R1, R2)
novo shoots traits
(organogenesis)
Immature embryo Somatic Plant Increased variation in R1 Hildebrand et al. (1989)
embryogenesis morphology, (mainly leaf morphology;
lipid lipid composition); not
composition inherited to R2
190
Cotyledonary node- Organogenesis Qualitative Lanceolate leaves, leaf and Freytag et al. (1989)
and epicotyl-derived traits pod variegation, determinate
N. Kuchuk et al.
6
Immature cotyledons Direct somatic Qualitative Complete and partial Amberger et al. (1992)
embryogenesis traits sterility, wrinkled and
191
plantlets (R0)
Immature zygotic Somatic Chromosome Tetraploid plants (R0) Kysely et al. (1987)
Biotechnology
6
traits habit; dark green leaves;
oblong leaflets; first flower
Continued
Table 6.4. Continued.
176
Species Type of in vitro culture Plant regeneration via Trait Response (value/description) References
Color profile: Disabled
Shoot apex-derived Shoot organogenesis Nuclear DNA Genotype-dependent, Cecchini et al. (1992)
callus variation tissue culture induced
DNA content variation in
regenerated plants (R0)
Immature zygotic Somatic Qualitative and Significantly altered leaf Stejskal and Griga
embryo-derived embryogenesis, quantitative and flower morphology; (1992); Griga et al.
callus, young organogenesis traits sterility; lethality (R0); pods (1995); Griga and Létal
leaflet-derived callus and seeds per plant; crude (1995)
protein content (R1–R4)
Arachis hypogaea Cotyledon callus Unknown Resistance to Enhanced resistance of R2 Venkatachalam et al.
Cercosporidium plants (1998)
personatum
192
culture filtrate
N. Kuchuk et al.
Phaseolus vulgaris Shoot meristems Meristem Resistance to Resistant regenerants (not Gantotti et al. (1985)
proliferation phaseolotoxin proven)
6
infected with shoot formation Phytophthora in regenerants and fungal
Biotechnology 177
193
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:15 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
194
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:16 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 179
with three cotyledons and three unifoliate leaves. Variation in fatty acid
composition was also higher in the R1 generation than in control material.
Variation in both morphological and seed composition characters, how-
ever, was not observed in the following R2 generation, probably because of
unstable epigenetic effects.
Stephens et al. (1991) observed a wrinkled leaf variant in the R2 genera-
tion of a soybean line regenerated through organogenesis. Observations of
progeny from selfed normal and variant derivatives of this line suggested
genetic instability for this trait. Reciprocal crosses indicated that the mutant
trait was inherited cytoplasmically. The unusual segregation ratios were
attributed to organelle segregation and to cytoplasmic inheritance.
Amberger et al. (1992) regenerated 475 plants of nine soybean
cultivars, via direct somatic embryogenesis from immature cotyledons. The
R1, R2 and R3 progeny from the regenerated plants were scored for qualita-
tive variation and inheritance of variant phenotypes. These included partial
sterility (R0, R1, R2), complete sterility (R0), abnormal leaf morphology (R0,
R1, R2, R3) chlorophyll chimaeras (R2, R3), chlorophyll deficiencies (R2,
R3), changes in growth habit (R2, R3), yellow edges on cotyledons (R3), no
unifoliates (R3), dwarf plants (R2), yellow–green plants (R3) and isozyme
variants (R2). Inheritance studies of chlorophyll-deficient, curled-leaf and
wrinkled leaf plants confirmed that these traits were genetically controlled.
Although none of the variants exhibited any obvious agronomically
favourable characteristics, the study resulted in the identification of novel
variants that may prove useful in the dissection of the soybean genome.
New variants included a malate dehydrogenase null and an aconitase null
(Amberger et al., 1992), curled leaves, lethal chlorophyll deficiencies, no
unifoliates and yellow-edged cotyledons. Similarly, Stephens et al. (1991)
observed an unusual segregation of wrinkled-leaf mutation that could be
considered as a cytoplasmically inherited trait.
Mathews et al. (1986) regenerated mung bean (V. radiata) plants from
de-embryonated cotyledons. Considerable variation was observed in the
R2 population, 7% of the R1 plant progenies segregating for chlorophyll
mutations and another 7% for viable morphological mutations. The chlo-
rophyll mutations included chlorina and xantha types, which were lethal
under field conditions. The viable mutations included those with penta-
foliate leaves, sterility, and green seed coat and cotyledon colour. None of
these mutants was found in the control population. The mutation rate per
100 R2 plants was 1.8% for the chlorophyll and for the viable mutants.
195
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:16 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
196
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:17 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 181
197
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:18 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
198
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:18 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 183
199
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:19 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Herbicide tolerance
200
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:19 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 185
201
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:20 A3867: AMA: Hedley: First Proof:30-Oct-00 6
186
Color profile: Disabled
Transformation
Species Gene/trait incorporated method References
Glycine max 5-Enolpyruvyl-shikimate-3-phosphate synthase (EPSP); herbicide Agrobacterium Hinchee et al. (1988)
tolerance (glyphosate) – Roundup®
Phosphinotricin acetyl transferase (PAT, bar); herbicide tolerance (PPT) Bombardment Christou and Swain
– Basta® (1990)
202
Lysine-feedback-insensitive bacterial DHDPS/ dapA and AK/lysC; Bombardment Falco et al. (1995)
fivefold lysine increase in the seed
N. Kuchuk et al.
Methionine-rich 2S albumin from Brazil nut; high oleic acid; modified Agrobacterium James (1997)
oil; virus resistance bombardment
Synthetic cryIAc gene (B.t.) – resistance to insect larvae (four species) Bombardment Stewart et al. (1996)
6
DNA into ovary
Bean pod mottle virus (BPMV) coat protein gene – virus resistance Agrobacterium Di et al. (1996)
203
Lupinus angustifolius Sulphur-rich sunflower seed albumin (enhanced methionine level in Agrobacterium Molvig et al. (1997);
Biotechnology
204
Cicer arietinum Seed-derived embryos Agrobacterium Organogenesis Fontana et al. (1993)
Vicia narbonensis (narbon Epicotyl segments Agrobacterium Somatic embryogenesis Pickardt et al. (1995)
N. Kuchuk et al.
6
Lupinus angustifolius Maturing embryo axis Agrobacterium Organogenesis Pigeaire et al. (1997)
Biotechnology 189
Insect resistance
There are two general strategies for genetically engineered insect resis-
tance in plants. Firstly, the incorporation of genes from specific bacteria,
which encode proteins that are toxic to insects. Secondly, the incorporation
of genes encoding plant-derived inhibitors of protein, or carbohydrate,
digestion in insects (Galun and Breiman, 1997).
AMYLASE INHIBITORS Seeds of the common bean (P. vulgaris) are naturally
resistant to bruchid beetles (e.g. cowpea weevil, Callosobruchus maculatus),
because of the presence of an α-amylase inhibitor (αAI-Pv), a seed protein
that is toxic to the larvae. Other legumes (e.g. pea, chickpea, cowpea, Azuki
bean), however, do not contain this α-amylase inhibitor-derived tolerance.
205
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:22 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Shade et al. (1994) and Schroeder et al. (1995) transferred the gene encod-
ing αAI-Pv into pea, using Agrobacterium-mediated transformation based
on the co-cultivation of embryonic axis segments and regeneration via
organogenesis (Schroeder et al., 1993). The α-amylase inhibitor gene was
stably expressed in the transgenic pea seeds up to the T5 seed generation.
αAI-Pv accumulated in the pea seeds up to 3% of the soluble protein, a level
that was higher than that normally found in beans (1–2%). In the T5 seed
generation, the development of pea weevil (Bruchus pisorum) larvae was
blocked at an early stage. Seed damage was minimal and seed yield was not
significantly reduced in the transgenic plants. In addition to the resistance
of transgenic pea plants to the bruchid beetles attacking the crop growing
in the field (B. pisorum), the transformed peas also showed complete resis-
tance to bruchid species that damage stored seeds, in particular cowpea
weevil (C. maculatus) and Azuki bean weevil (Callosobruchus chinensis; Shade
et al., 1994). Although αAI-Pv also inhibits human α-amylase, it is reported
that cooked peas should not have a negative impact on human energy
metabolism.
Dillen et al. (1997) transformed Phaseolus acutifolius with a
genomic fragment encoding the P. vulgaris arcelin-5a-protein, using an
Agrobacterium-mediated approach. It is believed that this seed storage
protein confers resistance to the insect Zabrotes subfasciatus, a major pest
of P. vulgaris. Arcelin-5a was produced at high levels in the seeds and the
authors suggest using of P. acutifolius as a ‘bridging’ species to introduce
transgenes into the economically more important species P. vulgaris.
Virus resistance
Virus coat protein-mediated resistance, based on the concept of ‘cross
protection’ and antisense-RNA derived resistance, has been reported for
grain legumes. The concept of cross protection is derived from the fact that
the infection of a given crop plant with mild strains of viruses and viroids
prevents or reduces the symptoms caused by a subsequent virulent strain.
Rather than using the whole virus, however, only part of the viral genome,
encoding the coat protein (CP), is integrated into the plant genome. The
antisense RNA (a mirror sequence of an mRNA sequence) may provide
viral tolerance by interfering either with the translation of viral mRNAs, or
with the replication of the viral genome (Greenberg and Glick, 1993; Galun
and Breiman, 1997).
Di et al. (1996) transformed soybean, using an Agrobacterium-mediated
approach, with a bean pod mottle virus (BPMV) CP-gene and found that
30% of the R2 plants derived from one transgenic line were resistant
to BPMV infection. Similarly, Xu et al. (1996) obtained A. rhizogenes trans-
formed transgenic soybean plants with the integrated gene for soybean
mosaic virus (SMV) coat protein. Arago et al. (1996) introduced the
antisense sequence of AC1, AC2, AC3 and BC1 genes, from the bean
golden mosaic geminivirus, to P. vulgaris transgenic plants, using particle
206
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:23 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 191
Nutritional quality
Galun and Breiman (1997) categorized nutritional quality of crops into
four groups: fatty acids, lipids and oils; carbohydrates; proteins and pig-
mentation. Within grain legumes, most reports relate to the improvement
of oils and proteins by genetic transformation. Weber et al. (1998), how-
ever, has reported changes in carbohydrate metabolism in transgenic Vicia
narbonensis.
In proteins, the strategy has concentrated mainly on improving
amino acid composition, in particular, the methionine (Met) and lysine
(Lys) content. Saalbach et al. (1994) transformed V. narbonensis, a close
relative of faba bean (V. faba), with the methionine-rich 2S albumin gene of
the Brazil nut (Bertholletia excelsa). This synthetic gene, controlled by the
CaMV 35S promoter, however, was highly expressed only in leaves and
roots and was hardly detectable in the seeds. Later, the transformation
protocol was improved by fusing the 2S albumin gene with the seed-specific
legumin B4 promoter from V. faba (Pickardt et al., 1995; Saalbach et al.,
1995a,b). Transformation was carried out using the Agrobacterium-mediated
approach together with a regeneration protocol using somatic embryo-
genesis from callus. Transformed calli were selected for kanamycin
resistance and the induced somatic embryos were screened for GUS activity
and cloned by multiple shoot regeneration. Fertile R0, R1 and R2 transgenic
plants were obtained and seed-specific gene expression was found in
transformants with the legumin B4 promoter/2S albumin gene fusion.
Analysis of the R2 plants indicated a Mendelian inheritance of the 2S
albumin gene. Some transformants exhibited a threefold increase in the
methionine content of the salt-soluble protein fraction extracted from
seeds. The same gene was introduced, using the biolistic process, into
P. vulgaris and stable transgenic bean plants were generated (Arago et al.,
1996). Again the gene was inherited in a Mendelian fashion in most of the
transgenic bean lines.
Falco et al. (1995) increased the lysine content in soybean seeds by
circumventing the normal feedback regulation of two enzymes of the bio-
synthetic pathway, aspartokinase (AK) and dihydropicolinic acid synthase
(DHDPS). Lysine-feedback-insensitive bacterial DHDPS and AK enzymes,
encoded by the Corynebacterium dapA gene and a mutant E.coli lysC gene,
respectively, were expressed in transgenic soybean seeds, following trans-
formation via particle bombardment and by fusion with the GUS reporter
gene. The result was a ten- to several hundredfold increase in free lysine
and up to a fivefold increase in the total seed lysine content, lysine contrib-
uting up to 33% of the total seed amino acid content. The lysine content in
R2 and R3 seeds remained at least as high as that observed in the R1 seed,
207
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:24 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
6.5.5 Field trials with transgenic grain legume plants and commercialized
transgenic legume crops
During the period 1986–1997, c. 25,000 transgenic crop field trials were
conducted in 45 countries, on more than 60 crops covering 10 traits.
Seventy-two per cent of all transgenic crop field trials were conducted in the
USA and Canada followed in descending order by Europe, Latin America
and Asia, with a few conducted in Africa. The most frequent crops in these
trials were maize, tomato, soybean, canola, potato and cotton and the most
frequent traits were herbicide tolerance, insect resistance, product quality
and virus resistance (James, 1997). Recently, a great deal of progress has
been achieved in the transformation and in the commercialization of cool
season legume species in Australia (Atkins et al., 1998; Hamblin et al., 1998).
Four co-operating laboratories/companies are now able to transform
seven legume species (Lupinus angustifolius, Lupinus albus, Lupinus luteus,
C. arietinum, P. sativum, V. faba and L. culinaris), almost all of which have
been field tested (Table 6.7).
The situation with commercialized transgenic crops in 1996 and 1997 is
shown in Tables 6.7, 6.8 and 6.9. From the data it is evident that soybean is
the only grain legume representative within recently commercialized
transgenic crops. On the other hand, soybean, in particular herbicide-
tolerant varieties, has a leading position in this context. The progress in
208
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:24 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 193
Table 6.7. Traits already commercialized in field trials, and under development
for selected crops, 1997–1998 (James, 1997; Hamblin et al., 1998).
209
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:25 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Table 6.8. Global area (millions of hectares) of transgenic crops grown in 1996
and 1997, by crop (James, 1997).
1996 1997
Increase
Crop Area % Area % 1996/7 ratio
Table 6.9. Global area (millions of hectares) of transgenic crops grown in 1996
and 1997, by trait (James, 1997).
1996 1997
Increase
Trait Area % Area % 1996/7 ratio
210
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:25 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 195
211
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:26 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
212
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:26 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 197
and B, have been identified in maize (Stinard et al., 1993), rice (Mizuno
et al., 1993) and pea (Burton et al., 1995). The proteins of these two families
are structurally related and are similar to glycogen-branching enzymes
from E. coli bacteria, which have been used to produce highly branched
starch in transformed potato (Shewmaker et al., 1994).
In the embryos of peas containing the r mutation (see Chapter 7),
the percentage of amylopectin is reduced from about 65% to about 35%.
This relative decrease in amylopectin has been associated with decreased
branching activity and the loss of one branching enzyme isoform (Smith,
1988). Antisense experiments inhibiting isoform B activity in potato,
however, resulted in no significant modification of starch synthesis, or of
the amylose : amylopectin ratio (Kossmann et al., 1991; Mueller-Roeber and
Kossmann, 1994).
Recently, different lines of transgenic potatoes have been generated
where the expression of starch biosynthetic genes has been repressed, using
an antisense RNA technique. In this way plants have been obtained that
synthesize a wide variety of structurally modified starches. These starches
are currently being assessed for their applicability in different industrial
processes in the food and non-food sectors (Kossmann et al., 1997).
213
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:27 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
214
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:27 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 199
215
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:28 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
these genes are present as operons in association with other genes involved
in galactoside utilization. Genes have been isolated from a number of
sources including Bacillus stearothermophillus (Ganter et al., 1988), E. coli
(Aslandis et al., 1989), Streptoccoccus mutans (Aduse-Opoku et al., 1991) and
Pedicoccus pentosaceous (Gonzales and Kunka, 1986).
In processing soybean and cowpea meal, α-galactosidases from various
sources including Bifidiobacterium sp. (Sakai et al., 1987), Aspergillus awamori
(Smiley et al., 1976), Aspergillus niger (Somiari and Balogh, 1995) and Lacto-
bacillus fermentum (Garro et al., 1996) have been used to remove flatulence
factors. More information concerning the utilization of α-galactosidases in
processing has been discussed elsewhere (see Chapter 4).
The development of transgenic grain legume plants with seed
specific expression of α-galactosidases could be a tool to overcome the
RFO problem. At present, genes encoding seed-specific proteins from
soybean (Okamura et al., 1986), pea (Ellis et al., 1988) and bean (Green-
wood and Grispefls, 1985) have been cloned and their promoters could
be used to construct chimaeric α-galactosidase genes with seed-specific
expression. The physiological role of the antinutritional carbohydrates
within the plant, however, is not clear. As discussed earlier (see Chapter 5),
there is a hypothesis suggesting that the α-galactosides have an important
role in seed maturation and in resistance to desiccation. In which case,
an α-galactosidase functioning during seed maturation could be
disadvantageous for seed quality. A better strategy could be α-galactosidases
that could be activated by external factors, preferably after seed harvesting.
From this point of view, genes encoding thermostable enzymes from
hyperthermophiles could be of interest for this purpose. One group of
hyperthermophilic bacteria is the genus Thermotoga. To date, Thermotoga
spp. are the only known hyperthermophiles capable of growing on
cellulose. They produce a multiplicity of hydrolases, which are involved
in the metabolism of various polysaccharide substrates. Recently an
α-galactosidase from Thermotoga neapolitana has been used for the hydrolysis
of guar (galactomannan) gum (McCutchen et al., 1996). This enzyme has
a temperature optimum close to 100°C, its activity decreasing with lower
temperatures.
It may be possible to transfer the thermostable α-galactosidase
chimaeric gene, under the control of a seed-specific promotor, to commer-
cial legume breeding lines. Hypothetically, the activity of this enzyme in
transgenic seeds would be low at temperate climatic conditions. The
accumulation of the RFO in transgenic seeds, therefore, could be at a
similar level to the non-transferred control cultivar and, hence, normal
seed quality would not be reduced. After harvesting, during seed
processing the enzyme could be activated by high temperature and could
then decrease the content of α-galactosides. In addition, this process
could be applied to canned green peas, where it is impossible to remove the
RFO by other methods.
216
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:29 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 201
Plant cell suspension culture provides a defined and controlled system for
the study of cell metabolism. Some advantages of cell suspension cultures
for metabolic studies, compared with the whole plant are: rapid and
uniform growth with the possibility of controlling nutritional conditions;
ease of extraction and purification of large quantities of metabolic
products; a more efficient use of labelled compounds and a lack of inter-
fering microorganisms. Although the metabolic systems of cell cultures and
of whole plants are frequently similar, some quantitative differences do
occur.
The aim of this section is to illustrate how cell suspension cultures can
be used to study the metabolism and biochemistry of plant cell walls. In
particular, the possibility of using cell suspensions from legumes as a model
system for studying their plant cell wall metabolism.
The plant cell wall is a highly dynamic structure playing an important role
in growth and development, morphology, cell-to-cell communication and
transport processes (Fry, 1989; Hayashi, 1989; Bowles, 1990; Sakurai, 1991;
Takeuchi et al., 1994; Heredia et al., 1995; Roberts, 1996; Ishii, 1997;
Driouich and Staehelin, 1997). It enables the plant cell to resist internal
and/or external pressures, provides a structural barrier to some molecules
and protects against invasion by insects and by pathogens (Bowles, 1990).
The cell cytoplasm obtains its metabolic substrates through the cell wall
and excretes other substances across it. The cell wall is a complex structure,
which contains mainly carbohydrates, proteins, lignins and water as well
as other substances embedded in it, such as cutin, suberin and certain
inorganic compounds.
Morphologically, three zones can be differentiated in plant cell walls,
the middle lamella, the primary wall and the secondary wall. The middle
lamella is the most external of the three zones and acts as a separating
panel between two cells. It consists almost exclusively of pectic substances.
The primary cell wall consists of cellulose microfibrils, complex polysaccha-
rides and N- and O-linked glycoproteins. Most plant cells have only a pri-
mary wall and the middle lamella, but some specialized cells exist that also
have the secondary wall. This is a supplementary wall with a predominantly
mechanical function.
It has been established (Mauch and Staehelin, 1989; Bolwell, 1993;
Penel and Greppin, 1996) that there are many glycoproteins and soluble
217
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:29 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
proteins with various enzyme activities, located in either the cell wall or
the extracellular space. Some of the enzymes are needed for modifying
the organization of the macromolecular network, others are involved in
processes such as defence reactions against pathogens (Bowles, 1990; Cote
and Hahn, 1994). The cell wall hydrolysis enzymes (cellulases, pectinases
and a series of glycosidases, such as α- and β-galactosidases, α- and
β-mannosidases, etc.) are involved in processes such as fruit maturation and
softening, as well as in responses to pathogen exposure and other stress
factors (Fry, 1989; Bowles, 1990). Peroxidases are known to take part in the
formation of links between lignin, proteins, hemicellulose and ferulic acid
(Bowles, 1990).
The structural components of primary plant cell walls are cellulose micro-
fibrils, other complex polysaccharides and N- and O-linked glycoproteins
(Heredia et al., 1995). Of these components, only cellulose microfibrils are
synthesized at the cell surface by plasma membrane enzymes. All other
types of cell matrix molecules are produced by Golgi-based enzyme systems
(Driouich et al., 1993; Driouich and Staehelin, 1997) and are transported,
via secretory vesicles, to the cell surface. Recent biochemical investigations
have led to the identification and partial characterization of Golgi-localized
glycosyltransferases, involved in the synthesis of xyloglucans, pectic polysac-
charides and glucoronoxylans (Camiranel et al., 1987; Gibeat and Carpita,
1994).
Immunolabelling experiments, with libraries of anti-polysaccharide
antibodies, have been used to outline the spatial organization of polysac-
charides (Moore et al., 1991; Zhang and Staehelin, 1992). It is important to
remember, however, that all of these studies carried out with antibodies
are directed against specific sugar domains of the polysaccharides and not
against specific glycosyltransferases. This is because none of these enzymes
has been fully purified to date, although some of them have been identified
and partially characterized (Driouich and Staehelin, 1997). The isolation of
such enzymes and the generation of specific antibodies against them is a
major problem that still has to be resolved.
218
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:30 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 203
219
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:30 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
and they play major roles in plant development processes and plant–
pathogen interactions. The structural characterization of these molecules is
a major experimental challenge. Knowledge of the structure of oligosac-
charides and the biological responses to these signals, now has progressed
to the point where detailed studies on their mode of action can be
undertaken.
In spite of the complexity of working with cell walls, knowledge of cell wall
polymer structure has expanded considerably in recent years (Becker et al.,
1974; Sakurai, 1991; Mutafschiev et al., 1993; McCann and Roberts, 1994;
Heredia et al., 1995). Knowledge of the biosynthesis and the functions of
the polymers, however, remains limited.
Plant cell suspension culture offers new possibilities for investigating
some aspects of cell wall metabolism (Bolwell, 1985; Butenko, 1985; Morris
et al., 1985; Dwight Camper and McDonald, 1989; Wink, 1994). In this type
of culture, single cells, or clumps of cells, grow and multiply suspended
in liquid media. Plant cell suspension cultures provide a model system for
studying various molecular, physiological and genetical problems, which
can be manipulated in ways that cannot be applied to whole plants. Cell
suspensions can be cultivated in flasks on shakers or in bioreactors, which
allows control of the nutritional and environmental conditions. As a result,
controlled growth and morphological development can be ensured. Cell
growth is rapid and more uniform than for cells within the plant. Further-
more, they offer a possibility for the controlled supply of metabolites, and
every cell in such a suspension has direct access to the external medium
containing these metabolites. Plant cell suspensions also offer a standard
system that allows the results obtained by different research groups to be
compared.
It has been established (Wink, 1994) that plant cell suspension is a
physiologically complex system, in which both the cell biomass and the
culture medium are included. The culture medium is not only the source
of all necessary nutrients, but also a functional cell compartment with a
number of different metabolites sequested in it (Olson et al., 1969; Van
Huystee and Lobazzewski, 1982; Wink, 1984, 1994; Morris et al., 1985;
Konno et al., 1986, 1987, 1989; Kawasaki, 1989; Uchiyama et al., 1993; Van
Huystee et al., 1994; Dolaptchiev et al., 1996; Ilieva et al., 1996a,b; Sims et al.,
1996). It is well known that suspension cultured plant cells secrete various
polysaccharides and glycoproteins into the culture medium (Olson et al.,
1969; Kawasaki, 1989; Uchiyama et al., 1993; Sims et al., 1996). These
secreted substances are thought to be components of the middle lamella,
derived from primary cell walls, two elements that are difficult to separate
220
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:31 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 205
221
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:31 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
222
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:32 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
Biotechnology 207
223
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:32 A3867: AMA: Hedley: First Proof:30-Oct-00 6
Color profile: Disabled
Composite Default screen
7
Breeding
G. Engqvist
andet Agronomy
al.
225
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:33 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
Fig. 7.1. Crop of the ‘semi-leafless’ pea held up by the tendrils intertwining.
226
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:38 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
Traditional field pea breeding techniques often start with making crosses in
a greenhouse in the autumn, growing the bulked first filial generation – F1
seeds in a greenhouse in the spring and then growing the F2 in the field the
following summer. If a pedigree breeding method is followed, the F2 will
usually be planted spaced out and the first selections, based on individual
plants, will then be carried out. Pedigree plant selections will be made at
the F3, often during an off-season cultivation at a location in the Southern
Hemisphere (if the breeding programme is based in the Northern
Hemisphere). The F4 of each pedigree line would then be grown in an
observation plot at the home station location. At this stage a vigorous
227
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:39 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
selection for stem stiffness and seed quality would be performed between
the lines. The F5 selected lines could then again be grown at an off-season
location in the Southern Hemisphere. At the F5 stage, selection based on
agronomic characters would be made.
The selection process at the F5 is often preceded by progeny tests of the
F4 generation in the greenhouse for certain diseases and this information is
then used to aid the F5 selection. The first comparative yield trials take
place at the F6 stage together with the start of elite seed production. The
first major comparative yield trial and replicated trials in other countries
will be carried out at the F7. The official trials can start at the F8 and
continue through to the F10.
The pedigree method allows an effective and relatively quick search
through a population following a cross, although it is rather labour
intensive and demanding and, therefore, expensive.
The main alternative to the pedigree breeding method is the bulk method,
in which the F2 is grown as a bulk. The F3 is then grown in plots for yield
determination at the home station, with two or three replicates of each plot
being grown depending on the amount of available seed. Part of the F3 seed
is sown later in the same season as spaced plants, to facilitate individual
plant selection. The F3 yield plots are harvested and weighed. Those
populations that show the highest relative yield are identified and single
plant selections are made on the spaced plants from the same population.
Preliminary yield estimates of the lines selected in the F3 start in F5.
The schedules outlined for the pedigree and bulk selection methods may
sometimes be changed. For example, repeat plant selections may be made
to increase homogeneity. Also, when a cross is made to incorporate plant
disease resistance characters, the F2 may be put directly through a selection
procedure to identify plants with the desired character. If and when the
breeder wishes to search through a large number of populations the selec-
tion work needs to be simplified. In this instance the bulked populations
can be grown for several years, delaying the plant selection until the F4 or
F5 generation. This has the advantage that the selected plants are more
homozygous and the derived lines more uniform than when selections are
made in the F2. At the same time the bulked populations can be put in
to comparative yield trials, preferably under severe conditions, and those
228
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:39 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
229
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:40 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
230
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:40 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Table 7.2. Management centres within Europe for the main grain legume databases.
Color profile: Disabled
231
Institute of Plant Breeding JOK Finland 17
Agrobiologische Institute LIN Austria 33
Nordic Gene Bank for Agricultural and Horticultural Plants NGB Sweden 3055
Breeding and Agronomy
7
Institute for Plant Production and Qualification TAP Hungary 934
Vegetable Crops Research Institute, Research Station Ujmajor UJM Hungary 748
understanding the synthesis of starch have been made following the identi-
fication and development of mutants within the starch biosynthetic path-
way. As well as aiding fundamental studies on starch synthesis, such mutants
are also available to breeders for the development of crops where the seed
contains starches with improved nutritional characteristics, or new uses.
Other pathways that lend themselves to genetic characterization, that
could provide breeders with new sources of variation, are those leading to
the synthesis of soluble carbohydrates, in particular the raffinose family of
oligosaccharides (RFO). Information on the known variation for both the
RFO and starch biosynthetic pathways is outlined in the following sections.
Starch
Two mutations, r and rb, affecting the content and composition of starch
have been known for many years and were first identified by their affect on
the shape of the dry mature seed, which in both cases appeared wrinkled.
Mendel (1865) used a naturally occurring mutation at the r locus in his
epoch making studies on inheritance. This mutation is the genetic change
that has brought about the development of the vined pea crop, which is
harvested when immature and then either quick-frozen, quick-dried or
canned. A second, apparently naturally occurring, mutation at the rb locus
was first genetically characterized by Kooistra (1962), who also determined
that r and rb were independent loci, even though the presence of recessive
genes at either locus gave rise to wrinkled seeds.
The presence of mutations at either the r or rb locus results in a
reduction in the starch content of the dry seed to about 35%, compared
with about 50% found in the non-mutant wild type. In addition to affecting
starch content, the presence of these mutations also affects starch composi-
tion. Starch from r mutant seed contains about 70% amylose, while starch
from the rb mutant contains about 20% amylose, compared with the
wild-type level of about 30% amylose (Wang et al., 1998). It is now known
that the mutation at the r locus is in a gene encoding a starch-branching
enzyme (Fig. 7.2; Bhattachatryya et al., 1990; Martin and Smith, 1995) and
results in a lack of activity of this enzyme during seed development.
The mutation at the rb locus reduces the activity of ADP-glucose
232
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:41 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
Fig. 7.2. The pathway of starch biosynthesis in pea (Wang et al., 1998).
233
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:46 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
r 10 27–36 60–75
rb 8 30–37 23–32
rug3 5 1–12 12
rug4 3 38–43 31–33
rug5 3 29–35 43–52
lam 5 39–49 4–10
aStarchis given as a percentage of the dry weight.
bAmylose is given as a percentage of the starch
on a dry weight basis.
contents ranged from 27 to 36% and 60 to 75%, for the r alleles, and from
30 to 37% and 23 to 32%, for the rb alleles, respectively (Table 7.4).
The remaining mutants were assigned to four previously unidentified
loci (rug3, rug4, rug5 and lam). The presence of mutations at the rug3
locus resulted in seeds containing only 1–12% starch, depending on the
particular mutant allele, and the starch had relatively low amounts
of amylose (Table 7.4). Mutations at this locus decrease the activity of
plastidial phosphoglucomutase (Fig. 7.2; Harrison et al., 1998). The seeds
from plants containing this mutation are viable even though they possess
very low or no starch and the resulting plants appear to grow normally
(Harrison et al., 1998).
Seeds from the rug4 mutants are only mildly wrinkled and the starch and
amylose contents are decreased to only 38–43% and 31–33%, respectively
(Table 7.4). It is now known that mutations at the rug4 locus result in a
dramatic reduction in the activity of sucrose synthase (Craig et al., 1999), an
enzyme that is outside of the dedicated starch biosynthetic pathway (Fig. 7.2).
Mutations at the rug5 locus result in a reduction in starch content to
29–35% and an increase in the proportion of amylose in the starch to
43–52% (Table 7.4). Mutants containing the mutations at the rug5 locus
have a reduced level of one of the major soluble starch synthases (Fig. 7.2;
Craig, 1998).
The sixth group of alleles isolated from the chemical mutagenesis
programme carried out by Wang et al. (1990) differed from the other five
in that seed shape and starch content were more similar to the wild type
(Table 7.4). The composition of the starch, however, was similar to that of
low amylose or ‘waxy’ mutants identified in maize and other species, hence
the gene symbol lam, for low amylose. Mutants containing the lam mutation
lack activity for a major granule-bound starch synthase (Denyer et al., 1995).
The presence of all of these new mutants affecting starch gives breeders
an opportunity to develop pea lines with seeds containing a range of
starches or, in the case of the rug3 mutants, seeds with little or no starch.
234
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:47 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
The physical properties of the starches produced by these new mutants are
described in Chapter 4. Having identified a range of loci affecting starch, it
then becomes possible to combine the genes in different ways to produce
double or perhaps treble mutants that could extend further the available
variation and potential uses.
Saponins
Although not present in large quantities in legume seeds, saponins are
considered to be antinutritional. Genetic variation for saponin type has
235
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:47 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
been found within soybean seeds and a genetic model linking structure (of
the sugar chains) and five genes; Sa-1a, Sg-1b, Sg-2, Sg-3 and Sg-4, has been
proposed by Tsukamoto et al. (1993). Variation for soyasapogenol B has
been reported in a study of 19 P. vulgaris varieties (Burbano et al., 1999).
This now opens up the possibility of breeders selecting lines that have low
amounts of saponins with antinutritional properties and higher amounts of
those saponins with known health benefits, improving the quality of their
seed.
Fig. 7.3. (A) Plant breeding selection methods for chemical component. Selection
from gene banks. Opposite page: (B) Selection from F2 populations.
236
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:50 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
237
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:58 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
238
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:00:58 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
Fig. 7.4. Miniature drill used for sampling from single seeds. The insert shows a
hole bored through a pea seed with the resultant flour sample – 30 mg, more than
enough for chemical or physical analyses. The seed can be grown normally to
produce a plant.
239
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:02 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
been carried out using mid-infrared (MI), which it has been suggested
could yield more chemical information than NI. Both methods are based
on the selective absorption and vibration of specific chemical bonds within
molecules. The fundamental vibrations result from absorption in the MI
while the second and third overtones occur in the NI.
240
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:03 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
241
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:03 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
The availability of water to crops during the time when the seeds are devel-
oping is a major factor determining yields. It is known, for example, that
water deficits during the time of seed filling in soybean crops decreases
seed size. This may result from a reduction in the supply of assimilates from
the maternal plant and/or an inhibition of seed metabolism. Experiments
have been carried out to determine whether it is maternal or zygotic factors
that limit seed growth at times of water deficit (Bernal-Lugo and Leopold,
1992). When water was withheld from greenhouse grown soybean plants
during the linear seed-filling period, leaf water potential decreased rapidly,
inhibiting canopy photosynthesis completely within 3 days. Seed dry weight
continued to increase, however, at or near the control rate. The level of
total extractable carbohydrates in leaf, stem and pericarp tissue decreased
242
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:03 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
243
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:04 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
Table 7.5a. The registration requirements of several European countries for new
varieties of pea – agronomic characters.
Czech Republic
Denmark
Germany
Moldova
Romania
Hungary
Bulgaria
Ukraine
Sweden
Estonia
Poland
France
Spain
Italy
Agronomic character UK
Yield ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓
Earliness of ripening ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓
Shortness of straw ✓ ✓ ✓ ✓ ✓ ✓ ✓
Height of crop canopy at harvest ✓ ✓ ✓
Standing ability ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓
Ease of combining ✓ ✓ ✓ ✓ ✓
One-step combining ✓ ✓
Height of lower pod attachment ✓ ✓
Pod shattering or spill at harvest ✓ ✓ ✓
Resistance to:
Drought ✓ ✓
Alternaria alternata ✓ ✓
Ascochyta pisi/Mycosphaerella pinodes ✓ ✓ ✓ ✓ ✓ ✓ ✓
Black rust ✓ ✓
Colletotrichum ✓ ✓
Downy mildew (Peronospora pisi) ✓ ✓ ✓ ✓ ✓ ✓ ✓
Erysiphe pisi ✓ ✓ ✓ ✓ ✓
Pea enation virus ✓ ✓ ✓ ✓
Pea wilt (race 1) Fusarium sp. ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓
Winter hardening – cold tolerance ✓ ✓
1000 seed weight ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓
244
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:05 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
Table 7.5b. The registration requirements of several European countries for new
varieties of pea – processing characters.
Czech Republic
Denmark
Germany
Moldova
Romania
Hungary
Bulgaria
Ukraine
Sweden
Estonia
Poland
France
Spain
Italy
UK
Processing character
Table 7.5c. The registration requirements of several European countries for new
varieties of pea – chemical characters.
Czech Republic
Denmark
Germany
Moldova
Romania
Hungary
Bulgaria
Ukraine
Sweden
Estonia
Poland
France
Spain
Italy
UK
Chemical character
245
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:06 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
246
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:07 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
The seed market and the food industry strongly appreciate grain colour
stability (evaluated by Bulgaria, Czech Republic, Sweden and Ukraine),
either green or yellow-seeded varieties being preferred (Table 7.5b). The
time of harvest is important, therefore, because strong sunshine can lead to
the bleaching of seed colour (tested in Hungary). In addition, seed samples
have to be free from waste and stain, as well as the effects of fungal diseases
(e.g. blemishes, contaminants) and pests (e.g. damaged by pea moth cater-
pillar and pea seed beetle). These characters are regarded as important for
canning and packet sales, and meeting these requirements results in higher
payments for such samples.
Some marrowfat varieties may need special agronomic measures to
ensure high quality produce. Samples which do not meet the technological
requirements mentioned above (except colour) may be suitable for the
micronizing market. The micronising process produces a high protein feed
for use in certain dried animal feedstuff and pet foods. Another criterion
for marrowfat peas might be large, even-sized seed, which is tested through
grading (Table 7.5b). It is necessary for a variety to be graded in only one
large group and not to be divided into more size groups (a high number of
size groups is analysed in Ukraine).
247
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:07 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
248
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:07 A3867: AMA: Hedley: First Proof:30-Oct-00 7
Color profile: Disabled
Composite Default screen
8
Manipulating
C. Hedley Grain Legume Carbohydrates
249
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:08 A3867: AMA: Hedley: First Proof:30-Oct-00 8
Color profile: Disabled
Composite Default screen
234 C. Hedley
the human diet (see Chapter 3). It is likely, however, that removal of these
compounds by plant breeders would have an adverse affect on the growth
and development of the plant and seed. Since the RFO are believed to be
involved in protection against abiotic stress and may have an important role
in germination (see Chapter 5). As with the protein inhibitors, the RFO
also may have a positive affect in the diet by promoting beneficial changes
in the gut flora, in particular increasing the proportion of Bifidobacteria,
which have been implicated in protection against colon cancer (see
Chapter 3).
With the possible exception of the storage proteins, nutritionists have
been more interested in the effect of the antinutritional, or non-nutritional
components in legume seeds, rather than the nutritional components. The
main nutritional carbohydrate in most legume seeds is starch. It is known
that existing legume starches are digested more slowly than those from
other species, in particular from cereals, and that this can have a positive
effect in reducing the glycaemic index in humans, which is an advantage to
those people with type 2 diabetes (see Chapter 3). The lack of information,
in general, on starch digestion and, more specifically, on why legume
starches are digested more slowly, however, creates the problem of not
knowing the nutritional consequences of changing the starch content or
composition in legume seeds.
Also, there is very little information on the effect of manipulating
starch on the growth and development of the plant or seed. Studies using
pea seed mutants affected in starch content, composition (see Chapter 7)
and granular structure (see Chapter 4) have given some information on the
possible consequences of manipulating legume starches. For example, it is
known from using these lines that blocking starch synthesis completely
in peas, by introducing a mutation at the rug3 locus, appears to have a
minimal affect on the growth of the plant and on the viability of the seed
(Wang et al., 1998). There is also no evidence that changing the chemical
composition and the granular structure of pea starches has any adverse
affects on the seed biology. One interesting observation, following the
introduction of mutations directly affecting starch, however, is that they
have pleiotropic effects on other seed components, in particular, proteins
(Casey et al., 1998a,b), lipids (Jones et al., 1995) and soluble carbohydrates
(Jones, 2000). For example, introducing recessive alleles at the r and rb loci
in pea gives a small percentage increase in the protein content, a large
difference in the protein composition and a large percentage increase in
the lipid content of the seed (Wang and Hedley, 1993).
The third carbohydrate category, after the soluble carbohydrates
and starch is the ‘fibre’ fraction, which contains a multitude of soluble
and insoluble compounds, generally characterized by nutritionists as non-
digestible elements in the diet. Many of the fibre components are looked
on as beneficial within the human diet, mainly from a health point of view
(see Chapter 3). Within the plant, most of these compounds are associated
250
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:08 A3867: AMA: Hedley: First Proof:30-Oct-00 8
Color profile: Disabled
Composite Default screen
As mentioned above and in Chapter 3, the main problems with the soluble
carbohydrates lie with the inability to digest the RFO, with the resulting
reduction in nutritional value for animal feed and the problem of
flatulence in humans. Any changes in the content and composition of the
RFO would also need to take into consideration their role in the plant,
mentioned above and in Chapter 5.
The first requirement of any conventional breeding programme
designed to genetically manipulate plant characteristics, such as the
251
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:09 A3867: AMA: Hedley: First Proof:30-Oct-00 8
Color profile: Disabled
Composite Default screen
236 C. Hedley
252
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:09 A3867: AMA: Hedley: First Proof:30-Oct-00 8
Color profile: Disabled
Composite Default screen
8.2.2 Starch
8.2.3 Fibre
253
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:10 A3867: AMA: Hedley: First Proof:30-Oct-00 8
Color profile: Disabled
Composite Default screen
238 C. Hedley
8.3 Conclusions
Defining strategies for improving or manipulating grain legume carbohy-
drates will only result in increased scientific and technological activity if
grain legume crops are considered worth developing from an economic
point of view. As stated in the introduction to this book, in general the
consumption of grain legumes has been declining for many years in the
developed parts of the world. A shrinking market gives rise to decreased
interest from producers and a reduction in available funding for research
and development. This downward spiral can only be stopped and perhaps
reversed by creating new opportunities for grain legume crops and by
adding value to the raw materials derived from them.
It is very important to get away from the ‘poor man’s meat’ identity that
grain legumes have had in the past. The health benefits from consuming
more grain legumes in the diet are well documented and with current
awareness of diet and health issues in western populations this should be a
good ‘selling point’. Another current concern of western populations is the
effect that intensive farming is having on the environment. An increased
acreage of grain legumes and reintroducing these species into farming
rotation systems would reduce the chemical inputs required for other crops
such as cereals. A more sustainable agricultural system would reduce the
level of fertilizers reaching water supplies and have a positive effect on
reducing chemical pollution.
An alternative, or perhaps additional, strategy for increasing the use
and production of grain legumes is to consider the seeds as a source of raw
materials for the processing industry, rather than as an entity to be eaten as
a vegetable. The three major constituents of the starchy legumes, protein,
starch and fibre, all have useful functional properties that can be readily
utilized in food products. Procedures have already been developed for
isolating these three fractions (see Chapter 4) relatively easily, from pea.
Considering the two carbohydrate fractions, pea starch has unique pasting
properties, having a stable development and high end-point viscosity,
compared with equivalent amounts of cereal and tuber starch. Most legume
starches have good gelling properties, although this is usually accompanied
by a high level of syneresis, which could be a negative property. As
254
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:10 A3867: AMA: Hedley: First Proof:30-Oct-00 8
Color profile: Disabled
Composite Default screen
255
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:10 A3867: AMA: Hedley: First Proof:30-Oct-00 8
Color profile: Disabled
Composite Default screen
References
References
References
AACC American Association of Cereal Chemists (1995) Sugars. In: AACC Methods,
Vol. 2, Section 80.
Abbas, I.R., Scheerens, J.C. and Berry, J.W. (1987) Tepary bean starch. Part III.
In vitro Digestibility. Starch/Stärke 39, 280–284.
Abdel-Gawad, A.S. (1993) Effect of domestic processing on oligosaccharide content
on same dry legume seeds. Food Chemistry 46, 25–31.
Abia, R., Buchanan, C.J., Saura-Calixto, F. and Eastwood, M.A. (1993) Structural
changes during the retrogradation of legume starches modify in vitro fermenta-
tion. Journal of Agricultural Food Chemistry 41 (11), 1856–1863.
Abreu, J.M.F. and Bruno-Soares, A.M. (1998) Chemical composition, organic
matter digestibility and gas production of nine legumes grains. Animal Feed
Science and Technology 70, 49–57.
Acevedo, E. and Bressani, R. (1989) Ingestion de fibra dietetica en los paises del
istmo controamericano: implicationes nutricionales. Archivos Latinoamericanos
de Nutricion 39, 392–404.
Acevedo, E., Velazquez-Coronado, L. and Bressani, R. (1994) Changes in dietary
fibre content and its composition as affested by processing of black beans
(Phaseolus vulgaris Tamazulapa veriety). Plant Foods for Human Nutrition 46,
139–145.
Adams, C.A., Rinne, R.W. and Fjerstad, M.C. (1980) Starch deposition and carbo-
hydrate activities in developing and germinating soya bean seeds. Annals of
Botany 45, 577–582.
Adams, C.A., Broman, T.H. and Rinner, W. (1981) Starch metabolism in develop-
ing and germinating soybean seeds is independent of α-amylase activity. Annals
of Botany (London) 48, 433–440.
Adsule, N.R., Lawande, K.M. and Kadam, S.S. (1989) Pea. In: Salunkhe, D.K. and
Kadam, S.S. (eds) CRC Handbook of World Food Legumes: Nutritional Chemistry,
241
257
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:11 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
242 References
Processing Technology and Utilization, Vol. I. CRC Press, Boca Raton, Florida,
pp. 215–251.
Aduse-Opoku, J., Tao, L., Ferretti, J. and Russell, R. (1991) Biochemical and genetic
analysis of Streptococcus mutans – galactosidase. Journal of General Microbiology 137,
757–764.
Ahmed, A.P. and Labavitch, J.M. (1977) A simplified method for accurate determi-
nation of cell wall uronide content. Journal of Food Biochemistry 1, 361.
Ahmed, R., Gupta, S.D. and Ghosh, P.D. (1987) The cytological status of plants
regenerated from shoot-meristem culture of Pisum sativum. Plant Breeding 98,
306–311.
Äkerberg, A.K.E., Liljeberg, H.G.M., Granfeldt, Y.E., Drews, A.W. and Björck, M.E.
(1998) An in vitro method, based on chewing, to predict resistant starch
content in foods parallel determination of potentially available starch and
dietary fibre. Journal of Nutrition 128, 651–660.
Akinyele, I.O. and Akinlosotu, A. (1991) Effect of soaking, dehulling and fermenta-
tion on the oligosaccharides and nutrient content of cowpeas (Vigna
unguiculata). Food Chemistry 41, 43–53.
Akpa, A.D. and Archer, S.A. (1994) Interaction between Pseudomonas syringae pv.
Pisi and isolated pea mesophyll protoplasts. Acta Phytopathology and Entomology
(Hungary) 29, 95–104.
Aldington, Z. and Fry, S.C. (1993) Oligosaccharins. Advances in Botany Research 19,
1–101.
Altaf, N. and Ahmad, M.S. (1990). Chickpea (Cicer arietinum L.). In: Bajaj, Y.P.S.
(ed.) Biotechnology in Agriculture and Forestry, Vol. 10. Legumes and Oilseed Crops 1.
Springer-Verlag, Berlin, pp. 100–113.
Althers, S., Stirm, S. and Jacobsen, H.J. (1993) Immunobiochemical analysis of a
nuclear protein marker for regeneration potential in higher plants. Journal of
Plant Physiology 141, 415–422.
Aman, P. (1979) Carbohydrates in raw and germinated seeds from mung bean and
chick pea. Journal of the Science of Food and Agriculture 30, 869–875.
Amberger, A.L., Palmer, R.G. and Shoemaker, R.C. (1992) Analysis of culture-
induced variation in soybean. Crop Science 32, 1103–1108.
Ambrose, M.J., Wang, T.L., Cook, S.K. and Hedley, C.L. (1987) An analysis of seed
development in Pisum sativum. IV. Cotyledon cell populations in vivo and
in vitro. Journal of Experimental Botany 38, 1909–1920.
Amuti, K.S. and Pollard, C.J. (1977) Soluble carbohydrates of dry and developing
seeds. Phytochemistry 16, 529–532.
Anandarajah, K. and McKersie, B.D. (1990) Manipulating the desiccation tolerance
and vigor of dry somatic embryos of Medicago sativa L. with sucrose, heat shock
and abscisic acid. Plant Cell Reports 9, 451–455.
Andersen, K.E., Buskov, S., Bjergegaard, C., Sorensen, H., Sorensen, J.C. and
Sörensen, S. (1997) Analyses of dietary fibre associated non-carbohydrate
constituents based on supercritical fluid extraction. In: Abstract European Food
Chemistry IX. Interlaken, Switzerland, pp. 209–214.
Anderson, T.A. (1982) Recent trends in carbohydrate consumption. Annual Review
Nutrition 2, 113–132.
Anderson, J.W. and Bridges, S.R. (1988) Dietary fibre content in selected foods.
The American Journal of Clinical Nutrition 47, 440–447.
258
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:12 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 243
Andersson, H. (1992) The ileostomy model for the study of carbohydrate digestion
and carbohydrate effects on sterol excretion in man. European Journal of Clinical
Nutrition 46 (Suppl. 2), 69–76.
Angel, C.R., Sell, J.L. and Zimmerman, D.R. (1988) Autolysis of α-galactosides
of defected soy flakes: influence on nutritive value for chickens. Journal of
Agricultural Food Chemistry 36, 542–546.
Angur, Ch. Lu In, Sakai, K., Ogata, T., Sinay, P., Darvill, A.G. and Albersheim, P.
(1992) Further studies of the ability of xyloglucan oligosaccharides to inhibit
auxin-stimulated grown. Plant Physiology 99, 180–185.
Anonymous (1994) Analysis of carbohydrates by anion exchange chromatography
with pulsed amperometric detection. Dionex Technical Note 20, Dionex Corpora-
tion, Sunnyvale, California.
AOAC Association of Official Analytical Chemists (1984) Sugars and sugar
products. In: Official Methods of Analysis, Vol. 14, Section 31, 573.
Apostol, I., Heinstein, P.F. and Low, P.S. (1989) Rapid stimulation of an oxidative
burst during elicitation of cultured plant cells. Role in defense and signal
transduction. Plant Physiology 90, 109–116.
Arago, F.J.L., Barros, L.M.G., Brasileiro, A.C.M., Ribeiro, S.G., Smith, F.D., Sanford,
J.C., Faria, J.C. and Rech, E.L. (1996) Inheritance of foreign genes in trans-
genic bean (Phaseolus vulgaris L.) co-transformed via particle bombardment.
Theoretical and Applied Genetics 93, 142–150.
Arcioni, S., Pezzotti, M. and Damiani, F. (1987) In vitro selection of alfalfa plants
resistant to Fusarium oxysporum f. sp. Medicaginis. Theoretical and Applied Genetics
74, 700–705.
Arentoft, A.M. and Sorensen, H. (1992) Alpha galactosides and dietary fibres
in relation to pea quality: methods of oligosaccharide analyses. In: AEP (ed.)
Proceedings of the 1st Conference on Grain Legumes, Angers, France. AEP, Paris,
France, pp. 457.
Arentoft, A.M., Michaelsen, S. and Sorensen, H. (1993) Determination of oligosac-
charides by capillary zone electrophoresis. Journal of Chromatography A 652,
517–524.
Aslandis, C., Schmidt, K. and Schmidt, R. (1989) Nucleotide sequences and operon
structure of plasmid-borne genes mediating uptake and utilization of raffinose
in Escherichia coli. Journal of Bacteriology 171, 6753–6763.
Asp, N.G. (1992) Preface. European Journal of Clinical Nutrition 46, 1.
Asp, N.G. (1996) Dietary carbohydrates: classification by chemistry and physiology.
Food Chemistry 57 (1), 9–14.
Asp, N.G. and Björck, I. (1992) Resistant starch. Trends in Food Science and Technology
3, 111–114.
Asp, N.G. and Johansson, C.G. (1984) Dietary fibre analysis. Nutrition Abstracts
Reviews 54, 735–752.
Asp, N.G., Johansson, C.-G., Hallmer, H. and Siljeström, M. (1983) Rapid enzymatic
assay of insoluble and soluble dietary fibre. Journal of Agricultural Food Chemistry
31, 476–482.
Asp, N.G., van Amelsvoort, J.M.M. and Hautvast, J.G.A.J. (eds) (1995) Proceedings
of the concluding plenary meeting of EURESTA – including the final reports of
the working groups. European FLAIR Concerted Action No II (COST 911) on:
Physiological implications of the consumption of resistant starch in man.
Wageningen, The Netherlands, IBSN 90-9008390-1.
259
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:12 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
244 References
Asp, N.G., van Amelsvoort, J.M.M. and Hautvast, J.G.A.J. (1996) Nutritional implica-
tions of resistant starch. Nutrition Research Reviews 9, 1–3.
Atanassov, A.I. and Mehandjiev, A.D. (1979) In vitro induced morphogenesis in pea
embryos. Comptes Rendus de l’Academie Bulgare de Sciences 32, 115–118.
Atkins, C.A. and Smith, P.M.C. (1997) Genetic transformation and regeneration
of legumes. In: Legocki, A., Bothe, H. and Puhler, A. (eds) Biological Fixation of
Nitrogen for Ecology and Sustainable Agriculture. Springer-Verlag, Berlin, Germany,
pp. 283–304.
Atkins, C.A., Smith, P.M.C., Gupta, S., Jones, M.G.K. and Caligari, P.D.S. (1998)
Genetics, cytology and biotechnology. In: Gladstones, J.S., Atkins, C.A. and
Hamblin, J. (eds) Lupins as Crop Plants: Biology, Production and Utilization. CAB
International, Wallingford, pp. 67–92.
Attia, R.S., El-Tabey Shehata, A.M., Aman, M.E. and Hamza, M.A. (1994) Effect of
cooking and decortication on the physical properties, the chemical composi-
tion and the nutritive value of chickpea (Cicer arietinum L.). Food Chemistry 50,
125–131.
Asworth, E.N., Stirm, V.E. and Volenec, J.J. (1993) Seasonal variations in soluble
sugars and starch within woody stems of Cornus sericea L. Tree Physiology 13,
379–388.
Aumaitre, A., Peiniau, J., Bengala Freire, J. and Seve, B. (1992) Sensitivity of the
weaned piglet to pea antinutritional factors: effect of technical treatments. In:
AEP (ed.) Proceedings of the 1st European Conference on Grain Legumes, Anger. AEP,
Paris, pp. 487–488.
Bach Knudsen, K.E. (1997) Carbohydrates and lignin contents of plant materials
used in animal feeding. Animal Feed Science and Technology 67, 319–338.
Bach Knudsen, K.E. and Li, B.W. (1991) Determination of oligosaccharides in
protein-rich feedstuffs by gas-liquid chromatography and high performance
liquid chromatography. Journal of Agricultural and Food Chemistry 39, 689–694.
Bachmann, M. (1993) Metabolism of the raffinose family oligosaccharides in
leaves of Ajuga reptans L. Cold induction, sink to source transition and
compartmentation. PhD thesis, University of Zurich, pp. 125.
Bachmann, M. and Keller, F. (1995) Metabolism of the raffinose family oligosaccha-
rides in leaves of Ajuga reptans L. Inter- and intracellular compartmentation.
Plant Physiology 109, 991–998.
Bachmann, M., Matile, P. and Keller, F. (1994) Metabolism of the raffinose family
oligosaccharides in leaves of Ajuga reptans L. Cold acclimation, translocation,
and sink to source transition: discovery of chain elongation enzyme. Plant
Physiology 105, 1335–1345.
Bajaj, Y.P.S. (1985) In vitro induction of genetic variability in ground nut. Proceedings
International Workshop Cytogenetics of Arachis. ICRISAT, Patancheru, pp. 165.
Bajaj, Y.P.S. and Dhanju, M.S. (1979) Regeneration of plants from apical meristem
tips of some legumes. Current Science 48, 906–907.
Bajaj, Y.P.S. and Gosal, S.S. (1987) Pollen embryogenesis and chromosomal
variation in cultured anthers of chickpea. Chickpea Newsletter 17, 12–13.
Bajaj, Y.P.S. and Gosal, S.S. (1981) Induction of genetic variability in grain-legumes
through tissue culture. In: Rao, A.N. (ed.) Proceedings COSTED Symposium on
Tissue Culture of Economically Important Plants, Singapore, pp. 25–41.
260
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:13 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 245
Bajaj, Y.P.S. and Saettler, A.W. (1968) Effect of culture filtrates of Pseudomonas
phaseolicola on the growth of excised roots and callus tissue cultures of bean.
Phytopathology 58, 1041–1042.
Bajaj, Y.P.S. and Saettler, A.W. (1970) Effect of halo toxin-containing filtrates of
Pseudomonas phaseolicola on the growth of bean callus tissue. Phytopathology 70,
1065–1067.
Bajaj, Y.P.S., Singh, H. and Gosal, S.S. (1980) Haploid embryogenesis in anther
cultures of pigeon-pea (Cajanus cajan). Theoretical and Applied Genetics 58,
157–159.
Bajaj, Y.P.S., Ram, A.K., Labana, K.S. and Singh, H. (1981) Regeneration of geneti-
cally variable plants from the anther-derived callus of Arachis hypogaea and A.
villosa. Plant Science 23, 35–39.
Baker, C.M., Durham, R.E., Burns, J.A., Parrott, W.A. and Wetzstein, H.Y. (1995)
High frequency somatic embryogenesis in peanut (Arachis hypogeae L.) using
mature dry seed. Plant Cell Reports 15, 38–42.
Bakhsh, A., Jones, D.A., Lambert, N. and Hedley, C.L. (1991) Genetic variation for
storage product composition in lentil (Lens culinaris). Aspects of Applied Biology
(Production and protection of grain legumes) 27, 279–281.
Bakhsh, A., Bird, J., Lambert, N., Bacon, J., Chambers, S., Jones, A., Wang, T.L.
and Hedley, C.L. (1992) Quantitative variation for seed storage product
composition in lentil (Lens culinaris). In: AEP (ed.) Proceedings of the 1st European
Conference on Grain Legumes, Angers. AEP, Paris, pp. 153–154.
Barna, K.S. and Wakhlu, A.K. (1993) Somatic embryogenesis and plant regenera-
tion from callus cultures of chickpea (Cicer arietinum L.). Plant Cell Reports 12,
521–524.
Barna, K.S. and Wakhlu, A.K. (1994) Whole plant regeneration of Cicer arietinum
from callus via organogenesis. Plant Cell Reports 13, 510–513.
Barna, K.S. and Wakhlu, A.K. (1995) Direct somatic embryogenesis and plantlets
regeneration of chickpea. In Vitro Cellular and Developmental Biology – Plant 31,
137–139.
Barnett, J.E.G., Rasheed, A. and Corina, D.L. (1973) Partial reactions of D-glucose
6-phosphate-1L-myo inositol 1-phosphate cyclase. Biochemical Journal 131,
21–30.
Bartels, D., Singh, M. and Salamini, F. (1988) Onset of desiccation tolerance during
development of the barley embryo. Planta 175, 485–492.
Bartle, K.D. (1993) Introduction to the theory of chromatic separations with
reference to gas chromatography. In: Baugh, P.J. (ed.) Gas Chromatography: a
Practical Approach. IRL Press, Oxford, pp. 1–14.
Barwale, U.B. and Widholm, J.M. (1987) Somaclonal variation in plants regener-
ated from cultures of soybean. Plant Cell Reports 6, 365–368.
Barwale, U.B. and Widholm, J.M. (1990) Soybean: Plant regeneration and soma-
clonal variation. In: Bajaj, Y.P.S. (ed.) Biotechnology in Agriculture and Forestry,
Vol. 10. Legumes and Oilseed Crops 1. Springer-Verlag, Berlin, pp. 114–133.
Bastianelli, D., Grosjean, F., Peyronnet, C., Duparque, M. and Regnier, J.M. (1998)
Feeding value of pea (Pisum sativum L.) 1. Chemical composition of different
categories of pea. Animal Science 67, 609–619.
Bean, S.J., Gooding, P.S., Mullineaux, P.M. and Davies, D.R. (1997) A simple system
for pea transformation. Plant Cell Reports 16, 513–519.
261
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:13 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
246 References
262
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:14 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 247
Berry, C.S., Anson, I., Miles, M., Morris, V.J. and Russel, P.L. (1988) Physico-
chemical characterisation of resistant starch from wheat. Journal of Cereal Science
8, 203–206.
Bewley, D. and Oliver, M.J. (1992) Desiccation tolerance in vegetative plant tissues
and seeds: protein synthesis in relation to desiccation and a potential role
for protection and repair mechanisms. In: Osmond, C.B., Somero, G. and
Bolis, C.L. (eds) Water and Life: A Comparative Analysis of Water Relationships at the
Organismic, Cellular and Molecular Levels. Springer-Verlag, Berlin, pp. 141–160.
Bewley, J.D. and Black, M. (1978) Physiology and Biochemistry of Seeds. Development,
Germination and Growth, 1st edn. Springer-Verlag, New York.
Bewley, J. D. and Black, M. (1985) Seeds. Physiology of Development and Germination,
1st edn. Plenum Press, New York.
Bewley, J.D. and Black, M. (1994) Seeds. Physiology and Development and Germination,
2nd edn. Plenum Press, New York.
Bewley, J.D. and Reid, J.S. (1985) Mannans and glucomannans. In: Dey, P.M. and
Dixon, R.A. (eds) Biochemistry of Storage Carbohydrates in Green Plants. Academic
Press, London, pp. 289–304.
Bhat, U.R., Paramahans, S.V. and Taranathan, R.N. (1983) Scanning electron
microscopy of enzyme-digested starch granules. Starch 35, 261–265.
Bhattacharyya, M.K., Smith, A.M., Ellis, T.H.N., Hedley, C.L. and Martin, C. (1990)
The wrinkled-seed character of peas described by Mendel is caused by a
transposon-like insertion in a gene encoding starch branching enzyme. Cell 60,
115–122.
Bhatty, R.S. (1990) Cooking quality of lentils: the role of structure and composition
of cell walls. Journal of Agricultural and Food Chemistry 38, 376–383.
Bhave, M.R., Lawrence, S., Barton, C. and Hannah, L.C. (1990) Identification and
molecular characterization of shrunken-2 cDNA clones of maize. Plant Cell 2,
581–588.
Biermann, C.J. (1988) Introduction to analysis of carbohydrates by gas-liquid
chromatography (GLC). In: Bierman, C.J. and McGinnis, G.D. (ed.) Analysis of
Carbohydrates by GLC and MS. CRC Press, Boca Raton, Florida, pp. 1–17.
Biliaderis, C.G., Page, C.M., Maurice, T.J. and Juliano, B.O. (1986) Thermal charac-
terisation of rice starches: a polymeric approach to phase transitions of
granular starch. Journal of Agricultural Food Chemistry 34, 6–14.
Biliaderis, C.G. and Tonogai, J.R. (1991) Influence of lipids on the thermal and
mechanical properties of concentrated starch gels. Journal of Agricultural Food
Chemistry 39, 833–840.
Binarova, P., Nedelnik, J., Fellner, M. and Nedbalkova, B. (1990) Selection for
resistance to filtrates of Fusarium spp. in embryogenic cell suspension culture
of Medicago sativa L. Plant Cell Tissue and Organ Culture 22, 191–196.
Binder, R.G. and Haddon, W.F. (1984) Analysis of O-methylinositols by gas–liquid
chromatography-mass spectrometry. Carbohydrate Research 129, 21–32.
Binding, H. (1986) Regeneration from protoplasts. In: Vasil, I.K. (ed.) Cell Culture
and Somatic Cell Genetics of Plants, Vol. 3. Academic Press, Orlando, Florida,
pp. 259–274.
Bingham, S. and Selvendran, R.R. (1983) Workshop report: dietary fibre estima-
tion. In: Wallace, G. and Bell, L. (eds) Fibre in Human and Animal Nutrition. The
Royal Society of New Zealand, pp. 119–121.
263
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:15 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
248 References
264
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:15 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 249
Bochicchio, A., Rizzi, E., Balconi C., Vernieri, P. and Vazzana, C. (1994) Sucrose
and raffinose contents and acquisition of desiccation tolerance in immature
maize embryos. Seed Science Research 4, 123–126.
Bogracheva, T.Ya., Leontiev, S.P. and Genin, Ya.V. (1994) The effect of solutes on
the Gelatinisation of smooth pea starch. Carbohydrate Polymers 25, 227.
Bogracheva, T.Ya., Davydova, N.I., Genin, Ya.V. and Hedley, C.L. (1995) Mutant
genes at the r and rb loci affect the structure and physico-chemical properties of
pea seed starches. Journal of Experimental Botany 46, 1905–1913.
Bogracheva, T.Ya., Ring, S., Morris, V., Lloyd, J.R., Wang, T.L. and Hedley, C.L.
(1997) The use of mutants to study the structural and functional properties
of pea starch. In: Richmond, P., Frazier, P.J. and Donald, A.M. (eds) Starch:
Structure and Function. Royal Society of Chemistry, Cambridge, pp. 230–237.
Bogracheva, T.Ya., Morris, D.G., Ring, S.G. and Hedley, C.L. (1998) The granular
structure of C-type pea starch and its role in gelatinisation. Biopolymers 45,
232–332.
Bogracheva, T.Y., Cairns, P., Noel, T.R., Hulleman, S., Wang, T.L., Morris, V.J.,
Ring, S.G. and Hedley, C.L. (1999) The effect of mutant genes at the r, rb, rug3,
rug4, rug5 and lam loci on the granular structure and physio-chemical proper-
ties of pea seed starch. Carbohydrate Polymers 39, 303–314.
Bohmer, P., Meyer, B. and Jacobsen, H.-J. (1995) Thidiazuron-induced high
frequency of shoot induction and plant regeneration in protoplast derived
pea callus. Plant Cell Reports 15, 26–29.
Bol, J.F., Linthorst, H.J.M. and Cornelissen, B.J.C. (1990) Plant pathogenesis
related proteins induced by virus infection. Annual Reviews of Phytopathology 28,
113–138.
Bolwell, G.P. (1985) Use of tissue cultures for studies of vascular differentiation.
In: Dixon, R.A. (ed.) Plant Cell Culture – a Practical Approach. JRL Press, London,
pp. 107–125.
Bolwell, P. (1993) Dynamic aspects of the plant extracellular matrix. International
Reviews of Cytology 146, 261–323.
Borejszo, Z. and Khan, K. (1992) Reduction of flatulence-causing sugars by
high temperature extrusion of pinto bean high starch fractions. Journal of Food
Science 57, 771–777.
Botham, R.L., Cairns, P., Morris, V.J. and Ring, S.G. (1995a) Physicochemical
characterisation of resistant starch in ileostomy effluent. In: Asp, N.G., van
Amelsvoort, J.M.M. and Hautvast, J.G.A.J. (eds) Proceedings of the Concluding
Plenary Meeting of EURESTA – Including the Final Reports of the Working Groups.
ATO, Wageningen, pp. 71–72.
Bothman, R.L., Cairns, P., Morris, V.J., Ring, S.G., Englyst, H.N. and Cummings,
J.H. (1995b) A physicochemical characterization of chick pea starch resistant
to digestion in the human small intestine. Carbohydrate Polymers 26, 85–90.
Bouché-Pillon, S., Fleurat-Lessard, P., Serrano, R. and Bonnemain, J.L. (1994)
Asymmetric distribution of the plasma-membrane H+-ATPase in embryos of
Vicia faba L. with special reference to transfer cells. Planta 193, 392–397.
Bourdillon, A. (1998) Advantages and constraints of grain legumes for the feed
market. In: AEP (ed.) Opportunities for High Quality, Health and Added-value Crops
to Meet European Demands. Proceedings 3rd European Conference on Grain Legumes.
AEP, Paris, France, p. 5.
265
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:16 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
250 References
266
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:17 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 251
Burbano, C., Muzquiz, M., Ayet, G., Cuadrado, C. and Pedrosa, M.M. (1999) Evalua-
tion of antinutritional factors of selected varieties of Phaseolus vulgaris. Journal of
the Science of Food and Agriculture 79, 1468–1472.
Burchfield, H.P. and Storrs, E.E. (1962) Biochemical Applications of Gas Chromatogra-
phy. Academic Press, New York, pp. 119–120.
Burn, J., Chapman, P.D., Bishop, D.T. and Mathers, J. (1998) Diet and cancer
prevention: the concerted action polyp prevention (CAPP) studies. Proceedings
of the Nutrition Society, 57, 183–186.
Burrows, W.J. and Carr, D.J. (1970) Cytokinin content of pea seeds during their
growth and development. Physiologia Plantarum 23, 1064–1070.
Burton, R.A., Bewley, J.D., Smith, A.M., Bhattacharyya, M.K., Tatge, H., Ring, S.,
Bull, V., Hamilton, W.D.P. and Martin, C. (1995) Starch branching enzymes
belonging to distinct enzymes families are differentially expressed during pea
embryo development. The Plant Journal 7, 3–15.
Buta, J.G. and Galletti, G.C. (1989) FT-IR investigation of lignin components in
various agricultural lignocellulosic by-products. Journal of the Science of Food and
Agriculture 49, 37–43.
Butenko, R.G. (1985) Some features of cultured plant cells. In: Butenko, R.G. (ed.)
Plant Cell Culture. MIR Publishers, Moscow, pp. 11–34.
Caffrey, M., Fonseca, V. and Leopold, A.C. (1988) Lipid–sugar interactions:
relevance to anhydrous biology. Plant Physiology 86, 754–758.
Cairns, P., Bogracheva, T.Ya, Ring, S.G., Hedley, C.L. and Morris, V.J. (1997) Deter-
mination of the polymorphic composition of smooth pea starch. Carbohydrate
Polymers 31, 275–282.
Camiranel, A., Brummell, D.A. and MacLachlan, G.A. (1987) Fucosylation of
xyloglucan: localization of the transferase in dicotisoms in pea stem cells. Plant
Physiology 84, 753–756.
Canibe, N. and Bach Knudsen, K.E. (1997) Digestibility of dried and toasted pea in
pigs. 1. Ileal and total tract digestibility of carbohydrates. Animal Feed Science and
Technology 64, 293–310.
Capitani, T.A., Trzasko, P., Zallie, J.P. and Mason, W.R. (1996) Starch based lipid
mimetic for foods. US Patent 5512311.
Carr, D.J. and Skene, K.G.M. (1961) Diauxic growth curves of seeds, with special
reference to french beans (Phaseolus vulgaris L.). Australian Journal of Biological
Sciences 14, 1–12.
Carré, B., Prevotei, B. and Leclercq, B. (1984) Cell wall content as a predictor of
metabolisable energy value of poultry feedingstuffs. British Poultry Science 25,
561–572.
Carré, B., Escartin, R., Melcion, J.P., Champ, M., Roux, G. and Leclerco, B. (1987)
Effect of pelleting and association with maize or wheat on the nutritive value of
smooth pea (Pisum sativum) seeds in adult cockerels. British Poultry Science 28,
219–229.
Carré, B., Beaufils, E. and Melcion, J.-P. (1991) Evaluation of protein and starch
digestibilities and energy value of pelleted or unpelleted pea seeds from winter
or spring cultivars in adult and young chickens. Journal of Agricultural and Food
Chemistry 39, 468–472.
Carré, B., Gomez, J. and Chagneau, A.M. (1995) Contribution of oligosaccharide
and polysaccharide digestion, and excreta losses of lactic acid and short chain
267
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:17 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
252 References
fatty acids, to dietary metabolisable energy value in broiler chickens and adult
cockerels. British Poultry Science 36, 611–629.
Carré, B., Melcion, J.P., Widiez, J.L. and Biot, P. (1998) Effect of various processes of
fermentation grinding and storage of peas on the digestibility of pea starch in
chickens. Animal Feed Sciences and Technology 71, 19–33.
Carrouee, B. (1995) Grain legumes in the European Union: current status and
prospect. In: AEP (ed.) Improving production and utilisation of grain legumes.
Proceedings of 2nd European Conference on Grain Legumes. AEP, Paris, France,
pp. 2–4.
Casey, R., Domoney, C., Forster, C., Hedley, C., Hitchin, E. and Wang, T. (1998)
The effect of modifying carbohydrate metabolism on seed protein gene
expression in peas. Journal of Plant Physiology 152, 636–640.
Casey, R., Forster, C., Hedley, C.L., Hitchin, E. and Wang, T.L. (1998a) The rela-
tionship between carbohydrate metabolism and seed protein composition in
peas. In: AEP (ed.) Proceedings of the 3rd European Conference on Grain Legumes:
Opportunities for High Quality, Healthy and Added-value Crops to Meet European
Demands. AEP, Paris, France, pp. 84–85.
Casey, R., Domoney, C., Forster, C., Hedley, C.L., Hitchin, E. and Wang, T.L.
(1998b) The effect of modifying carbohydrate metabolism on seed protein
gene expression in peas. Journal of Plant Physiology 152, 636–640.
Cassidy, A., Bingham, S.A. and Cummings, J.H. (1994) Starch intake and colorectal
cancer risk: an international comparison. British Journal of Cancer 69, 937–942.
Castillo, E.M., de Lumen, B.O., Reyes, P.S. and de Lumen, H.Z. (1990) Raffinose
synthase and galactinol synthase in developing seeds and leaves of legumes.
Journal of Agricultural and Food Chemistry 38, 351–355.
Cavallini, A. and Natali, L. (1991) Nuclear DNA variability within Pisum sativum
(Leguminosae): cytophotometric analyses. Plant Systematics and Evolution 173,
179–185.
Cecchini, E., Natali, L., Cavallini, A. and Durante, M. (1992) DNA variations in
regenerated plants of pea (Pisum sativum L.). Theoretical and Applied Genetics 84,
874–879.
Cegla, G.P. and Bell, K.R. (1977) High pressure liquid chromatography for the anal-
ysis of soluble carbohydrates in defatted oilseed flours. Journal of the American
Oil Chemists Society 54, 150–152.
Cerletti, P., Eynard, L. and Guerrieri, N. (1993) Resistant starch: a negative or
positive entity in foods? Agro Food Industry Hi-Technology 4, 27–29.
Cerning-Beroard, J. and Filiatre-Verel, A. (1976) A comparison of the carbohydrate
composition of legume seeds: horsebeans, peas and lupines. Cereal Chemistry 53,
968–978.
Cerning-Beroard, J. and Filiatre-Verel, A. (1979) Etude comparee de la composition
glucidique des graines de pois lisee et ride. Lebesmittel-Wissenschaft und
Technologie 12, 273–280.
Champ, M. (1992) Determination of resistant starch in foods and food products:
interlaboratory study. European Journal of Clinical Nutrition 46, 51–62.
Champ, M. (1995) Definition, analysis, physical and chemical characterization
of RS. In: Asp, N.G., van Amelsvoort, J.M.M. and Hautvast, J.G.A.J. (eds) Proceed-
ings of the Concluding Plenary Meeting of EURESTA – Including the Final Reports of
the Working Groups. European FLAIR Concerted Action No. 11(COST 911),
Brussels, pp.1–14.
268
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:18 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 253
269
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:19 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
254 References
270
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:19 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 255
271
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:20 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
256 References
272
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:21 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 257
Dey, P.M. (1990) Oligosaccharides. In: Dey, P.M. and Harbourne, J.B. (eds) Methods
in Plant Biochemistry, Vol. 2. Carbohydrates. Academic Press, New York, pp.
189–218.
Dhir, S.K., Dhir, S. and Widholm, J.M. (1991) Plantlet regeneration from immature
cotyledon protoplasts of soybean (Glycine max L.). Plant Cell Reports 10, 39–43.
Dhir, S.K., Dhir, S. and Widholm J.M. (1992) Regeneration of fertile plants from
protoplasts of soybean (Glycine max L. Merr): genotypic differences in culture
response. Plant Cell Reports 11, 285–289.
Di, R., Purcell, V., Collins, G.B. and Ghabrial, S.A. (1996) Production of transgenic
soybean lines expressing the bean pod mottle virus coat protein precursor
gene. Plant Cell Reports 15, 746–750.
Dierick, N. and Decuyper, J. (1998) Effect of exogenous carbohydrase supple-
mentation to high pea (Pisum sativum, L.) and soy bean meal diets on digestion
and nutrient excretion in growing pigs. In: AEP (ed.) Opportunities for High
Quality, Healthy and Added-Value Crops to Meet European Demands. Proceedings of
the 3rd European Conference on Grain Legumes, Valladolid, Spain. AEP, Paris,
pp. 12–13.
Dillen, W., de Clerq, J., Goossens, A., van Montagu, M. and Angenon, G. (1997)
Agrobacterium-mediated transformation of Phaseolus acutifolius A. Gray. Theoreti-
cal and Applied Genetics 94, 151–158.
Dinehskumar, V., Kirti, P.B., Sachan, J.K.S. and Chopra, V.L. (1995) Picloram
induced somatic embryogenesis in chickpea (Cicer arietinum L.). Plant Science
109, 207–213.
Dittrich, P. and Brandl, A. (1987) Revision of the pathway of D-pinitol formation in
Leguminosae. Phytochemistry 26, 1925–1926.
Dolaptchiev, L., Bakalova, A., Vassileva, P., Ilieva, M. and Michneva, M. (1996)
Tobacco plant cell cultures with high content of extracellular phospho-
hydrolases. Enzyme Microbiology and Technology 19, 384–388.
Domoney, C. (1999) Inhibitors of legume seeds. In: Shewry, P.R. and Casey, R.
(eds) Seed Proteins. Kluwer Academic Publishers, Dordrecht, the Netherlands,
pp. 635–655.
Donovan, J.W. (1979) Phase transitions of the starch–water system. Biopolymers 18,
263–275.
van Doorne, L.E., Marahall, L.E. and Kirkwood, R.C. (1995) Somatic embryogenesis
in pea (Pisum sativum L.): effect of explant, genotype and culture conditions.
Annals of Applied Biology 126, 169–179.
Dornbos, D.L. Jr. and McDonald, M.B. Jr (1986) Mass and composition of develop-
ing soybean seeds at five reproductive growth stages. Crop Science 26, 624–630.
Dostálová, J., Krejcová, A., Réblova, Z. and Pokorný, J. (1999a) Changes of
α-galactosides during soaking and cooking of peas (in Czech). Bulletin
Potravinárskeho Výskumu 38, 133–140.
Dostálová, J., Zátopková, M., Réblová, Z. and Pokorný, J. (1999b) Changes of
undigestible saccharides during food processing (in Czech). Zborník Príspevkov
Zjazdu Chemických Spolebnosti 51, 1–2.
Dostálová, J., Borovicková, L. and Pokorný, J. (2000) Modification of recipes
for reduction of daily α-galactoside intake from legumes (in Czech). Výyiva a
potraviny – Zpravodaj 55, 18–19.
Doublier, J.L. (1987) A rheological composition of wheat, maize, faba bean and
smooth pea starches. Journal of Cereal Science 5, 247–262.
273
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:21 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
258 References
Driouich, A. and Staehelin, L.A. (1997) The plant Golgi apparatus: structural
organization and functional properties. In: Berger, E.G. and Roth, J. (eds) The
Golgi Apparatus. Birkhauser Verlag, Basel, pp. 275–301.
Driouich, A., Faye, L. and Staehelin, L.A. (1993) The plant Golgi apparatus: a
factory for complex polysaccharides and glycoproteins. Trends in Biochemical
Sciences 18, 210–214.
Dunn, G. (1974) A model for starch breakdown in higher plants. Phytochemistry 13,
1341–1346.
Dunn, J.M. and Finocchiaro, E.T. (1997) Starch-based texturising agents and
methods of manufacture. US Patent 564243.
Dunn, J.M., Akliva, A.T. and Finocchiaro, E.T. (1996) Starch-based opacifying agent
for food beverages. US Patent 5571334.
Dwight Camper, N. and McDonald, S.K. (1989) Tissue and cell cultures as model
systems in herbicide research. Reviews of Weed Science 4, 169–190.
Dysseler, D. and Hoffem, D. (1994) Estimation of resistant starch intake in Europe.
In: Asp, N.G., van Amelsvoort, J.M.M. and Hautvast, J.G.A.J. (eds) Proceedings of
the Concluding Plenary Meeting of EURESTA, April 1994. European Fair-Concerted
Action no. 11 (COST 911). ATO, Wageningen, pp. 84–86.
Dysseler, P. and Hoffem, D. (1995) Ring test 1993–1994 for total and resistant starch
determination: results and discussion. In: Asp, N.G., van Amelsvoort, J.M.M.
and Hautvast, J.G.A.J. (eds) Proceedings of the Concluding Plenary Meeting of
EURESTA – Including the Final Reports of the Working Groups. ATO, Wageningen,
pp. 87–94.
Eapen, S. and George, L. (1993a) Somatic embryogenesis in peanut: influence of
growth regulators and sugars. Plant Cell Tissue and Organ Culture 35, 151–156.
Eapen, S. and George, L. (1993b) Plant regeneration from leaf disks of peanut and
pigeonpea: Influence of benzyladenine, indolacetic acid and indoleacetic
acid-aminoacid conjugates. Plant Cell Tissue and Organ Culture 35, 223–227.
Eapen, S. and George, L. (1994) Somatic embryogenesis in Cicer arietinum:
influence of genotype and auxins. Biologia Plantarum 36, 343–349.
Ebel, J., Ayers, A.R. and Albersheim, P. (1976) Host-pathogen interactions. XII.
Response of suspension-cultured soybean cells to the elicitor isolated from
Phytopthora megasperma var. Sojae, a fungal pathogen of soybean. Plant Physiology
57, 775–779.
Edje, O.T. and Burris, J.S. (1970). Physiological and biochemical changes in
deteriorating soybean seeds. Proceedings of the Association of Seed Analysis 60,
158–166.
Edwards, A., Fulton, D., Hylton, C., Jobling, S.A., Gidley, M., Roessner, U., Martin,
C. and Smith, A. (1998) A combined reduction in activity of starch synthases II
and III of potato has novel effects on the starch of tubers. Plant Journal 17,
251–261.
Eerlingen, R.C. and Delcour, J.A. (1995) Formation, analysis, structure and proper-
ties of type III enzyme resistant starch. Journal of Cereal Science 22, 129–138.
Eerlingen, R.C., Crombez, M. and Delcour, J.A. (1993a) Enzyme resistant starch. I.
Quantitative and qualitative influence of incubation time and temperature of
autoclaved starch on resistant starch formation. Cereal Chemistry 70, 339–344.
Eerlingen, R.C., Deceunick, M. and Delcour, J.A. (1993b) Enzyme resistant starch.
II. Influence of amylose chain length on resistant starch formation. Cereal
Chemistry 70, 345–350.
274
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:22 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 259
Eerlingen, R.C., Cillen, G. and Delcour, J.A. (1994a) Enzyme resistant starch. IV.
Effect of endogenous lipids and added dodecyl sulphate on formation of
resistant starch. Cereal Chemistry 71, 170–177.
Eerlingen, R.C., Jacobs, H. and Delcour, J.A. (1994b) Enzyme resistant starch. V.
Effect of retrogradation of waxy maize starch on enzyme susceptibility. Cereal
Chemistry 71, 351–355.
Eeuwens, C.J. and Schwabe, W.W. (1975) Seed and pod wall development in Pisum
sativum L. in relation to extracted and applied hormones. Journal of Experimental
Botany 26, 1–14.
Eisenberg, F. Jr. (1967) D-myo-Inositol 1-phosphate as product of cyclization of
glucose-6-phosphate and substrate for a specific phosphatase in rat testis.
Journal of Biological Chemistry 242, 1375–1382.
Elliasson, A.-C. (1980) Effect of water on the gelatinisation of wheat starch.
Starch/Stärke 32, 270–272.
Elliasson, A.-C. (1988) Physical and chemical characteristics of legume starches. ISI
Atlas of Science: Animal and Plant Sciences, pp. 89–94.
Ellis, D. (1995) Genetic transformation of somatic embryos. In: Bajaj, Y.P.S. (ed.)
Biotechnology in Agriculture and Forestry 30, Somatic Embryogenesis and Synthetic
Seed 1. Springer-Verlag, Berlin, pp. 207–220.
Ellis, J.R., Shirsat, A.H., Hepher, A., Yarwood, J.N., Gatchause, J.A., Croy, R.R.D. and
Boulter, D. (1988) Tissue specific expression of a pea legumin gene in seeds of
Nicotiana lumbaginifolia. Plant Molecular Biology 10, 203–214.
Englyst, H.N. and Cummings, J.H. (1985) Digestion of polysaccharides of some
cereal foods in the human small intestine. American Journal of Clinical Nutrition
42, 778–787.
Englyst, H.N. and Cummings, J.H. (1986) Digestion of the carbohydrates of banana
(Musa paradisiaca sapientum) in the human small intestine. American Journal of
Clinical Nutrition 44, 42–50.
Englyst, H.N. and Cummings, J.H. (1987) Digestion of polysaccharides of potato in
the small intestine of man. American Journal of Clinical Nutrition 45, 423–431.
Englyst, H.N. and Cummings, J.H. (1988) Improved method for measurement of
dietary fibre as non-starch polysaccharides in plant foods. Journal of Association
of Official Analytical Chemists International 71, 808–814.
Englyst, H.N., Wiggins, H.S. and Cummings, J.H. (1982) Determination of the
non-starch polysaccharides in plant foods by gas–liquid chromatography of
constituent sugars as alditol acetate. Analyst 107, 307–318.
Englyst, H.N., Kingman, S.M. and Cummings, J.H. (1992) Classification and
measurement of nutritionally important starch fractions. European Journal of
Clinical Nutrition 46 (Suppl. 2) 33S–50S.
Eriksson, T. (1985) Protoplast isolation and culture. In: Fowke, L. and Constable, F.
(eds) Plant Protoplasts. CRC Press, Boca Raton, Florida, pp. 1–20.
Escarpa, A., Gonzales, M.C., Manas, M., Garcia-Diz, L. and Saura-Calixto, F. (1996)
Resistant starch formation: standardization of a high-pressure autoclave
process. Journal of Agricultural Food Chemistry 44, 924–928.
Eskin, N.A.M., Johnsons, S., Vaisey-Genser, M. and McDonald, B.E. (1980) A study
of oligosaccharides in a selected group of legumes. Canadian Institute Food
Science and Technology Journal 13, 40–42.
Evans, D.A. and Bravo, J.E. (1984) Protoplast isolation and culture. In: Evans, D.A.
(ed.) Handbook of Plant Cell Cultures, Vol. 1. Globe, UK, pp. 124–176.
275
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:23 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
260 References
Evans, I.D. and Haisman, D.R. (1982) The effect of solutes on the gelatinization
temperature range of potato starch. Starch/Stärke 34, 224–231.
Ezhova, T.A., Bagrova, A.M., Hartina, G.A. and Gostimski, S.A. (1989) Study of
inheritance of somaclonal variations in Pisum sativum L. Genetics (Moscow) 25,
878–885.
Faik, A., Chambat, G. and Joselean, J.P. (1995) Changes in wall-bound poly-
saccharidase activities during the culture cycle of a Rulus fruticosus cell
suspension. International Journal of Biological Macromolecules 17, 381–386.
Faisant, N., Champ, M., Colonna, P. and Buleon, A. (1993a) Structural discrepan-
cies in resistant starch obtained in vivo in humans and in vitro. Carbohydrate
Polymers 21, 205–209.
Faisant, N., Champ, M., Colonna, P. and Buleon, A. (1993b) Structural features of
starch that escapes digestion in the human small intestine. In: Bioavailability 93.
Proceedings of the Conference on Nutritional, Chemical and Food Processing Implications
of Nutrient Availability, 146–150.
Faisant, N., Champ, M., Colonna, P., Buleon, A., Molis, C., Langkilde, A.M.,
Schweizer, T., Fluorie, B. and Galmiche, J.P. (1993c) Structural features of
starch at the end of the human small intestine. European Journal of Clinical
Nutrition 47, 285–296.
Faisant, N., Buleon, A., Colonna, P., Molis, C., Lartigue, S., Galmische, J.P. and
Champ, M. (1995a) Digestion of raw banana starch in the small intestine of
healthy humans: structural features of resistant starch. British Journal of Nutrition
73, 11–123.
Faisant, N., Colonna, P., Buleon, A., Bouchet, B., Gallant, D.J. and Champ, M.
(1995b) Characteristics of starches escaping human small intestine digestion/
absorption. In: Asp, N.G., van Amelsvoort, J.M.M. and Hautvast, J.G.A.J. (eds)
Proceedings of the Concluding Plenary Meeting of EURESTA – Including the Final
Reports of the Working Groups. ATO, Wageningen, pp. 115–116.
Faisant, N., Gallant, D.J., Bouchet, B. and Champ, M. (1995c) Banana starch
breakdown in the human small intestine studied by electronic microscopy.
European Journal of Nutrition 49, 98–104.
Faisant, N., Planchot, F., Kozlowski, F., Pacouret, M.-P., Colonna, P. and Champ, M.
(1995d) Resistant starch determination adapted to products containing high
level of resistant starch. Scientific Alimentation 15, 85–91.
Falco, S.C., Guisa, T., Locke, M., Mauvais, J., Sanders, C., Ward, R.T. and Webber, P.
(1995) Transgenic canola and soybean seeds with increased lysine. Bio-
Technology (New York) 13, 577–581.
FAO Food Balance Sheets (1996) Statistics Series No. 133. FAO, Rome.
FAO Yearbook Production (1996) 49.
FAO Yearbook Production (1998) 51.
FAO Yearbook Production (2000) 53.
Farmer, E.E., Miloshok, T.D., Saxton, M.J. and Ryan, C.A. (1991) Oligosaccharide
signalling in plants. Specificity of oligouronid-enhanced plasma membrane
protein phosphorylation. Journal of Biological Chemistry 266, 3140–3145.
Faulks, R.M. and Bailey, A.L. (1990) Digestion of cooked starches from different
food sources by procine α-amylase. Food Chemistry 36, 191–203.
Faulks, R.M., Southon, S. and Livesey, G. (1989) Utilisation of α-amylase resistant
maize and pea starch in the rat. British Journal of Nutrition 61, 291–300.
276
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:23 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 261
Feeney, R.E. and Whitaker, J.R. (1982) The Maillard reaction and its prevention. In:
Cherry, J.P. (ed.) Food Protein Deterioration: American Chemical Society Symposium
No. 206. American Chemical Society, Washington, DC, pp. 201–230.
Fehr, W.R., Caviness, C.E., Burmood, D.T. and Pennington, J.S. (1971) Stage of
development descriptions for soybeans, Glycine max (L.) Merrill. Crop Science 11,
929–931.
Fellows, R.J., Egli, D.B. and Legget, J.E. (1978) A pod leakage technique for phloem
translocation studies in soybean (Glycine max (L.) Merr.). Plant Physiology 62,
812–814.
Fenwick, G.R., Price, K.R., Tsukamoto, C. and Okubo, K. (1991) Saponins. In: Toxic
Substances in Crop Plants, pp. 285–327.
Ferket, P.R. (1991) Effect of diet on gut microflora of poultry. Zootecnica Inter-
national 7–8, 44–49.
Fernandez, J.A. and Batterham, E.S. (1995) The nutritive value of lupin-seed and
dehulled lupin-seed meals as protein sources for growing pigs as evaluated by
different technique. Animal Feed Sciences and Technology 53, 279–296.
Fernandez-Romero, D., Moral, R., Gil, J. and Moreno, M.T. (1998) The effect of
TDZ on shoot regeneration from faba bean (Vicia faba). In: AEP (ed.) Opportu-
nities for High Quality, Healthy and Added-Value Crops to Meet European Demands.
Proceedings 3rd European Conference on Grain Legumes, Valladolid, Spain. AEP,
Paris, p. 366.
Fernandez-Romero, D., Moreno, M.T., Cubero, J.I. and Gil, J. (1995). Plantlet
regeneration of chickpea (C. arietinum L.). In: AEP (ed.) Improving Production
and Utilisation of Grain Legumes. Proceedings 2nd European Conference on Grain
Legumes, Copenhagen, Denmark. AEP, Paris, p. 435.
Fidanza, F.A.A., Zuccorini, M.G.P and Corvi, R.R. (1982) Contenuto in fibra
alimentare di alcuni alimenti. La revista della Societa Italiana di Scienza dell
Alimentazione 11, 3–7.
Finch-Savage, W.E., Hendry, G.A.F. and Atherton, N.M. (1994) Free radical activity
and loss of viability during drying of desiccation-sensitive tree seeds. Proceedings
of the Royal Society of Edinburgh Section B (Biological Sciences) 102B, 257–260.
Finocchiaro, E.T. (1996) Starch containing reduced fat peanut butter and method
of manufacture. US Patent 5549923.
Flandre, F. and Sangwan-Norreel, B.S. (1995) Direct somatic embryogegenesis from
axis of mature pea (Pisum sativum L.). Revue de Cytologie et de Biologie Vegetales le
Botaniste 18, 39–60.
Fleming, S.E. (1980) A study of relationships between flatus potential and carbohy-
drate distribution in legume seeds. Journal of Food Science 46, 794–798.
Fleming, S.E. (1981) Influence of cooking method on digestibility of legume cereal
starches. Journal of Food Science 47, 1–3.
Flinn, A.M. and Pate, J.S. (1968) Biochemical and physiological changes during
maturation of fruit of the field pea (Pisum arvense L.). Annals of Botany 32,
479–495.
Flis, M., Sobotka, W. and Zdunczyk, Z. (1997) Effect of variety and dehulling on
nutritional value of white lupin seeds for growing pigs. Journal of Animal and
Feed Sciences 6, 521–531.
Fontana, G., Santini, L., Caretto, S., Frugis, G. and Mariotti, D. (1993) Genetic
transformation in the grain legume Cicer arietinum L. (chickpea). Plant Cell
Reports 12, 194–198.
277
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:24 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
262 References
278
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:25 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 263
Frias, J., Vidal-Valverde, C., Kozlowska, H., Tabera, T., Honke, J. and Hedley, C.L.
(1996c) Natural fermentation of lentils. Influence of time, flour concentration
and temperature on the kinetics of monosaccharides, disaccharide and
alpha-galactosides. Journal of Agricultural Food Chemistry 44, 579–584.
Frias, J., Diaz-Pollan, C., Hedley, C.L. and Vidal-Valverde, C. (1996d) Evolution and
kinetics of monosaccharides, disaccharide and alpha-galactosides during
germination of lentils. Zeitschrift für Lebensmitteln Untersuchungen und Forschungs
202, 35–39.
Frias, J., Sotomayor, C., Diaz-Pollan, C., Hedley, C.L., Fenwick, R.G. and Vidal-
Valverde, C. (1996e) Influence of milling, concentration and temperature
during soaking on soluble carbohydrate content and trypsin inhibitor activity
in lentils seeds. In: Fenwick, R.G., Hedley, C., Richards, R.L. and Khokhar, S.
(eds) Agri-Food Quality. An Interdisciplinary Approach. The Royal Society of
Chemistry, Thomas Graham House, Science Park, Milton Road, Cambridge,
pp. 284–288.
Frias, J., Fornal, J., Ring, S.G. and Vidal-Valverde, C. (1998) Effect of germination
on physico-chemical properties of lentil starch and its components. Food Science
and Technology (LWT), in press.
Frias, J., Vidal-Valverde, C., Sotomayor, C., Diaz-Pollan, C. and Urbano, G. (1999)
Influence of processing on available carbohydrate content and antinutritional
factors of chickpeas. European Food Research and Technology, 210, 340–345.
Frias, J., Vidal-Valverede, C., Sotomayor, C., Diaz-Pollan, C. and Urbano, C. (2000)
Effect of processing on available and non available carbohydrates content and
trypsin inhibitor activity on chick pea. European Food Research and Technology
(in press).
Frokiaer, H., Nielson, D, Sorensen, H., Sorensen, J.C. and Sorensen, S. (1998)
Determinations of soyasaponins in grain legumes by enzyme linked immuno-
sorbent assay based on monoclonal antibodies. In: AEP (ed.) Proceedings of the
3rd European Conference on Grain Legumes, Valladolid. Opportunities for High Quality,
healthy and added-value crops to meet European demands. AEP, Paris, pp. 334–335.
Frolova, L. (1986) Cytogenetic characteristics of a cell population of Vicia faba in
vitro. Biologisches Zentralblatt 105, 105.
Frolova, L.V. and Shamina, Z.B. (1974) Cytogenetic characteristic of tissue culture
of plants from the Leguminaceae family. Cytology and Genetics (Kiev) 8, 413–418.
Frolova, L.V. and Shamina, Z.B. (1978) (in Russian) The kinetics of cell population
of Vicia faba cultured in vitro. Citologija 20, 551–556.
Frühbeck, G., Monreal, I. and Santidrian, S. (1997) Hormonal implications of the
hypocholesterolemic effect of intake of field bean (Vicia faba L.) by young men
with hipercholesterolemia. American Journal of Clinical Nutrition 66, 1452–1460.
Fry, S.C. (1988) Wall polymers: Extraction and fractionation. In: Wilkins, M. (ed.)
The Growing Plant Cell Wall: Chemical and Metabolic Analysis. Longman Scientific
and Technical, Harlow, UK, pp. 49–101.
Fry, S.C. (1989) Cellulases, hemicellulases and auxinstimulated growth: a possible
relationship. Physiologia Plantarum 75, 532–536.
Fry, S.C. (1994) Chemical analysis of components of the primary cell wall. In:
Harris, N. and Oparka, K.J. (eds) Plant Cell Biology: a Practical Approach. Oxford
University Press, Oxford, pp. 199–220.
Fry, S.C., Aldington, Z., Hetherington, P.R. and Aitkai, J. (1993) Oligosaccharides as
signals and substrates in the plant cell wall. Plant Physiology 103, 1–5.
279
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:25 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
264 References
Fu, J.H., Li, L.C., Yuan, H.L., Yue, S.X. and Zhu, L.H. (1993) Expression of the
atrazine-resistant transgenic soybean plants in field tests. Acta Agronomica Sinica
19, 497–500.
Gai, J.Y. and Guo, Z. (1997) Efficient plant regeneration through somatic embryo-
genesis from germinated cotyledon of the soybean. Soybean Genetics Newsletter
24, 41–44.
Galau, G.A., Jakobsen, K.S. and Hughes, D.W. (1991) The controls of dicots late
embryogenesis and early germination. Physiologia Plantarum 81, 280–288.
Gale, J. and Boll, W.G. (1978) Growth of bean cells in suspension culture in the
presence of NaCl and protein-stabilizing factors. Canadian Journal of Botany 57,
777–782.
Gallant, D.J., Bouchet, B., Buleon, A. and Perez, S. (1992) Physical characteristics of
starch granules and susceptibility to enzymatic degradation. European Journal of
Clinical Nutrition, 46 (Suppl. 2), 3–16.
Galun, E. and Breiman, A. (1997) Transgenic Plants. Imperial College Press,
London.
Gamborg, O.L., Davies, B.P. and Stahlhut, R.W. (1983) Cell division and differentia-
tion in protoplasts from cell cultures of Glycine species and leaf tissue of
soybean. Plant Cell Reports 2, 213–215.
Gamborg, O.L., Shyluk, J. and Kartha, K.K. (1975) Factors affecting the isolation
and callus fromation in protoplasts from the shoot apices of Pisum sativum L.
Plant Science 4, 285–292.
Gamborg, O.L., Constabel, F. and Shyluk, J. (1974) Organogenesis in callus from
shoot apices of Pisum sativum. Physiologia Plantarum 30, 125–128.
Ganter, C., Bock, A., Buckel, P. and Mattes, R. (1988) Production of thermostable,
recombinant alpha-galactosidase suitable for raffinose elimination from sugar
beet syrup. Journal of Biotechnology 8, 301–310.
Ganter, J.L.M.S., Correa, J., Reicher, F., Heyraud, A. and Rinaudo, M. (1991) Low
molecular weight carbohydrates from Mimosa scabrella seeds. Plant Physiology
and Biochemistry 29, 139–146.
Gantotti, B.V., Kartha, K.K. and Patil, S.S. (1985) In vitro selection of phaseolotoxin
resistant plants using meristem culture of bean (Phaseolus vulgaris). Phyto-
pathology 75, 1316–1317.
Garcia-Alonso, A., Goni, I. and Saura-Callixto, F. (1998) In vitro starch availability
in raw and cooked legumes (lentils, chick peas and beans). In: AEP (ed.) Oppor-
tunities for High Quality, Healthy and Added-value Crops to Meet European
Demands.Proceedings of the 3rd European Conference on Grain Legumes, Valladolid,
Spain. AEP, Paris, pp. 22–23.
Garcia-Olmedo, R., Diaz, A. and Villanueva, J. (1985) Estudio de la fibra alimentaria
en legumbres cocinadas segun recetas tipicas de la cocina espanola. Anales de
Bromatologia 37, 79–90.
Garleb, K.A., Bourquin, L.D. and Fahey, G.C. (1991) Galactouronate in pectic
substances from fruits and vegetables: comparison of anion exchange HPLC
with Pulsed amperometric detection to standard colorimetric procedure.
Journal of Food Sciences 56, 423.
Garro, M., de Valdez, G.F., Oliver, G. and de Giorgi, G. (1996) Purification of alpha-
galactosidase from Lactobacillus ferentum. Journal of Biotechnology 45, 103–109.
Gatel, F. and Champ, M. (1998) Grain legumes in human and animal nutrition – up
to date results and question marks. In: AEP (ed.) Opportunities for High Quality,
280
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:26 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 265
Health and Added-value Crops to Meet European Demands. Proceedings of 3rd Euro-
pean Conference on Grain Legumes. AEP, Paris, pp. 7–11.
Gaudreault, P.R. and Webb, J.A. (1981) Stachyose synthesis in leaves of Cucurbita
pepo. Phytochemistry 20, 2629–2633.
Gdala, J. (1998) Composition, properties, and nutritive value of dietary fibre of
legume seeds. A review. Journal of Animal and Feed Sciences 7, 131–149.
Gdala, J. and Buraczewska, L. (1997) Ileal digestibility of pea and faba beans
carbohydrate in growing pigs. Journal of Animal Feed Sciences 6, 235–245.
Gdala, J., Buraczewska, L., Wasilewko, J. and Grala, W. (1995) Ileal digestibility of
nutrients in pigs fed diets with whole or dehulled faba beans (Vicia faba, L.)
supplemented with enzymes. In: AEP (ed.) Improving Production and Utilisation
of Grain Legumes. Proceedings of 2nd European Conference on Grain Legumes,
Copenhagen, Denmark. AEP, Paris, pp. 264.
Gdala, J., Jansman, A.J.M., Buraczewska, L., Huisman, J. and van Leeuwen, P.
(1997a) The influence of α-galactosidase supplementation on the ileal digest-
ibility of lupin seed carbohydrates and dietary protein in young pigs. Animal
Feed Sciences and Technology 67, 115–125.
Gdala, J., Johansen, H.N., Bach Knudsen, K.E., Knap, I.H., Wagner, P. and
Jorgensen, O.B. (1997b) The digestibility of carbohydrates, protein and fat
in the small and large intestine of piglets fed non-supplemented and enzyme
supplemented diets. Animal Feed Sciences and Technology 65, 15–33.
Gatel, F. and Champ, M. (1998) Grain legumes in human and animal nutrition – up
to date results and question marks. Opportunities for High Quality, Health and
Added-value Crops to Meet European Demands. Proceedings of 3rd European Conference
on Grain Legumes, Valladolid, Spain. AEP, Paris, pp. 7–11.
Gee, M., McComb, E.A. and McCready, R.M. (1958) A method for the characteriza-
tion of pectic substances in fruit and sugar-beet marks. Food Research 27, 72.
Gelvin, S.B. and Schilperoort, R.A. (1995) Plant Molecular Biology Manual. Kluwer,
Dordrecht.
George, L. and Eapen, S. (1993) Influence of genotype and explant on somatic
embryogenesis in peanut. Oleagineux 48, 361–364.
George, L. and Eapen, S. (1997). Influence of cytokinins and explant type on plant
regeneration in chickpea (Cicer arietinum L.). Physiology and Molecular Biology of
Plants 3, 129–134.
Gerarde, J. (1636) The Herball; or, Generall Historie of Plantes, new edn. Norton and
Whittakers, London, pp. 1219–1221.
Ghosh, P. and Sharma, A.K. (1979) Chromosome analysis in suspension culture of
Vigna sinensis var. Black and Pisum sativum L. Caryologia 32, 419–424.
Gibeaut, D.M. and Carpita, N.C. (1994) Biosynthesis of plant cell wall polysaccha-
rides. Review FASEB Federation of American Societies for Experimental Biology Journal
8, 904–915.
Gidley, M. and Bociek, S.M. (1985) Molecular organisation of starch. A 13C
CP/MAS NMR study. Journal of American Chemical Society 107, 7040–7044.
Gidley, M. and Robinson, G. (1990) Techniques for studying interactions between
polysaccharides. In: Dey, P.M. (ed.) Methods in Plant Biochemistry, Vol. 2. Carbohy-
drates. Academic Press, London.
Gill, R. and Saxena, P.K. (1992) Direct somatic embryogenesis and regeneration of
plants from seedling explants of peanut (Arachis hypogeae): promotive role of
thidiazuron. Canadian Journal of Botany 70, 1186–1192.
281
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:27 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
266 References
282
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:27 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 267
Gram, T., Mattsson, O. and Joersbo, M. (1996) Division frequency of pea proto-
plasts in relation to starch accumulation. Plant Cell Tissue and Organ Culture 45,
179–183.
Grant, J.E., Cooper, P.A., McAra, A.E. and Frew, T.J. (1995) Transformation of peas
(Pisum sativum L.) using immature cotyledons. Plant Cell Reports 15, 254–258.
Grant, J.E., Pither-Joyce, M.D., Fifield, W., Cooper, P.A., Erasmuson, S.K. and
Timmerman-Vaughan, G.M. (1998a) Resistance to alfalfa mosaic virus in trans-
genic pea (Pisum sativum L.). In: Abstracts IX International Congress Plant Tissue
Cell Culture, Jerusalem, p. 147.
Grant, J.E., Pither-Joyce, M., Fifield, W., Cooper, P.A. and Timmerman-Vaughan, G.
(1998b) In: AEP (ed.) Opportunities for High Quality, Healthy and Added-Value
Crops to Meet European Demands. Proceedings of 3rd European Conference on Grain
Legumes, Valladolid, Spain. AEP, Paris, pp. 372–373.
Gray, J. (1991) Starches and Sugars. A Comparison of their Metabolism in Man. ILSI
Europe, Springer-Verlag, London, pp. 1–21.
Gray, L.E., Guan, Y.Q. and Widholm, J.M. (1986) Reaction of soybean callus to
culture filtrates of Phialophora gregata. Plant Science 47, 45–55.
Graybosch, R.A., Edge, M.E. and Delannay, X. (1987) Somaclonal variation in
soybean plants regenerated from the cotyledonary node tissue culture system.
Crop Science 27, 803–805.
Green, J.L. and Angell, C.A. (1989) Phase relations and vitrification in saccharide–
water solutions and the trehalose anomaly. Journal of Physical Chemistry 93,
2880–2882.
Green, R.V., Gordon, S.H., Jackson, M.A., Bennett, G.A., McClelland, J.F. and Jones,
R.W. (1992) Detection of fungal contamination in corn: potential of FT-IR PAS
and DRS. Journal of Agricultural and Food Chemistry 40, 883–886.
Greenberg, B.M. and Glick, B.R. (1993) The use of recombinant DNA technology
to produce genetically modified plants. In: Glick, B.R. and Thompson, J.E.
(eds) Methods in Plant Molecular Biology and Biotechnology. CRC Press, Boca
Raton, Florida, pp. 1–10.
Greenwood, J.S. and Chrispefls, M.J. (1985) Correct targeting of the bean storage
protein phaseolin in the seeds of transformed tobacco. Plant Physiology 79,
65–71.
Griga, M. (1993) Some factors affecting somatic embryogenesis efficiency in
soybean (Glycine max L. Merr). Biologia Plantarum 35, 179–189.
Griga, M. (1996) Agronomic performance of pea (Pisum sativum L.) somaclones
regenerated from embryogenic and organogenic cultures. In: Extended Abstracts
COST 822 WG, Mechanisms and Markers of Regeneration and Genetic Stability,
Turku, Finland, pp. 10–11.
Griga, M. (2000) Morphological alterations in sterile mutant of Pisum sativum
obtained via somatic embryogenesis. Biologia Plantarum, 43, 161–165.
Griga, M. and Klenoticova, H. (1994) Plant regeneration in callus cultures of faba
bean (Vicia faba). Rostlinna Vyroba 40, 697–709.
Griga, M. and Letal, J. (1995) Somaclonal variation study of pea (Pisum sativum L.).
In: Recent Advances in Plant Biotechnology, International Conference of the Institute of
Plant Genetics. 95-IPG, Nitra, Slovakia, pp. 130–135.
Griga, M. and Letal, J. (1996) Improved protocol for pea (Pisum sativum L.) somatic
embryogenesis and field performance of pea somaclones. In: Abstracts 2nd
Asia–Pacific Conference Plant Cell Tissue Culture, Beijing, p. 4.
283
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:28 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
268 References
Griga, M. and Novak, F.J. (1990) Pea (Pisum sativum L.). In: Bajaj, Y.P.S. (ed.) Bio-
technology in Agriculture and Forestry, Vol. 10. Legumes and Oil Seed Crops 1.
Springer-Verlag, Berlin, pp. 65–99.
Griga, M. and Slama, J. (1997) Direct pea somatic embryogegenesis and screening
Pisum genotypes for embryogenic capacity. In: Mechanisms and Markers of
Regeneration and Genetic Stability. COST Extended Abstracts 3rd Meeting of
COST 822 Working Group, Gieben, Belgium, pp. 226–227.
Griga, M. and Stejskal, J. (1993) Organogenesis and somatic embryogenesis in vitro
in pea (Pisum sativum L., P. arvense L.) and field evaluation of somaclones – a
preliminary report. In: Book of Abstracts XXIII Annual ESNA Meeting, Halle,
p. 127.
Griga, M. and Stejskal, J. (1994) Micropropagation of pea (Pisum sativum L.) – in
vitro system and its practical applications. In: Lumsden, P.J., Nicholas, J.R. and
Davies, W.J. (eds) Physiology, Growth and Development of Plants in Culture. Kluwer,
Dordrecht, pp. 278–283.
Griga, M., Tejklova, E., Novak, F.J. and Kubalakova, M. (1986) In vitro clonal
propagation of Pisum sativum L. Plant Cell Tissue and Organ Culture 6, 95–104.
Griga, M., Stejskal, J. and Klenoticova, H. (1992) Plant regeneration in soybean,
pea and faba bean via somatic embryogenesis. In: AEP (ed.) Proceedings of 1st
European Conference on Grain Legumes, Angers. AEP, Paris, pp. 107–108.
Griga, M., Stejskal, J. and Beber, K. (1995) Analysis of tissue culture-derived
variation in pea (Pisum sativum L.) – preliminary results. Euphytica 85, 335–339.
Grosjean, F. and Gatel, F. (1986) Pea for pigs. Pig News Information 7, 443–448.
Gross, P. and ap Rees, T. (1986) Alkaline inorganic pyrophosphatase and starch
synthesis in amyloplasts. Planta 167, 140–145.
Gruchala, L. and Pomeranz, Y. (1993) Enzyme-resistant starch: studies using
differential scanning calorimetry. Cereal Chemistry 70, 163–170.
Gueguen, J. (1983) Legume seed protein extraction, processing, and end product
characteristics. Qualitas Plantarum Plant Food and Human Nutrition 32, 267–303.
Gujska, E. and Khan, K. (1991) Functional properties of extrudates from high
starch fractions of navy and pinto beans and corn meat blended with legume
high protein fractions. Journal of Food Science 56, 431–435.
Guo, H. (1991) Protoplast isolation, culture and differentiation responses in
soybean. MSc thesis, Kentucky University, p. 122.
Gupta, S. (1975) Morhogenetic response of haploid callus tissue of Pisum sativum
(var. B22). Indian Agriculturist 19, 11–21.
Gurr, M.I. and Asp, N.G. (1994) Dietary Fibre, 2nd edn. ILSI Press, Washington, DC.
Haase, N.U. and Kempf, W. (1991) Ein Beitrag zur Biosynthese amylosereicher
Erbsestarken. Starch 43, 2–5.
Haase, N.U. and Shi, H.L. (1991) A characterization of faba bean starch (Vicia faba
L.). Starch 43, 205–208.
Haddon, L.E. and Northcote, D.H. (1976) The effect of growth conditions and
origin of tissue on the ploidy and morphogenetic potential of tissue cultures of
bean (Phaseolus vulgaris L.). Journal of Experimental Botany 27, 1031–1051.
Hague, A., Manning, A.M., Hanlon, K.A., Huschtscha, L.I., Hart, D. and Paraskeva,
C. (1993) Sodium butyrate induces apoptosis in human colonic tumour cee
lines in a p53-independent pathway. Implications for the possible role of
dietary fibre in the prevention of large-bowel cancer. International Journal of
Cancer 55, 498–505.
284
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
14 November 2000 14:32:14 A3867: AMA: Hedley: First Proof:14-Nov-00 References
Color profile: Disabled
Composite Default screen
References 269
Halbach, T., Kiesecker, H., Jacobsen, H.J. and DeKathen, A. (1998) Tissue culture
and engineering of lentil (Lens culinaris Medik). In: AEP (ed.) Opportunities for
High Quality, Healthy and Added-Value Crops to Meet European Demands. Proceedings
3rd European Conference on Grain Legumes, Valladolid, AEP, Paris, p. 376.
Hall, J.M. (1989) A review of total dietary fiber methodology. Cereal Foods World 34,
526–528.
Halmer, P. (1985) The mobilisation of storage carbohydrates in germinated seeds.
Physiologie Vegetale 23, 107–125.
Halperin, W. (1986) Attainment and retention of morphogenetic capacity in vitro.
In: Vasil, I.K. (ed.) Cell Culture and Somatic Cell Genetics in Vitro, vol. 3. Academic
Press, New York, pp. 3–48.
Ham, K.S., Kauffmann, S., Albersheim, P. and Darvill, A.G. (1991) Host pathogen
interactions XXXIX. A soybean pathogenesis related phytoalexin elicitor-active
heat-stable fragments from fungal walls. Molecular Plant–Microbe Interactions 44,
545–552.
Hamblin, J., Barton, J., Atkins, C., Jones, M., Smith, P., Wylie, S., Schroeder, H.,
Molvig, L., Tabe, L. and Higgins, T.J. (1998) The development and status of
transgenic pulses in Australia. In: AEP (ed.) Opportunities for High Quality,
Healthy and Added-Value Crops to Meet European Demands. Proceedings 3rd European
Conference on Grain Legumes, Valladolid. AEP, Paris, pp. 70–71.
Hammat, N. and Davey, M.R. (1988) Isolation and culture of soybean hypocotyl
prototplasts. In Vitro Cellular and Developmental Biology 24, 601–604.
Hammat, N., Kim, H-I., Davey, M.R., Nelson, R.S. and Cocking, E.C. (1987) Plant
regeneration from cotyledon protoplasts of Glycine canescence and G. cladestina.
Plant Science 48, 129–135.
Hammat, N., Ghose, T.K. and Davey, M.R. (1986) Regeneration in legumes. In:
Vasil I.K. (ed.) Cell Culture and Somatic Cell Genetics in vitro, Vol. 3. Academic
Press, New York, pp. 67–95.
Hanke, D.E. and Northcote, D.H. (1974) Cell wall formation by soybean callus
protoplasts. Journal of Cell Science 14, 29–50.
Harley, S.M. and Beevers, L. (1989) Coated vesicles are involved in the transport of
storage proteins during seed development in Pisum sativum L. Plant Physiology
91, 674–678.
Harris, N. (1976) Starch grain breakdown in cotyledon cells of germinating mung
bean seeds. Planta 129, 271–272.
Harrison, C.J. (1996) The rug-3 locus of pea. PhD thesis, University of East Anglia,
Norwich.
Harrison, C.J., Hedley, C.L. and Wang, T.L. (1998) Evidence that the rug3 locus of
pea (Pisum sativum L.) encodes plastidial phosphoglucomutase confirms that
the imported substrate for starch synthesis in pea amyloplasts is glucose-6-
phosphate. The Plant Journal 13, 753–762.
Hartman, C.L., McCoy, T.J. and Knous, T.R. (1984) Selection of alfalfa (Medicago
sativa) cell lines and regeneration of plants resistant to the toxin(s) produced
by Fusarium oxysporum f. sp. Medicaginis. Plant Science Letters 34, 183–194.
Hartman, C.L., Secor, G.A., Venette, J.R. and Albaugh, D.A. (1986) Response of
bean calli to filtrate from Pseudomonas syringae pv. Phaseolicola and correlation
with whole plant disease reaction. Physiology Molecular Plant Pathology 28,
353–358.
285
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:29 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
270 References
Harzic, N., Guilloteau, A. and Huyghe, C. (1998) In vitro shoot formation of Lupinus
albus from cotyledonary node. In: AEP (ed.) Opportunities for High Quality,
Healthy and Added-Value Crops to Meet European Demands. Proceedings 3rd European
Conference on Grain Legumes, Valladolid. AEP, Paris, p. 369.
Hauxwell, A.J., Corke, F.M.K., Hedley, C.L. and Wang, T.L. (1990) Storage protein
gene expression is localised to regions lacking mitotic activity in developing
pea embryos. An analysis of seed development in Pisum sativum XIV. Develop-
ment 110, 283–289.
Hayashi, T. (1989) Xyloglucans in the primary cell wall. Annual Review of Plant
Physiology and Plant Molecular Biology 40, 139–168.
Heatley, M.E. and Smith, R.H. (1996) Whole plant regeneration from shoot apex of
Arachis hypogaea. In Vitro Cellular and Developmental Biology – Plant 32, 115–118.
Hedley, C.L. and Ambrose, M.J. (1980) An analysis of seed development in Pisum
sativum L. Annals of Botany 46, 89–105.
Hedley, C.L. and Ambrose, M.J. (1981) Designing ‘leafless’ plants for improving
yields of dried pea crops. Advances in Agronomy 34, 225–277.
Hedley, C.L. and Smith, C.M. (1985) Genetic variation for pea-seed development.
In: Hebblethwaite, P.D., Heath, M.C. and Dawkins, T.C.K. (eds) The Pea Crop:
a Basis for Improvement. Butterworths, Sevenoaks, UK, pp. 329–338.
Hedley, C.L., Smith, C., Ambrose, M.J., Cook, S. and Wang, T.L. (1986) An analysis
of seed development in Pisum sativum. II. The effect of the r-locus on the
growth and development of the seed. Annals of Botany 58, 371–379.
Hedley, C.L., Lloyd, J.R., Ambrose, M.J. and Wang, T.L. (1994) An analysis of seed
development in Pisum sativum. XVII. The effect of the rb locus alone and in
combination with r on the growth and development of the seed. Annals of
Botany 74, 365–371.
Hedley, C.L., Bogracheva, T.Ya., Lloyd, J.R. and Wang, T.L. (1996) Manipulation of
starch composition and quality in pea seeds. In: Fenwick, G.R., Hedley, C.L.,
Richards, R.L. and Khokhar, S. (eds) Agri-Food Quality, an Interdisciplinary
Approach. The Royal Society of Chemistry, Cambridge, pp. 138–148.
Heim, U., Weber, H., Baumlein, H. and Wobus, U. (1993) A sucrose-synthase gene
of Vicia faba. Expression pattern in developing seeds in relation to starch
synthesis and metabolic regulation. Planta 191, 394–401.
Hendry, G.A.F. (1993) Oxygen, free radical processes and seed longevity. Seed
Science Research 3, 141–153.
Hendry, G.A.F., Finch-Savage, W.E., Thrope, P.C., Atherton, N.M., Buckland, S.H.,
Nilson, K.A. and Seel, W.E. (1992) Free radical processes and loss of seed
viability during desiccation in the recalcitrant species Quercus rubur L. New
Phytologist 122, 273–279.
Heredia, A., Jiménez, A. and Guillén, R. (1995) Review. Composition of plant cell
walls. Zeitschrift für Lebensmittel-Unteruchung und-Forschung 200, 24–31.
Hermann, E.M. and Shannon, L.M. (1985) Accumulation and subcellular localisa-
tion of alpha-galactosidase-hemagglutinin in developing soybean cotyledons.
Plant Physiology 77, 886–890.
Herrera-Estrella, L., Leon, P., Olsson, O. and Teeri, T.H. (1995) Reporter genes
for plants. In: Gelvin, S.B. and Schilperoort, R.A. (eds) Plant Molecular Biology
Manual. Kluwer, Dordrecht, pp. 1–32.
286
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:30 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 271
Herselman, L. and Mienie, C.M.S. (1995) The production of disease free dry bean
seed through meristem tip culture. Centro International de Agricultura Tropical,
Cali, Columbia. Workshop Proceedings. CIAT, SACCAR 188–190.
Hesemann, C.U. (1980) Haploid cells in calli from anther culture of Vicia faba.
Zeitschrift für Pflanzenzuchtung 84, 18–22.
Hildebrand, D.F., Adams, T.R., Dahmer, M.L., Williams, E.G. and Collins, G.B.
(1989) Analysis of lipid composition and morphological characteristics in
soybean regenerants. Plant Cell Reports 7, 701–703.
Hildebrandt, L.A. and Marlett, J.A. (1991) Starch bioavailability in the upper
gastrointestinal tract of colectomized rats. Journal of Nutrition 121, 679–686.
Hill, J. (1756) The British Herbal, an History of Plants and Trees, Natives of Britain,
Cultivated for Use, or Raised for Beauty. Osborne, J., Shipton, J., Hodges, J.,
Newbery, B., Collins, S., Crowder and H. Woodgate, London, 279 pp.
Hill, L.M. and Smith, A.M. (1991) Evidence that glucose-6-phosphate is imported as
the substrate for starch synthesis by plastids of developing pea embryos. Planta
185, 91–96.
Hiller, K., Keipert, M. and Linzer, B. (1966) Triterpensaponine. Die Pharmazie 21,
713–751.
Hinchee, M.A.W., Connor-Ward, D.V., Newell, C.A., McDonnell, R.E., Sato, S.J.,
Gaser, C.S., Fischhoff, D.A., Re, D.B., Fraley, R.T. and Horsch, R.B. (1988)
Production of transgenic soybean plants using Agrobacterium-mediated DNA
transfer. Bio-Technology (New York) 6, 915–922.
Hirasawa, E. (1989) Auxin induce α-amylase activity in pea cotyledons. Plant
Physiology 91, 484–486.
Hizukuri, S. and Takagi, T. (1984) Estimation of the molecular weight for amylase
by the low angle laser-light-scattering technique combined with high-
performance chromatography. Carbohydrate Research 134, 1–10.
Ho, L.C. (1988) Metabolism and compartmentation of imported sugars in sink
organs in relation to sink strength. Annual Review of Plant Physiology and Plant
Molecular Biology 39, 355–378.
Hoch, G., Peterbauer, T. and Richter, A. (1999) Purification and characterization
of stachyose synthase from lentil (Lens culinaris) seeds: galactopinitol and
stachyose synthesis. Archives of Biochemistry and Biophysics 366, 75–81.
Hoekstra, F.A., Crowe, L.M. and Crowe, J.H. (1989) Differential desiccation
sensitivity of corn and Pennisetum pollen linked to their sucrose contents. Plant
Cell and Environment 12, 83–91.
Hoekstra, F.A., Haigh, A.M., Tetteroo, F.A.A. and Roekel, van T. (1994) Changes
in soluble sugars in relation to desiccation tolerance in cauliflower seeds. Seed
Science Research 4, 143–147.
Hoffmann-Ostenhof, O. and Pittner, F. (1982) Biosynthesis of myo-inositols and its
isomers. Canadian Journal of Chemistry 60, 1863–1871.
Holliday, M.J. and Klarman, W.L. (1979) Expression of disease reaction types in
soybeans callus from resistant and susceptible plants. Phytopathology 69, 576.
Honda, S., Nishimura, Takahashi, M.Y., Chiba, H. and Kayehi, K. (1982) A manual
method for the spectrophotometric determination of reducing carbohydrates
with 2-cyanoacetamide. Analytical Biochemistry 119, 194–199.
Hoover, R. and Sosulski, F. (1985a) Studies on the functional characteristics and
digestibility of starches from Phaseolus vulgaris biotypes. Starch 37, 181–191.
287
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:31 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
272 References
288
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:31 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 273
Ilieva, M., Kratchanova, M., Pavlova, E., Dimova, Ts., Petrova, D., Pavlov, A. and
Michneva, M. (1996c) Polygalacturonase and pectinmethylesterase activities
during growth of Nicotiana tabacum 1508 cell suspension. In: Proceedings of XVIII
International Carbohydrate Symposium, Milan, pp. 21–26.
Imberty, A. and Perez, S. (1988) A revisit to the three dimensional structure of
B-type starch. Biopolymers 27, 1205–1221.
Imberty, A., Chanzy, H. and Perez, S. (1988) The double-helical nature of the
crystalline part of A-starch. Journal of Molecular Biology 201, 365–378.
Ioshikava, M., Takenchi, I. and Horino, O. (1990) A mechanism for ethylene-
induced disease resistance in soybean: enhanced synthesis of an elicitor-
releasing factor 3-endoglucanase. Physiology and Molecular Plant Pathology 37,
367–376.
Ishii, T. (1997) Structure and functions of feruloylated polysaccharides. Plant
Sciences 127, 111–127.
Islam, R. (1994) Somatic embryogenesis from immature cotyledons of chickpea
(Cicer arietinum L.). Pakistan Journal of Botany 26, 197–199.
Islam, R., Farooqui, H. and Riazuddin, S. (1993) In vitro organogenesis of chickpea
and its transformation by Agrobacterim tumefaciens. Plant Tissue Culture 3, 29–34.
Iyer, V., Salunkhe, D.K., Satha, S.K. and Rockland, L.B. (1980) Quick cooking beans
(Phaseolus vulgaris L.) II. Phytate, oligosaccharides and antienzymes. Qualities
Plantarum Foods for Human Nutrition 30, 45–52.
Iyengar, A.K. and Kulkarni, P.R. (1977) Oligosaccharide levels of processed
legumes. Animal Feed Sciences and Technology 14, 222–223.
Jackson, J. and Hobbs, S. (1990) Rapid multiple shoot production from cotyledon-
ary node explants of pea (Pisum sativum L.). In Vitro Cellular and Developmental
Biology – Plant 26, 835–838.
Jacobs, H., Eerlingen, R.C., Clauwert, W. and Delcour, J.A. (1995) Influence of
annealing on the pasting properties of starches from varying botanical sources.
Cereal Chemistry 72, 480–487.
Jacobsen, H.J. (1992) Biotechnology applied to grain legumes – current state
and approaches. In: AEP (ed.) Proceedings of 1st European Conference on Grain
Legumes, Angers. AEP, Paris, pp. 99–103.
Jakel, L., Knosche, R. and Gunther, G. (1990) Cytological investigations on tisuue
culture of Pisum sativum L. and Vicia faba L. cultured on herbicide-containing
selection media. Biologisches Zentralblatt 109, 159–167.
James, C. (1997) Global Status of Transgenic Crops in 1997. ISAAA, Ithaca, New York.
Jane, J., Kasemsuvan, T., Chen, J.F. and Juliano, B.O. (1996) Phosphorus in rice and
other starches. Cereal Foods World 41, 827–832.
Jansman, A.J.M., Huisman, J. and van der Poel, A.F.B. (1993) Ileal and faecal
digestibility of field beans (Vicia faba L.) varying in tannin content. Animal Feed
Sciences and Technology 42, 83–96.
Jarkova, T.V., Deineko, E.V. and Shumny, V.K. (1998) Screening of pea (Pisum
sativum L.) collection effects on the morphogenetic capacity in tissue culture.
Plant Tissue Culture and Biothechnology 4, 175–182.
Jelaska, S., Pevalek, B., Papes, D. and Devide, Z. (1981) Developmental aspects of
long-term callus culture of Vicia faba L. Protoplasma 105, 285–292.
Jeltema, M.A. and Zabik, M.E. (1979) Fiber components-quantification and
relationship to cake quality. Journal of Food Science 44, 1732–1735.
289
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:32 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
274 References
Jeltema, M.A. and Zabik, M.E. (1980) Revised method for quantitative fibre
components. Journal of the Science of Food and Agriculture 31, 820–829.
Jenkins, D.J.A., Cuff, D., Wolever, T.M.S., Knowland, D., Thompson, L., Cohen, Z.
and Prokipchuk, E. (1987) Digestibility of carbohydrate foods in an
ileostomate: relationship to dietary fiber, in vitro digestibility, and glycemic
response. American Journal of Gastroenterology 82, 709–717.
Jesh, P. and Harzburg, B. (1997) Kartoffeln sind ihre Starke. Kartoffelnbau 9/10,
348–351.
Jia, S. (1982) Factors affecting the division frequency of pea mesophyll protoplasts.
Canadian Journal of Botany 60, 2192–2196.
Jia-Ping, Z., Roth, E.J., Terzaghi, W. and Lark, K.G. (1981) Isolation of sodium
dependent variants from haploid soybean cell culture. Plant Cell Reports 1,
48–51.
Jin, H., Hartman, G.L., Huang, Y.H., Nickell, C.D. and Widholm, J.M. (1996)
Regeneration of soybean plants from embryogenic suspension cultures treated
with toxic culture filtrate of Fusarium solani and screening of regenerants for
resistance. Phytopathology 86, 714–718.
Johansen, H.N., Glitse, V. and Bach Knudsen, K.E. (1996) Influence of extraction
solvent and temperature on quantitative determination of oligosaccharides
from plant materials by high-performance liquid chromatography. Journal of
Agricultural and Food Chemistry 44, 1470–1474.
Johansson, C.-G., Siljeström, M. and Asp, N.G. (1984) Dietary fibre in bread and
corresponding flours – formation of resistant starch during baking. Zeitschrift
für Lebensmittel Untersuchung und Forschung 179, 24–28.
John, M., Roehring, H., Smidt, J., Walden, R. and Schell, J. (1997) Cell signalling by
oligosaccharides. Current Trends in Plant Science 2, 111–115.
Jones, D.A., Barber, L.M., Arthur, A.E. and Hedley, C.L. (1995) An analysis of seed
development in Pisum sativum L. XVI. Assessing variation for fatty-acid content
by use of a non-destructive technique for single seed analysis. Plant Breeding
114, 81–83.
Jones, D.A., Urban, J. and Copikova, J. (1999a) A micro-analytical method for the
determination of starch and amylose/amylopectin content in pea seeds.
Biologia Plantarum 42, 303–308.
Jones, D.A., Dupont, M.S., Ambrose, M.J., Frias, J. and Hedley, C.L. (1999b) The
discovery of compositional variation for the raffinose family of oligosaccharides
in pea seeds. Seed Science Research 9, 305–310.
Jones, H., Karp, A. and Jones, M.K. (1989) Isolation, culture, and regeneration of
plants from potato protoplasts. Plant Cell Reports 8, 307–311.
Jones, R.G. (2000) Carbon partitioning into carbohydrates through pea seed
development. PhD thesis, University of East Anglia, UK.
Jood, S., Chauhan, B.M. and Kapoor, A.C. (1988) Contents and digestibility of
carbohydrates of chickpea and black gram as affected by domestic processing
and cooking. Food Chemistry 30, 113–127.
Jood, S., Mehta, U., Singh, R. and Bhat, C.M. (1985) Effect of processing on flatus
producing factors in legumes. Journal of Agricultural and Food Chemistry 33,
268–271.
Jorgensen, H., Zhao, X.O. and Eggum, B.O. (1996) The influence of dietary fibre
and environmental temperature on the development of the gastrointestinal
290
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:33 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 275
291
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:33 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
276 References
Kataki, P.K., Horbowicz, M., Taylor, A.G. and Obendorf, R.L. (1997) Changes in
sucrose, cyclitols and their galactosyl derivatives with seed ageing. In: Ellis,
R.H., Black, M., Murdoch, A.J. and Hong, T.D. (eds) Basic and Applied Aspects of
Seed Biology. Kluwer, Dordrecht, pp. 515–522.
Kawasaki, S. (1989) Extensin secreted into the culture medium by tobacco cells. I.
Purification and some properties. Plant Cell Physiology 30, 259–265.
Keijbets, M.J.H. and Pilnik, W. (1974) Some problems in the analysis of pectin in
potato tuber tissue. Potato Research 17, 169.
Kelkar, M., Shastri, P. and Rao, B.Y. (1996) Effect of processing on in vitro carbohy-
drate digestibility of cereals and legumes. Journal of Food Science and Technology,
India 33, 493–497.
Keller, F. and Ludlow, M.M. (1993) Carbohydrate metabolism in drought stressed
leaves of pigeon pea (Cajanus cajan). Journal of Experimental Botany 44,
1351–1359.
Kermode, A.R. (1997) Approaches to elucidate the basis of desiccation-tolerance in
seeds. Seed Science Research 7, 75–95.
Kerr, P.S. (1993) Soybean products with improved carbohydrate composition and
soybean plants. International Patent Publication Number WO 93/07742,
PCT/US/92/08958.
Kerr, P.S., Pearlstein, R.W., Becker-Manley, M.F. and Pierce, J.W. (1993) Nucleotide
sequences of galactinol synthase from zucchini and soybean. International
Patent Publication Number WO 93/02196, PCT/US92/06057.
Key, F.B. and Mathers, J.C. (1993) Complex carbohydrate and large bowel fermen-
tation in rats given wholemeal bread and cooked haricot beans (Phaseolus
vulgaris) fed in mixed diets. British Journal of Nutrition 69, 497–509.
Kim, H.-O. and Williams, P.C. (1990) Determination of starch and energy in feed
grains by near-infrared reflectance spectroscopy. Journal of Agricultural and Food
Chemistry 38, 682–688.
Kim, H.R., Hermansson, A.-M. and Eriksson, C.E. (1992) Structural characteristics
of hydroxypropyl potato starch granules depending on their molar substitu-
tion. Starch 44, 111–116.
Kim, W.J., Smit, C.J.B. and Nakayama, T.O.N. (1973) The removal of oligosaccha-
rides from soy beans. Lebensmittel Wissenschaft und Technologie 6, 201–204.
Kitagawa, I., Saito, M. and Taniayama, T. (1984a) Quantitative determination of
soyasaponins in soybeans of various origins and soybean products by means
of high-performance liquid chromatography. Yakagaku Zasshi 104, 275–279.
Kitagawa, I., Yoshikawa, M., Hayashi, T. and Taniayama, T. (1984b) Characteriza-
tion of saponin constituents in soybeans of various origins and quantitative
analysis of soyasaponins by gas-liquid chromatography. Yakagaku Zasshi 104,
162–168.
Klein, A.S., Montezinos, D. and Delmer, D.P. (1981) Cellulose and 1,3-glucan
synthesis during the early stages of wall regeneration in soybean protoplasts.
Planta 152, 105–114.
Kmita-Glazewska, H. and Kostyra, H. (1998) Chemical composition of soluble
and insoluble fractions of dietary fibre isolated from pea seeds. In: AEP (ed.)
Opportunities for High Quality, Healthy and Added-Value Crops to Meet European
Demands. Proceedings of 3rd European Conference on Grain Legumes, Valladolid.
AEP, Paris, p. 382.
292
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:34 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 277
293
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:35 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
278 References
294
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:35 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 279
Kurtzman, R.H. and Halbrook, W.U. (1970) Polysaccharides from dry navy beans,
Phaseolus vulgaris: its isolation and stimulation of Clostridium perfringens. Applied
Microbiology 20, 715–719.
Kvasnibka, F. (1995) Antinutritional saccharides in peas. Vyz. Potraviny 50, 3–4
(in Czech).
Kvasnibka, F., Ahmadová-Vavrousová, R., Mrskot, M., Velítek, J. and Kadlec, P.
(1994) Characteristics of pea varieties according to galactooligosaccharides
content. In: Fenwick, G.R., Hedley, C., Richards, R.L. and Khokhar, S. (eds)
Agricultural Food Quality. An Interdisciplinary Approach. The Royal Society of
Chemistry, Cambridge, pp. 153–156.
Kvasnibka, F., Ahmadová-Vavrousová, R., Frias, J., Price, K.R. and Kadlec, P. (1996)
A rapid HPLC method for determination of raffinose family oligosaccharides
in pea seeds. Journal of Liquid Chromatography 19, 135–147.
Kweon, M.R., Bhirud, P.R. and Sosulski, F.W. (1996) An aqueous alcoholic-alkaline
process of corn and pea starches. Starch/Stärke 48, 214–220.
Kysely, W. and Jacobsen, H.J. (1990) Somatic embryogenesis from pea embryos and
shoot apices. Plant Cell Tissue and Organ Culture 20, 7–14.
Kysely, W., Myers, J.R., Lazzeri, P.A., Collins, G.B. and Jacobsen, H.J. (1987) Plant
regeneration via somatic embryogenesis in pea (Pisum sativum L.). Plant Cell
Reports 6, 305–308.
Lacassagne, L., Francesch, M., Carre, B. and Melcion, J.P. (1988) Utilization of
tannin-containing and tannin-free faba beans (Vicia faba) by young chicks:
effects of pelleting feeds on energy, protein and starch digestibility. Animal Feed
Sciences and Technology 20, 59–68.
Lacassagne, L., Melcion, J.P., de Monredon, F. and Carre, B. (1991) The nutritional
values of faba bean flours varying in their mean particle size in young chickens.
Animal Feed Sciences and Technology 34, 11–19.
Lahuta, L.B., Górecki, R.J. and Rejowski, A. (1995) Accumulation of sugars in
maturing field bean (Vicia faba minor L.) and pea seeds (Pisum sativum L.) in
relation to seed viability. In: AEP (ed.) Improving Production and Utilisation of
Grain Legumes. Proceedings 2nd European Conference on Grain Legumes. AEP, Paris,
p. 44.
Lahuta, L.B., Jagielska, T., Górecki, R.J., Jones, A. and Hedley, C. (1998) Soluble
carbohydrates in desiccation tolerance of germinating pea seeds of different
isolines. In: AEP (ed.) Opportunities for High Quality, Healthy and Added-value
Crops to meet European Demands. Proceedings 3rd European Conference on Grain
Legumes, Valladolid. AEP, Paris, pp. 40–41.
Lai, F.-M. and McKersie, B.D. (1994) Regulation of starch and storage protein
accumulation in alfalfa (Medicago sativa L.) somatic embryos. Plant Science 100,
211–219.
Landgren, C.R. (1976) The influence of culture conditions on mitotic activity of
protoplats derived from Pisum root cortical explants. Protoplasma 87, 49–69.
Landgren, C.R. (1981) Gibberellic enhancement of the enzymatic release of Pisum
root cell protoplasts. Physiology Plantarum 52, 349–352.
Larkin, P.J. and Scowcroft, W.R. (1981) Somaclonal variation – a novel source of
variability from cell cultures for plant improvement. Theoretical and Applied
Genetics 60, 197–214.
295
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:36 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
280 References
Larsson, S., Johansson, L.A. and Svenningsson, M. (1993) Soluble sugars and mem-
brane lipids in winter wheats (Triticum aestivum L.) during cold acclimation.
European Journal of Agronomy 1, 85–90.
Latunde-Dada, A.O. and Lucas, J.A. (1983) Somaclonal variation and reaction to
Verticillium wilt in Medicago sativa L. plants regenerated from protoplast. Plant
Science Letters 32, 205–211.
Latunde-Dada, A.O. and Lucas, J.A. (1988) Somaclonal variation and resistance to
Verticillium wilt in lucerne, Medicago sativa L., plants regenerated from callus.
Plant Science 58, 111–119.
Leach, H.W., McCowen, L.D. and Schoch, T.J. (1959) Structure of the starch
granule. I. Swelling and solubility patterns of various starches. Cereal Chemistry
36, 534–544.
Le Bansky, B.P., McKnight, T.D. and Griffing, L.R. (1992) Purification and
characterization a secreted purple phosphatase from soybean suspension
cultures. Plant Physiology 99, 391–395.
Lee, E.Y.C. and Whelan, W.J. (1971) Glycogen and starch debranching enzymes. In:
Boyer, P.D. (ed.) The Enzymes, Vol. 5. Academic Press, New York, pp. 191–234.
Lee, S.C. and Prosky, L. (1994) Perspectives on new dietary fibre definition. Cereal
Foods World 39, 667.
Le Guen, M.-P., Peyronnet, C. and Gabon, G. (1997) Nitrogen ruminal
degradability of pea and lupin. Grain Legumes 17, 14–17.
Lehle, L. and Tanner, W. (1972) Synthesis of raffinose-type sugars. Methods in
Enzymology 28B, 522–530.
Lehmann, J. (1998) In: Carbohydrates Structure and Biology, translated by Haines, A.H.
Thieme, Stuttgart, p. 288.
Lehninger-Mertens, R. and Jacobsen, H.J. (1989a) Protoplast regeneration and
organogenesis from pea protoplasts. In Vitro Cellular and Developmental Biology –
Plant 25, 571–574.
Lehminger-Mertens, R. and Jacobsen, H.J. (1989b). Plant regeneration from pea
protoplasts via somatic embryogenesis. Plant Cell Reports 8, 379–382.
Leopold, A.C. and Vertucci, C.W. (1986) Physical attributes of desiccated seeds.
In: Leopold, A.C. (ed.) Membranes, Metabolism and Dry Organisms. Cornell
University Press, Ithaca, New York, pp. 22–34.
Leopold, A.C., Sun, W.Q. and Bernal-Lugo, I. (1994) The glassy state in seeds:
analysis and function. Seed Science Research 4, 267–274.
LePrince, O., Deltour, R., Thorpe, P.C., Atherton, N.M. and Hendry, G.A.F. (1990).
The role of free radicals and radical processing systems in loss of desiccation
tolerance in germinating maize (Zea mays L). New Phytologist 116, 573–580.
LePrince, O., Hendry, G.A.F. and McKersie, B.D. (1993) The mechanism of
desiccation tolerance in developing seeds. Seed Science Research 3, 231–246.
Leprince, O. and Walters-Vertucci, C. (1995) A calometric study of the glass transi-
tion behaviors in axes of bean seeds with relevance to storage stability. Plant
Physiology 109, 1471–1481.
Leske, K.L., Jevne, C.J. and Coon, C.N. (1993) Effect of oligosaccharide addition
on nitrogen-corrected true metabolizable energy of soy protein concentrate.
Poultry Science 72, 664–668.
Leske, K.L., Zhang, B. and Coon, C.N. (1995) The use of low alpha-galactoside
protein products as a protein source in chicken diets. Animal Feed Sciences and
Technology 54, 275–286.
296
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:36 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 281
Letzelter, N., Wilson, R., Jones, D.A. and Sinnaeve, G. (1995) Quantitative determi-
nation of the composition of individual pea seeds by Fourier Transform
infrared photoacoustic spectroscopy. Journal of Food Science and Agriculture 67,
239–245.
Lewandowicz, G. and Mczynski, M. (1990a) Chemiczna modyfikacja skrobi. Cz.I.
Modyfikacja skrobi ziemniaczanej. Chemik 1, 9–14.
Lewandowicz, G. and Mczynski, M. (1990b) Chemiczna modyfikacja skrobi. Cz.II.
Reaktywnosc skrobi róznych gatunków roslin. Chemik 3, 71–89.
Lewandowicz, G., Soral-Smietana, M. and Fornal, J. (1998) New RS preparations –
structure and physico-chemical properties. Proceedings of the VIII Inter-
national Starch Convention, Cracow (Poland), 16–19 June.
Li, B.W. and Schuhmann, P.J. (1980) Gas–liquid chromatography analysis of sugars
in ready-to-eat breakfast cereals. Journal of Food Sciences 45, 138–141.
Li, B.W., Schuhmann, P.J. and Wolf, W.J. (1985) Chromatographic determination
of sugars and starch in a diet composite reference material. Journal of Agricul-
tural and Food Chemistry 33, 531–536.
Libbenga, K.R. and Torrea, J.G. (1973) Hormone-induced endoreduplication prior
to mitosis in cultured pea root cortex cells. American Journal of Botany 60,
293–299.
Lichner, F.T. and Spanswick, R.M. (1981) Electrogenic sucrose transport in devel-
oping soybean cotyledons. Plant Physiology 67, 869–874.
Lim, H.T., Song, Y.N. and Yeoung, Y.R. (1994) Plant regeneration from immature
embryo and thin layer of pea (Pisum sativum L.) via callus induction, somatic
embryogenesis and organogenesis. Korean Journal of Breeding 26, 33–40.
Lin, T.-P. and Huang, N.-H. (1994) The relationship between carbohydrate compo-
sition of some tree seeds and their longevity. Journal of Experimental Botany 45,
1289–1294.
Lin, W. (1985a) Energetics of sucrose transport into protoplasts from developing
soybean cotyledons. Plant Physiology 78, 41–45.
Lin, W. (1985b) Linear sucrose transport in protoplasts from developing soybean
cotyledons. Plant Physiology 78, 649–651.
Lin, W., Schmitt, M.R., Hitz, W.D. and Giaquinta, R.T. (1984) Sugar transport into
protoplasts isolated from developing soybean cotyledons. Plant Physiology 75,
936–940.
Lin, W., Schmitt, M.R., Hitz, W.D. and Giaquinta, R.T. (1984) Sugar transport
into protoplasts isolated from developing soybean cotyledons. II. Protoplast
isolation and general characteristics of sugar transport. Plant Physiology 75,
936–940.
Lintas, C., Cappelloni, M. and Ruggeri, S. (1992) Processing and legume poly-
saccharides. In: AEP (ed.) Proceedings of 1st European Conference on Grain Legumes.
AEP, Paris, pp. 441–442.
List, G.R., Mounts, T.L., Orthoefer, F. and Neff, W.E. (1996) Potential margarine
oils from genetically modified soybeans. Journal of the American Oil Chemists
Society 73, 729–732.
Liu, J.J., Odegard, W. and de Lumen, B.O. (1995) Galactinol synthase from kidney
bean cotyledon and zucchini leaf. Plant Physiology 109, 505–511.
Lizotte, P.A., Henson, C.A. and Duke, S.H. (1990) Purification and characterization
of pea epicotyl α-amylase. Plant Physiology 92, 615–621.
297
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:37 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
282 References
Lloyd, J.R., Bull, V.J., Hedley, C.L. and Ring, S.G. (1996a) Determination of the
effect of r and rb mutations on the structure of amylose and amylopectin in pea
(Pisum sativum L.). Carbohydrate Polymers 29, 45–49.
Lloyd, J.R., Wang, T.L. and Hedley, C.L. (1996b) An analysis of seed development
in Pisum sativum XIX. Effect of mutant alleles at the r and rb loci on starch grain
size and on the content and composition of starch in developing pea seeds.
Journal of Experimental Botany 47, 171–180.
Locker, A. and Ilan, I. (1975) On the nature of the hormonal regulation of amylase
activity in cotyledons of germinating peas. Plant Cell Physiology 16, 449–454.
Loewus, F.A. (1990) Cyclitols. In: Dey, P.M. and Harbourne, J.B. (eds) Methods in
Plant Biochemistry, Vol. 2. Carbohydrates. Academic Press, New York, pp. 219–233.
Loewus, F. and Dickinson, D.B. (1982) Cyclitols. In: Loewus, F.A. and Tanner, W.
(eds) Encyclopaedia of Plant Physiology: Plant Carbohydrates 1: Intracellular Carbohy-
drates, Vol. 13A. Springer-Verlag, Berlin, pp. 193–206.
Loewus, F. and Loewus, M. (1980). Biochemistry of myo-inositol. In: Stumpf, P. and
Conn, E. (eds) Biochemistry of Plants, Vol. 3. Academic Press, New York,
pp. 43–76.
Loewus, F.A. and Loewus, M.W. (1983) myo-Inositol: its biosynthesis and metabo-
lism. Annual Review of Plant Physiology 34, 137–161.
Login, A., Piotrowicz-Cieslak, A. and Górecki, R.J. (1995) α-Galactosidase in
maturing and germinating yellow lupine seeds. In: Come, D., Duczmal, K.,
Górecki, R., Grzesik, M. and Lewak, S. (eds) Third French–Polish Symposium
‘Current problems of seed physiology’ 18–20 July 1995. ART, Olsztyn, Poland, p. 21.
Loiseau, J., Marche, C. and Deunff, Y. Le. (1995) Effect of auxins, cytokinins, carbo-
hydrates and aminoacids on somatic embryogenesis induction from shoot
apices of pea. Plant Cell Tissue and Organ Culture 33, 267–275.
Longstaff, M. and McNab, J.M. (1987) Digestion of starch and fibre carbohydrates
in peas by adult cockerels. British Poultry Science 28, 261–285.
Lorberth, R., Ritte, G., Willmitzer, L. and Kossmann, J. (1998) Inhibition of a
starch-granule-bound protein leads to modified starch and repression of cold
sweetening. Nature Biotechnology 16, 473–477.
Lorenz, K. (1979) The starch of the faba bean (Vicia faba). Comparison with wheat
and corn starch. Starch 31, 181–184.
Lorz, H. and Brown, P.T.H. (1986) Variability in tissue culture derived plants –
possible origins, advantages and drawbacks. In: Horn, W. (eds) Genetic Manipu-
lation in Plant Breeding. Walter de Gruyter Publishers, Berlin, pp. 513–534.
Low, A.G. (1985) Role of dietary fibre in pigs diets. In: Haresign, H. and Cole, D.J.A.
(eds) Recent Advances in Animal Nutrition. Butterworths, London, pp. 87–112.
Lowell, C.A. and Kuo, T.M. (1989) Oligosaccharide metabolism and accumulation
in developing soybean seeds. Crop Science 29, 459–465.
Lozovaya, V.V., Zabotina, O.A. and Widholm, I.M. (1996) Synthesis and turnover of
cell wall polysaccharides and starch in photosynthetic soybean suspension
cultures. Plant Physiology 111, 921–929.
Lu, D.Y., Cooper-Bland, S., Pental, D., Cocking, E.C. and Davey, M.R. (1983) Isola-
tion and sustained division of protoplasts from cotyledons of seedlings and
immature seeds of Glycine max L. Zeitschrift für Pflanzenphysiologie 111, 389–394.
Lu, H., Gai, J. and Ma, Y. (1993) Soybean protoplast culture under different
hormone conditions and plantlet regeneration. Acta Agronomica Sinica 19,
328–333.
298
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:38 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 283
Lutova, L.A. and Zabelina, Ye.K. (1988) Callus and shoot in vitro formation in differ-
ent forms of peas Pisum sativum L. Genetics (Moscow) 24, 1632–1640.
Lyons, A.J., Pridham, T.G. and Hesseltine, C.W. (1969) Survey of some Actino-
mycetales for α-galactosidase activity. Applied Microbiology 18, 579–583.
MacKintosh, C., Lyon, G.D. and MacKintosh, R.W. (1994) Protein phosphatase
inhibitors activate antifungal defence responses of soybean cotyledons and cell
cultures. Plant Journal 5, 137–147.
Macleod, M.R., Hedley, C.L., Martin, C.R. and Wang, T.L. (1991) Genetic analysis
of new wrinkled-seeded pea mutants at the r locus. Aspects of Applied Biology 27,
263–265.
Madhusudhan, M. and Tharanathan, R.N. (1995) Legume and cereal starches –
why differences in digestibility? Part 1: Isolation and composition of legume
(Greengram and Bengalgram) starches. Starch 49, 165–171.
Madhusudhan, M. and Tharanathan, R.N. (1996) Structural studies of linear and
branched fractions of chick pea and finger millet starches. Carbohydrate Research
284, 101–109.
Makkar, H.P.S., Becker, K., Abel, H. and Pawezik, E. (1997) Nutrients content,
rumen protein degradability and nutritional factors in some colour- and
white-flowering cultivars of Vicia faba beans. Journal of the Sciences Food and
Agriculture 75, 511–520.
Malik, K.A. and Saxena, P.K. (1991) Regeneration in Phaseolus vulgaris L. Promotive
role of N(6)-benzylaminopurine in cultures from juvenile leaves. Planta 184,
148–150.
Malik, K.A. and Saxena, P.K. (1992a) Somatic embryogenesis and shoot regenera-
tion from intact seedlings of Phaseolus vulgaris acutifolius A., P. aureus (L.)
Wilczek, P. coccineus L. and P. wrightii L. Plant Cell Reports 11, 163–168.
Malik, K.A. and Saxena, P.K. (1992b) Thidiazuron induced high frequency
shoot regeneration in intact seedlings of pea (Pisum sativum), chickpea (Cicer
aratinum) and lentil (Lens culinaris). Australian Journal of Plant Physiology 19,
731–740.
Malik, K.A. and Saxena, P.K. (1992c) Regeneration in Phaseolus vulgaris L.: High
frequency induction of direct shoot formation in intact seedlings by
N(6)-benzylaminopurine and thidiazuron. Planta 186, 384–389.
Mallee, F.M. (1995) Starch-based texturizing agent. US Patent 5470391.
Mallee, F.M., Stone, J.A. and Finocchiaro, E.T. (1996) Starch-based texturizing
agent. US Patent 554751.
Malmberg, R.L. (1979) Regeneration of whole plants from callus cultures of diverse
genetic lines of Pisum sativum L. Planta 146, 243–244.
Maltese, S., Atta, S. and Cousin, R. (1995) Determination of pea seed composition
by near infrared transmission spectroscopy. In: AEP (ed.) Improving Production
and Utilisation of Grain Legumes. Proceedings of 2nd European Conference on Grain
Legumes, Copenhagen. AEP, Paris, p. 372.
Mangat, B.S., Pelekis, M.K. and Cassells, A.C. (1990) Changes in the starch content
during organogenesis in in vitro cultured Begonia rex stem explants. Physiology
Plantarum 79, 267–274.
Manners, D.J. (1989) Review paper. Recent development in our understanding of
amylopectin structure. Carbohydrate Polymers 11, 87–112.
Marks, G.E. and Davies, D.R. (1979) The cytology of cotyledon cells and the induc-
tion of giant polytene chromosomes in Pisum sativum L. Protoplasma 101, 73–80.
299
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:39 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
284 References
Marshall, J., Sidebottom, C., Debet, M., Martin, C., Smith, A. and Edwards, A.
(1996) Identification of the major starch synthase in the soluble fraction of
potato tubers. Plant Cell 8, 1121–1135.
Martin, C. and Smith, A.M. (1995) Starch biosynthesis. The Plant Cell 7, 971–985.
Matthews, R.H. (1989) Legumes Chemistry, Technology and Human Nutrition. Marcel
Dekker, New York.
Mathews, V.H., Rao, P.S. and Bhatia, C.R. (1986) Somaclonal variation in cotyledon-
ary plants of mung bean. Zeitschrift für Pflanzenzucht 96, 169–173
Mauch, F. and Staehelin, L.A. (1989) Functional implications of the subcellular
localization of ethylene-induced chitinase and glucanase in bean leaves. Plant
Cell 1, 447–457.
Mauch, F., Mauch Mani, B. and Boller, T. (1988) Antifungal hydrolases in pea
tissue: II. Inhibition of fungal growth by combination of chitinase and
glucanase. Plant Physiology 88, 936–942.
Mazur, A., Remesy, C., Gueux, E., Levrat, A.-M. and Deminged, Ch. (1990) Effect of
diet rich in fermentable carbohydrates on plasma lipids levels and on lipo-
protein catabolism in rats. Journal of Nutrition 120, 1037–1045.
Margareta, E., Nyman, G.L., Svanberg, M.S.J. and Asp, N.G. (1994) Molecular
weight distribution and viscosity of water-soluble dietary fibre isolated from
green beans, brussels sprouts and green peas following different types of
processing. Journal of the Science of Food and Agriculture 66, 83–91.
McCabe, D.E., Swain, W.F., Martinell, B.J. and Christou, P. (1988) Stable trans-
formation of soybean (Glycine max) by particle acceleration. Bio-Technology (New
York) 6, 923–926.
McCann, M.C. and Roberts, K. (1994) Changes in cell wall architecture during cell
elongation. Journal of Experimental Botany 45 (special issue), 1683–1691.
McCutchen, C., Duffaud, G., Leduc, P., Peterson, A., Tayal, A., Khan, S. and Kelly,
R. (1996) Characterization of extremely thermostable enzymatic breakers
(alactosidase and annanase) from the hyperthermophilic bacterium Thermo-
toga neapolitana 5068 for hydrolysis of guar gum. Biotechnology Bioengineering 52,
332–339.
McDougall, G.J., Morrison, I.M., Stewart, D., Weyers, J.D.B. and Hillman, J.R.
(1993) Plant fibres: botany, chemistry and processing for industrial use. Journal
of the Science of Food and Agriculture 62, 1–20.
McIntyre, A., Gibson, P.R. and Young, G.P. (1993) Butyrate production from dietary
fibre and protection against large bowel cancer in a rat model. Gut 34, 386–391.
McKee, H.R., Robertson, R.N. and Lee, J.B. (1955) Physiology of the pea fruits. I.
The developing fruit. Australian Journal of Biological Sciences 8, 137–162.
McKently, A.H., Moore, G.A. and Garden, F.P. (1990) In vitro regeneration of
peanut. Crop Science 30, 192–196.
McKentley, A.H., Moore, G.A., Doostdar, H. and Niedz, R.P. (1995) Agrobacterium-
mediated transformation of peanut (Arachis hypogaea L.) embryo axes and the
development of transgenic plants. Plant Cell Reports 14, 699–703.
Mercier, C. (1979) Les α-galactosides des graines de leguminouses. In: INRA (ed.)
Les Materies Premiers et Alimentation des Volailess. Seances de Travail, Paris,
pp. 79–90.
Meredith, F.I., Thomas, C.A., Snook, M.E., Himmelsbach, D.S. and van Halbeek, H.
(1988) Soluble carbohydrates oligosaccharides and starch in lima bean seeds.
Journal of Food Science 53, 768–771.
300
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:39 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 285
301
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:40 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
286 References
sunflower seed albumine gene. Proceedings of the National Academy of Sciences USA
94, 8393–8398.
Monerri, C., Garcia Luis, A. and Guardiola, J.L. (1986) Sugar and starch changes in
pea cotyledons during germination. Physiologia Plantarum 67, 49–54.
Mongeau, R. and Brassard, R. (1994) Comparison and assessment of the difference
in total dietary fibre in cooked dry legumes as determined by five methods.
Journal AOAC International 77, 1197–1202.
Mongeau, R. and Brassard, R. (1995) Importance of cooking temperature and
pancreatic amylase in determination of dietary fibre in dried legumes. Journal
AOAC International 78, 1444–1449.
Monma, M., Sugimoto, T., Monma, M., Kawamura, Y. and Saio, K. (1991) Starch
breakdown in developing soybean seeds (Glycine max cv. Enrei). Agriculture
Biology Chemistry 55, 67–71.
Moore, P.J., Swords, K.M.M., Lynch, M.A. and Staehelin, L.A. (1991) Spatial
reorganization of the assembly pathways of glycoproteins and complex
polysaccharides in the Golgi apparatus of plants. Journal of Cell Biology 112,
589–602.
Morohashi, Y., Katoh, H., Kaneko, Y. and Matsushima, H. (1989) Control of α-
amylase development in cotyledons during and following germination of mung
bean seeds. Plant Physiology 91, 253–258.
Morris, P., Scragg, A.H., Smart, N.J. and Stafford, A. (1985) Secondary product
formation by cell suspension cultures. In: Dixon, R.A. (ed.) Plant Cell Culture – a
Practical Approach. IRL Press, Oxford, pp. 127–134.
Morris, V.J. and Miles, M.J. (1994) Birefringent techniques. In: Ross-Murphy, S.B.
(ed.) Physical Techniques for the Study of Food Polymers. Blackie Academic &
Professional, London, pp. 215–275.
Morrison, I.M. (1972) Improvements in the acetyl bromide technique to determine
lignin and digestibility and its application to legumes. Journal of the Science of
Food and Agriculture 23, 1463–1469.
Morrison, W.R. and Karkalas, J. (1990) Starch. In: Dey, P.M. (ed.) Methods in Plant
Biochemistry, Vol. 2. Academic Press, San Diego, pp. 323–352.
Moscoso, W., Rourne, M.C. and Hood, L.F. (1984) Relationships between the
hard-to-cook phenomenon in red kidney beans and water absorption,
puncture force, pectin, phytic acid, and minerals. Journal of Food Science 49,
1577–1583.
Mroginski, L.A. and Kartha, K.K. (1981) Regeneration of pea (Pisum sativum L. cv.
Century) plants by in vitro culture of immature leaflets. Plant Cell Reports 1,
64–66.
Mueller-Roeber, B., Kossmann, J., Hannah, L.C., Willmitzer, L. and Sonnerwald, U.
(1990) Only one of two different ADP-glucose pyrophosphorylase genes from
potato responds strongly to elevated levels of sucrose. Molecular General Genetics
224, 136–146.
Mueller-Roeber, B., Sonnerwald, U. and Willmitzer, L. (1992) Inhibition of ADP-
glucose pyrophosphorylase in transgenic potatos leads to sugar-storing tubers
and influences tuber formation and expression of tuber storage protein genes.
EMBO Journal 11, 1229–1238.
Muir, J.G. and O’Dea, K. (1992) Measurement of resistant starch: factors affecting
the amount of starch escaping digestion in vitro. American Journal of Clinical
Nutrition 56, 123–127.
302
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:40 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 287
Muir, J.G. and O’Dea, K. (1993) Validation of an in vitro assay for predication the
amount of starch that escapes digestion in the small intestine of humans.
American Journal of Clinical Nutrition 57, 540–546.
Muir, J.G., Birkett, A., Brown, I., Jones, G. and O’Dea, K. (1995) Food processing
and maize variety affects amounts of starch escaping digestion in the small
intestine. American Journal of Clinical Nutrition 61, 82–89.
Müller, H.L., Kirchgessner, M. and Roth, F.X. (1989) Energy utilisation of
intracaecally infused carbohydrates and casein in sows. In: Van der Honning, Y.
and Close, W.H. (eds) Energy Metabolism of Farm Animals. Pudoc, Wageningen,
pp. 123–126.
Muller-Rober, B. and Kossmann, J. (1994) Approaches to influence starch quantity
and starch quality in transgenic plants. Plant Cell Environment 17, 601–613.
Munster, van I.P., Tangerman, A. and Nagengast, F.M. (1994) Effect of resistant
starch on colonic fermentation, bile acid metabolism, and mucosal pro-
liferation. Digestive Diseases and Sciences 39, 834–842.
Murashige, T. and Skoog, F. (1962) A revised medium for rapid growth and
bioassays with tobacco tissue cultures. Physiologia Plantarum 15, 473–497.
Murch, S.J. and Saxena, P.K. (1997) Modulation of mineral and free fatty acid pro-
files during thidiazuron mediated somatic embryogenesis in peanut (Arachis
hypogeae L.). Journal of Plant Physiology 151, 358–361.
Murthy, B.N.S., Victor, J., Singh, R.P., Fletcher, R.A. and Saxena, P.K. (1996). In
vitro regeneration of chickpea (Cicer arietinum L.): stimulation of direct
organogenesis and somatic embryogenesis by thidiazuron. Plant Growth
Regulation 19, 233–240.
Mutaftschiev, S., Macaya, A., Prat, R., Devillers, P. and Golberg, R. (1993) Early
effects of plant cell wall fragments on plant cell growth. Plant Physiology and
Biochemistry 31, 459–467.
Muthukumar, B., Mariamma, M., Veluthambi, K. and Gnanam, A. (1996) Genetic
transformation of cotyledon explants of cowpea (Vigna unguiculata L. Walp.)
using Agrobacterium tumefaciens. Plant Cell Reports 15, 980–985.
Muzquiz, M., Rey, C., Cuadrado, C. and Fenwick, G.R. (1992) Effect of germination on
the oligosaccharide content of lupin species. Journal of Chromatography 607, 349–352.
Muzquiz, M., Burbano, C., Pedrosa, M., Folkman, W. and Gulewicz, K. (1999) Lupin
as potential source of raffinose family oligosaccharides. Preparative method for
their isolation and purification. Industrial Crops and Products 19, 183–188.
Naczk, M., Myhara, R.M. and Shahidi, F. (1992) Effects of processing on oligo-
saccharides of oilseed and legume protein meals. Food Chemistry 45, 193–197.
Naczk, M., Amarowicz, R. and Shahidi, F. (1997) α-Galactosides of sucrose in food:
composition, flatulence-causing effect, and removal. In: Shahidi F. (ed.)
Antinutrients and Phytochemical in Food. ACS, Washington, DC, Series 662,
pp. 127–151.
Naivikul, O. and D’Appolonia, B.L. (1978) Comparation of legumes and wheat
flour carbohydrates. I. Sugar analysis. Cereal Chemistry 55, 913–918.
Natali, L. and Cavalini, A. (1987a) Regeneration of pea (Pisum sativum L.) plantlets
by in vitro culture of immature embryos. Plant Breeding 99, 172–176.
Natali, L. and Cavalini, A. (1987b) Nuclear cytology of callus and plantlets regener-
ated from pea (Pisum sativum L.) meristems. Protoplasma 141, 121–125.
Natali, L. and Cavalini, A. (1989) Grafting in vitro regenerated pea (Pisum sativum
L.) shoots as a method for obtaining mature plants. Plant Breeding 102, 341–343.
303
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:41 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
288 References
Nauerby, B., Madsen, M., Christiansen, J. and Wyndaele, R. (1991) A rapid and effi-
cient regeneration system for pea (Pisum sativum), suitable for transformation.
Plant Cell Reports 9, 676–679.
Nawracal, J., Konieczny, G. and Kazmierczak, M. (1996) Regeneration of soybean
plants from embryonic axes. Soybean Genetics Newsletter 23, 117–119.
Nche, M.I.R., Nout, M.J.R. and Rombouts, F.M. (1994) Fast production by
Clostridium perfringens as a measure of the fermentability of carbohydrates and
processed cereal- legume foods. Food Microbiology 11, 21–29.
Nelke, M., Nowak, J., Wright, J.M. and McLean, N.L. (1993) DNA fingerprinting of
red clover (Trifolium pratense L.) with Jeffrey’s probes: detection of somaclonal
variation and other applications. Plant Cell Reports 13, 72–78.
Nestares, T., Urbano, G., Lopez-Frias, M. and Barrionuevo, M.J. (1997) Nutritional
assessment of magnesium from raw and processed chickpea (Cicer arietinum L.)
in growing rats. Journal of Agriculture Food Chemistry 45, 3138–3142.
Newell, C.A. and Luu, H.T. (1985) Protoplast culture and plant regeneration in
Glycine canescence F. J. Herm. Plant Cell Tissue and Organ Culture 4, 145–149.
Nicolas, P., Gertsch, I. and Parisod, C. (1984) Isolation and structure determination
of an α-D-galactosyl-α-D-galactosyl-α-D-galactosyl-D-pinitol from the chickpea.
Carbohydrate Research 131, 331–334.
Nielsen, S.V.A., Poulsen, G.B. and Larsen, M.E. (1991) Regeneration of shoots from
pea hypocotyl explants. Physiologia Plantarum 82, 99–102.
Nisbet, G.S. and Webb, K.J. (1990) Transformation in legumes. In: Bajaj, Y.P.S.
(ed.) Biotechnology in Agriculture and Forestry, Vol. 10. Legumes and Oilseed Crops 1.
Springer-Verlag, Berlin, pp. 38–48.
Nnanna, I.A. and Phillips, R.D. (1988) Changes in oligosaccharide content, enzyme
activities and dry matter during controlled germination of cowpeas (Vigna
unguiculata). Journal of Food Science 53 (6), 1782–1786.
Noah, L., Guillon, F., Bouchet, B., Buleon, A., Gallant, D.J., Colonna, P., Molis, C.,
Faisant, N., Gelmiche, J.P. and Champ, M. (1995) Digestion of carbohydrate
components of dry beans (Phaseolus vulgaris) in healthy humans. In: Proceedings
of the 2nd AEP Conference Improving Production and Utilisation of Grain Legumes,
Copenhagen. AEP, Paris, pp. 276–277.
Noah, L., Guillon, F., Bouchet, B., Buleon, A., Molis, Ch., Gratas, M. and Champ, M.
(1998) Digestion of carbohydrate from white beans (Phaseolus vulgaris L.)
in healthy humans. Journal of Nutrition 128, 977–985.
Nowak, J. and Steinkraus, K.H. (1988) Effect of tempen fermentation of peas on
their potential flatulence productivity as measured by gas production and
growth of Clostridium perfringens. Nutrition Report International 38, 1163–1171.
Nwokolo, E. and Smartt, J. (eds) (1996) Food and Feed from Legumes and Oilseeds.
Chapman & Hall, London.
Obendorf, R.L. (1997) Oligosaccharides and galactosyl cyclitols in seed desiccation
tolerance. Seed Science Research 7, 63–74.
Obendorf, R.L., Horbowicz, M. and Taylor, D.P. (1993) Structure and chemical
composition of developing buckwheat seed. In: Janick, J. and Simon, J.E. (eds)
New Crops. John Wiley & Sons, New York, pp. 244–251.
Obendorf, R.L., Moon, H., Hildebrand, D.F., Torisky, R. and Collins, G.B. (1996) A
comparison of pinitols in somatic and zygotic soybean embryos (Abstract).
Molecular and Cellular Biology of the Soybean 6, 40.
304
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:42 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 289
Obendorf, R.L., Horbowicz, M., Dickerman, A.M., Brenac, P. and Smith, E. (1998)
Soluble oligosaccharides and galactosyl cyclitols in maturing soybean seeds in
planta and in vitro. Crop Science 38, 78–84.
Ochatt, S., Pasquis, B., Pontecaille, C. and Rancillac, M. (1998) Rapid plant regener-
ation from juvenile explants of protein legumes. In: AEP (ed.) Opportunities for
High Quality, Healthy and Added-Value Crops to Meet European Demands. Proceedings
3rd European Conference on Grain Legumes, Valladolid. AEP, Paris, p. 367.
Odunfa, S.A. (1983) Carbohydrate changes in fermenting locust bean (Parkia
filicoidea) during iru fermentation. Qualitas Plantarum Plant Foods and Human
Nutrition 32, 3–10.
Oelck, M.M., Rao, P.S. and Ozias-Akins, P. (1983) Protoplast regeneration from
some legume species. Experimentia (supplement) 45, 50–51.
Ogawa, K., Watanabe, T., Ikeda, Y. and Kondo, S. (1997) A new glycoside, 1D-2-O-α-
D-galactopyranosyl-chiro-inositol from jojoba beans. Carbohydrate Research 302,
219–221.
Ogura, H. (1982) Studies on the genetic instability of cultured tissues and the
regenerated plants – effects of auxins and cytokinins on mitosis of Vicia faba
cells. In: Fujiwara, A. (ed.) Plant Tissue Culture. Maruzen, Tokyo, pp. 433–434.
Ogura, H. (1990) Chromosome variation in plant tissue culture. In: Bajaj, Y.P.S.
(ed.) Biotechnology in Agriculture and Forestry, Vol. 11. Somaclonal Variation in Crop
Improvement 1. Springer-Verlag, Berlin, pp. 49–84.
Okamura, J.K., Jofuku, K.D. and Goldberg, R.B. (1986) Soybean seed lectin
gene and flanking nonseed protein genes are developmentally regulated in
transformed tobacco plants. Proceedings of the National Academy of Sciences USA
89, 8240–8244.
Oku, T. (1994) Special physiological functions of newly development mono-
and oligosaccharides. In: Goldberg, I. (ed.) Functional Foods – Designer Foods,
Pharmafoods, Nutraceuticals. Chapman & Hall, New York, pp. 202–218.
Olive, M.R., Ellis, R.J. and Schuch, W.W. (1989) Isolation and nucleotide sequences
of cDNA clones encoding ADP-glucose pyrophosphorylase polypepetides from
wheat leaf and endosperm. Plant Molecular Biology 12, 525–538.
Olson, A.C., Evans J.J., Frederick, D.P. and Jansen, E.F. (1969) Plant suspension
culture media macromolecules. Pectic substances, protein and peroxidase.
Plant Physiology 44, 1594–1600.
Ondøej, M., Rakousky, S. and Schutzner, K. (1998) Transgenic crops tolerant to
herbicides. Czechoslovak Journal of Genetics and Plant Breeding 34, 31–42.
Orman, B.A. and Schumann, R.A. (1991) Comparison of near-infrared spectros-
copy calibration methods for the prediction of protein, oil and starch in maize
grain. Journal of Agricultural and Food Chemistry 39, 883–886.
Oruna-Concha, M.J., Gonzalez-Castro, M.J., Lopez-Hernandez, J. and Simal-Lozano,
J. (1996) Evolution of sugar, starch, pectin and insoluble fibre contents in
frozen green beans and padron peppers. Deutsche Lebensmittel-Rundschau 92,
278–281.
Ozcan, S., Barghchi, M., Firek, S. and Draper, J. (1992) High frequency adventitious
shoot regeneration from immature cotyledons of pea (Pisum sativum L.). Plant
Cell Reports 11, 44–47.
Ozias-Akins, P., Schnall, J.A., Anderson, W.F., Singsit, Ch., Clemente, T.E., Adang,
M.J. and Weissinger, A.K. (1993) Regeneration of transgenic peanut plants
from stably transformed embryogenic callus. Plant Science 93, 185–194.
305
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:42 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
290 References
Padgette, S.R., Kolacz, K.H., Dellannay, X., Re, D.B., LaVallee, B.J., Tinius, C.N.,
Rhodes, W.K., Otero, Y.I., Barry, G.F., Eichholtz, D.A., Peschke, V.M.,
Nida, D.L., Taylor, N.B. and Kishore, G.M. (1995) Development, identification
and characterization of a glyphosate soybean line. Crop Science 35, 1451–1461.
Pahl, H. (1998) Potential and constraints in production and end-uses of grain
legumes in Europe – grain legumes and economics. In: AEP (ed.) Opportunities
for High Quality, Health and Added-value Crops to Meet European Demands.
Proceedings of 3rd European Conference on Grain Legumes. AEP, Paris, pp. 3–5.
Pandey, R. and Ganapathy, P.S. (1984) Isolation of sodium chloride-tolerant callus
line of Cicer arietinum L. cv. BG-203. Plant Cell Reports 3, 45–47.
Papes, D., Jelaska, S., Tomaseo, M. and Devide, Z. (1978) Triploidy in callus culture
of Vicia faba L. investigated by the Giemsa C-banding technique. Experientia 34,
1016–1017.
Paredes-Lopez, O., Maza-Calvino, E.C., Gonzalez-Castaneda, J. and Montes-Rivera,
R. (1988) Starch 40, 11–15.
Parrott, W.A., Hoffman, L.M., Hildebrand, D.F., Williams, E.G. and Collins, G.B.
(1989) Recovery of primary transformants of soybean. Plant Cell Reports 7,
615–617.
Parrott, W.A., Bailey, M.A., Durham, R.E. and Mathew, H.V. (1992) Tissue culture
and regeneration in Legumes. In: Moss, J.P. (ed.) Biotechnology and Crop
Improvement in Legumes. ICRISAT, Patancheru, India, pp. 115–148.
Parrott, W.A., Merkle, S.A. and Williams, E.G. (1993) Somatic embryogenesis:
potential for use in propagation and gene transfer systems. In: Murray, D.R.
(ed.) Advanced Methods in Plant Breeding and Biotechnology. CAB International,
Wallingford, UK, pp. 58–200.
Parrott, W.A., All, J.N., Adang, M.J., Bailey, M.A., Boerma, H.R. and Stewart, C.N. Jr.
(1994) Recovery and evaluation of soybean plants transgenic for a Bacillus
thuringiensis var. Kurstaki insecticidal gene. In Vitro Cellular and Developmental
Biology – Plant 30, 144–149.
Paszkowski, J., Peterhans, A., Schlupmann, H., Basse, C., Lebel, E.G. and Masson, J.
(1992) Protoplasts as tool for plant genome modifications. Physiologia
Plantarum 85, 352–356.
Pate, J.S. (1984) The carbon and nitrogen nutrition of fruits and seed-case studies
of selected grain legumes. In: Murray, D.R. (ed.) Seed Physiology, Vol. 1. Develop-
ment. Academic Press, London, pp. 41–82.
Pate, J.S., Sharkey, P.J. and Lewis, O.A.M. (1974) Phloem bleeding from legume
fruits – a technique for study of fruit nutrition. Planta 120, 229–243.
Patel, A.R., Patel, M.R., Patel, N.R., Patel, K.C. and Patel, R.D. (1986) Graft copoly-
mers of starch and polyacrylonitrile. Effect of source. Starch/Stärke 38, 235–237.
Patrick, J.W. and McDonald, R. (1980) Pathway of carbon transport within
developing ovules of Phaseolus vulgaris L. Australian Journal of Plant Physiology
7, 671–684.
Patrick, J.W. and Offler, C.E. (1995) Post-sieve element transport of sucrose in
developing seeds. Australian Journal of Plant Physiology 22, 681–702.
Patzelt, W.J. (1974) Polarized-light Microscopy. Principles, Instruments, Applications.
Ernst Leitz, Wetzlar.
Paul, A.A. and Southgate, D.A.T. (1985) In: Widdowson, E.M. (ed.) The Composition
of Foods, 4th edn. Elsevier/North-Holland Biomedical Press, Amsterdam.
306
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:43 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 291
Pearson, D. (1976) Sugar and preserves. In: Pearson, D. (ed.) The Chemical Analysis
of Food. Churchill Livingstone, Edinburgh, pp. 107–157.
Pedroso, M.C. and Pais, M.S. (1995) Factors controlling somatic embryogenesis.
Plant Cell Tissue and Organ Culture 43, 147–154.
Penel, K. and Greppin, H. (1996) Pectin binding proteins: characterization of the
binding and comparison with heparin. Plant Physiology and Biochemistry 34,
479–488.
Perez, S. and Imberty, A. (1996) Structural features of starch. Carbohydrates in Europe
15, 17–21.
Periago, M.J., Ros, G. and Casas, J.L. (1997) Non-starch polysaccharides and in vitro
starch digestibility of raw and cooked chick peas. Journal of Food Science 62,
93–96.
Petek, F., Villarroya, E. and Courtois, J.E. (1966) Isolation of two galactosides of
myo-inositol from vetch seeds. Comptes Rendus de l’Academie des Sciences (Paris)
Serie D: Sciences Naturelles 263, 195–197.
Petek, F., Villarroya, E. and Courtois, J.E., (1969) Purification and properties of
α-galactosidase in germinating Vicia sativa seeds. European Journal of Biochemistry
8, 395–402.
Peterbauer, T. and Richter, A. (1998) Galactosylononitol and stachyose synthesis in
seeds of adzuki bean. Purification and characterization of stachyose synthase.
Plant Physiology 117, 165–172.
Peterbauer, T., Mucha, J., Mayer, U., Popp, M., Gloessl, J. and Richter, A. (1999)
Stachyose synthesis in seeds of adzuki bean (Vigna angularis): molecular
cloning and functional expression of stachyose synthase. The Plant Journal 20,
509–518.
Peters, J.E., Padock, E.F., Tegenkamp, I. and Tegenkamp, T. (1977) Haploid callus
cells from anthers of Phaseolus vulgaris. Phytomorphology 27, 79–85.
Petruzelli, L. and Taranto, G. (1989) Wheat ageing: the contribution of embryonic
and non-embryonic lesions to loss seed viability. Plant Physiology 76, 289–294.
Peyronnet, C., Bastianelli, D. and Gatel, F. (1996) Factors of pulse value. Grain
Legumes 11, 20–21.
Phansiri, S., Miyake, H., Maeda, E. and Taniguchi, T. (1993) Ultrastructural analysis
of cell wall formation and cell division of soybean (Glycine max) protoplasts.
Japanese Journal of Crop Science 62, 429–437.
Pharr, D.M., Hendrix, D.L., Robbins, N.S., Gross, K.C. and Sox, H.N. (1987)
Isolation of galactinol from leaves of Cucumis sativus. Plant Science 50, 21–26.
Phillips, R.D. (1989) Composition and flatulence-producing potential of commonly
eaten Nigerian and American legumes. Food Chemistry 33, 271–281.
Pickardt, T., Saalbach, I., Waddell, D., Meixner, M., Muntz, K. and Schieder, O.
(1995) Seed specific expression of the 2S albumin gene from Brazil nut
(Bertholetia excelsa) in transgenic Vicia narbonensis. Molecular Breeding 1, 295–301.
Pigeaire, A., Abernethy, D., Smith, P.M.C., Simpson, K., Fletcher, N., Lu, C.Y.,
Atkins, C.A. and Cornish, E. (1997) Routine transformation of a grain legume
crop (Lupinus angustifolius L.) via Agrobacterim tumefaciens-mediated gene
transfer to shoot apices. Molecular Breeding 3, 341–349.
Piotrowicz-Cieslak, A.I., Górecki, R.J. and Frias, J. (1995) Changes in galactosides
content during maturation of yellow lupin (Lupinus luteus L.) and lentil (Lens
culinaris L.) seeds. In: AEP (ed.) Improving Production and Utilisation of Grain
Legumes. Proceedings 2nd European Conference on Grain Legumes. AEP, Paris, p. 378.
307
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:44 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
292 References
Pizzoferrato, L., Capelloni, M., Carnovale, E., Clementi, A., Ceccarelli, G. and
Tolomeo, M. (1995) Evaluation of starch digestibility in mottled beans and
lentils. In: AEP (ed.) Improving Production and Utilisation of Grain Legumes.
Proceedings 2nd European Conference on Grain Legumes. AEP, Paris, p. 278.
Plant, A.R. (1984) Studies of the protein bodies of Lupinus angustifolius. Dissertation
Abstracts International, C- European Abstracts 45, 367.
Pokorny, J. and Dostalova, J. (1996) Legumes – their consumption and nutritional
value. Vyz. Potraviny 51, 133–135 (in Czech).
Poncet, C., Remond, D., Bernard, L. and Peyronnet, C. (1998) Nitrogen value of
protein rich leguminous seed in ruminants: ruminal degradability of raw
and extruded pea and lupin. In: AEP(ed.) Improving Production and Utilisation
of Grain Legumes. Proceedings of 3rd European Conference on Grain Legumes. AEP,
Paris, pp. 16–17.
Polanco, M.C. and Ruiz, M.L. (1997) Effect of benzylaminopurine on in vitro and
in vivo in lentil (Lens culinaris Medik). Plant Cell Reports 17, 22–26.
Polisetty, R., Paul, J., Deveshwar, J.J., Khetarpal, S., Suresh, K. and Chandra, R.
(1997) Multiple shoot induction by benzyladenine and complete plant
regeneration from seed explants of chickpea (Cicer arietinum L.). Plant Cell
Reports 16, 565–571.
Ponsamuel, J. Huhman, D.V., Cassidy, B.G. and Post-Beittenmiller, D. (1998) In
vitro regeneration via caulogenesis and brassin-induced shoot conversion of
dormant buds from plumular explants of peanut (Arachis hypogeae L. cv.
Okrum). Plant Cell Reports 17, 373–378.
Potrykus, I. and Spangenberg, G. (1995) Gene Transfer to Plants. Springer-Verlag,
Berlin.
Power, J.B. and Chapman, J.V. (1985) Isolation, culture and genetic manipulation
of plant protoplasts. In: Dixon, R.A. (ed.) Plant Cell Culture – a Practical
Approach. IRL Press, Oxford, pp. 37–66.
Preiss, J. and Levi, C. (1980) Starch biosynthesis and degradation. In: Pridham J.B.
(ed.) The Biochemistry of Plants, Vol. 3. Academic Press, New York, pp. 371–423.
Price, K.R., Lewis, J., Wyatt, G.M. and Fenwick, G.R. (1988) Flatulence causes,
relation to diet remedies. Nahrung 32, 609–623.
Pridham, J.B. and Dey, P.M. (1974) The nature and function of higher
plant α-galactosidases. In: Pridham, J.B. (ed.) Plant Carbohydrate Biochemistry.
Academic Press, New York, pp. 83–96.
Proceedings of the 1st European Conference on Grain Legumes (1992). AEP, Paris.
Proceedings of the 3rd European Conference on Grain Legumes. Opportunities for High Qual-
ity, Healthy and Added-Value Crops to Meet European Demands (1998). AEP, Paris.
Proceedings of the International Conference Euro Food Tox IV on Bioactive Substance in Food
of Plant Origin (1994). Oloztyn, Poland. FECS event 193.
Prosky, L., Asp, N.G., Furda, I., De Vries, J., Schweizer, T.F. and Harald, B.F. (1985)
Determination of total dietary fibre in foods and food products: collaborative
study. Journal of the AOAC 68, 677–679.
Puonti-Kaerlas, J. and Eriksson, T. (1988) Improved protoplast culture and regener-
ation of shoots in pea (Pisum sativum L.). Plant Cell Reports 7, 242–245.
Puonti-Kaerlas, J., Eriksson, T. and Engstrom, P. (1990) Production of transgenic
pea (Pisum sativum L.) plants by Agrobacterim tumefaciens-mediated gene trans-
fer. Theoretical and Applied Genetics 80, 246–252.
308
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:44 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 293
309
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:45 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
294 References
Reddy, N.R., Pierson, M.D., Sathe, S.K. and Salunkhe, D.K. (1984) Chemical,
nutritional and physiological aspects of dry bean carbohydrates. A Review of
Food Chemistry 13, 25–68.
Rees ap, T. and Morrell, S. (1990) Carbohydrate metabolism in developing
potatoes. American Potato Journal 67, 835–847.
Reeves, J.B. (1988) Near infrared spectroscopic analysis of lignin components in
sodium chlorite-treated and untreated forages and forage by-products. Journal
of Dairy Science 71, 388–397.
Reichert, R.D. (1982) Air classification of peas (Pisum sativum) varying widely in
protein content. Journal of Food Science 47, 1263–1267.
Reichert, R.D. and Youngs, C.G. (1978) Nature of the residual protein associated
with starch fractions from air-classified field peas. Cereal Chemistry 55, 469–480.
Reid, D.S., Carr, J.M., Sajjaanantakul, T. and Labavich, J.M. (1986) Effect of freez-
ing and frozen storage on the characteristics of pectin extracted from cell
walls. ACS Symposium Series, American Chemical Society 310, 200–216.
Reid, J.S.G. (1985) Galactomannans. In: Dey, P.M. and Dixon, R.A. (eds) Bio-
chemistry of Storage Carbohydrates in Green Plants. Academic Press, London,
pp. 205–288.
Reisch, B., Duke, S.H. and Bingham, E.T. (1980) The genetic control of bud
formation from callus cultures of diploid alfalfa. Plant Science 20, 71–77.
Rentz, A. and Stitt, M. (1993) Substrate specificity and product inhibition of
different forms of fructokinases and hexokinases in developing potato tubers.
Planta 190, 166–175.
Revilla, M.A. and Tarango, J.F. (1986a) Scanning electron microscopy study on
starch granules in lentil cotyledons. Starch 38, 199–202.
Revilla, M.A. and Tarango, J.F., (1986b) Ultrastructure of starch grain breakdown in
cotyledone cells. Starch 38, 379–381.
Revilleza, M.J.R., Mendoza, E.M.T. and Raymundo, L.C. (1990) Oligosaccharides in
several Philippine indigenous food legumes: determination, localization and
removal. Plant Foods for Human Nutrition 40, 83–93.
Roberts, K. (1996) Structures at the plant cell surface. Current Opinion in Cell Biology
2, 920–928.
Robertson, J.B. and Horvath, P.J. (1993) Detergent analysis of foods. In: Spiller,
G.A. (ed.) Handbook of Dietary Fibre in Human Nutrition. CRC Press, Boca Raton,
Florida, pp. 49–52.
Rochat, C., Wuilleme, S., Boutin, J.-P. and Hedley, C.L. (1995a) ADPGPPase activity
affects storage product metabolism in rbrb pea seed coats. In: AEP (ed.) Proceed-
ings of the 2nd European Conference on Grain Legumes, Copenhagen. Improving
Production and Utilisation of Grain Legumes. AEP, Paris, pp. 34–35.
Rochat, C., Wuilleme, S., Boutin, J.-P. and Hedley, C.L. (1995b) A mutation at the rb
gene, lowering ADPGPPase activity, affects storage product metabolism of pea
seed coats. Journal of Experimental Botany 46, 415–421.
Rodriguez, A., Martin, L., Alonso, J.R., Nicolas, G. and Villalobos, N. (1988)
Scanning electron microscope study of starch granule degradation in chick-pea
cotyledons. Part I. Influence of embryonic axis on degradation process. Starch
40, 211–214.
Roffs, C.H., Schon, H., Steffen, M. and Kindl, H. (1987) Cell suspension culture of
Arachis hypogea L. Model system for specific enzyme induction in secondary
metabolism. Planta 172, 238–244.
310
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:46 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 295
Roper, W. (1979) Growth and cytology of callus and cell suspension cultures of Vicia
faba. Zeitschrift für Pflanzen Physiology 93, 245–257.
Ross, J.K., English, C. and Perlmutter, C.A. (1985) Dietary fiber constituents of
selected fruits and vegetables. Journal of the American Dietetic Association 85,
1111–1116.
Rossi, M., Germondari, I. and Casini, P. (1984) Comparison of chickpea cultivars.
Chemical composition, nutritional evaluation and oligosaccharide content.
Journal of Agricultural and Food Chemistry 2, 811–814.
Roth, E.J., Frazier, B.L., Apuya, N.R. and Lark, K.G. (1989) Genetic variation in an
inbred plant: variation in tissue cultures of soybean [Glycine max (L.) Merrill].
Genetics 121, 359.
Rubluo, A., Kartha, K.K., Mroginski, L.A. and Dyck, J. (1984) Plant regeneration
from pea leaflets cultured in vitro and genetic stability of regenerants. Journal of
Plant Physiology 117, 119–130.
Rufty, T.W., Israel, D.W. and Wolk, R.J. (1984) Assimilation of 15NO −3 taken up by
plants in the light and in the dark. Plant Physiology 76, 769–775.
Rufty, T.W., Huber, S.C. and Volk, R.J. (1988) Alteration in leaf carbohydrate
metabolism in response to nitrogen stress. Plant Physiology 88, 725–730.
Russell, D.R., Wallace, K., Bathe, J., Martinell, B. and McCabe, D. (1993) Stable
transformation of Phaseolus vulgaris via electric-discharge mediated particle
acceleration. Plant Cell Reports 12, 165–169.
Russel, P.L., Berry, C.S. and Greenwell, P. (1989) Characterisation of resistant
starch from wheat and maize. Journal of Cereal Science 9, 1–15.
Ryan, C.A. and Farmer, E.E. (1991) Oligosaccharide signals in plants: a current
assessment. Annual Reviews in Plant Physiology and Plant Molecular Biology 42,
651–674.
Rybczynski, J. and Podyma, E. (1993a) Preliminary studies of plant regeneration via
somatic embryogenesis induced on immature cotyledons of white lupin
(Lupinus albus L.) Genetica Polonica 34, 249–257.
Rybczynski, J. and Podyma, E. (1993b) Micropropagation of some Lupinus species
from seedling explants. Genetica Polonica 34, 237–247.
Saalbach, I., Pickardt, T., Machemehl, F., Saalbach, G., Schieder, O.N. and Muntz,
K. (1994) A chimaeric gene encoding the methionine-rich 2S albumin of the
Brazil nut (Bertholletia excelsa H.B.K.) is stably expressed and inherited in
transgenic grain legumes. Molecular General Genetics 242, 226–236.
Saalbach, I., Pickardt, T., Waddell, D.R., Hillmer, S., Schieder, O.N. and Muntz, K.
(1995a) The sulphur-rich Brazil nut 2S albumin is specifically formed in
transgenic seeds of the grain legume Vicia narbonensis. Euphytica 85, 181–192.
Saalbach, I., Waddell, D., Pickardt, T., Schieder, O. and Muntz, K. (1995b) Stable
expression of the sulphur-rich 2S albumin gene in transgenic Vicia narbonensis
increases the methionine content of seeds. Journal of Plant Physiology 145,
674–681.
Saalbach, L., Hofmeister, J. and Baumlein, H. (1997) Plant seeds as bioreactors:
use of a heat-stable, bacterial amylase in transgenic seeds of legumes for
liquefaction of seed starch. In: Proceedings 5th Symp. Renewable Resources. Schrift.
Bundesminister. Ernahrung, Landwirschaft und Forsten. Reihe A, Angew.
Wissenschaft., pp. 182–184.
Saeed, M. and Duke, S.H. (1990) Amylases in pea tissues with reduced chloroplast
density and/or function. Plant Physiology 94, 1813–1819.
311
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:46 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
296 References
Sagara, A.P., Suhasini, K. and Krishnamurthy, K.V. (1993) Plant regeneration via
somatic embryogenesis in chickpea (Cicer arietinum L). Plant Cell Reports 12,
652–655.
Saini, H.S. (1989) Legume seed oligosaccharides. In: Huisman, J., van der Poel,
A.F.B. and Liener, I.E. (eds) Recent Advances of Research in Antinutritional Factors
in Legume Seeds. Pudoc, Wageningen, pp. 329–341.
Saini, H.S. and Knights, E.J. (1984) Chemical constitution of starch and
oligosaccharide content of ‘desi’ and ‘kabuli’ chickpea (Cicer arietinum) seed
types. Journal of Agricultural and Food Chemistry 32, 940–944.
Saini, H.S. and Gladstones, J.S. (1986) Variability in the total and component
galactosyl sucrose oligosaccharides of Lupinus species. Australian Journal of
Agricultural Research 37, 157–166.
Sakai, K., Tachiki, T., Kumagai, H. and Tochikura, T. (1987) Hydrolysis of alpha-
D-galactosyl oligosaccharides in soymilk by alpha-D-galactosidase of Bifido-
bacterium breve 203. Agricultural Biological Chemistry 51, 315–322.
Sakurai, N. (1991) Cell wall functions in growth and development – a physical and
chemical point of view. The Botanical Magazine, Tokyo 104, 235–251.
Salunkhe, D.K. and Kadam, S.S. (eds) (1989) CRC Handbook of World Food Legumes:
Nutritional Chemistry, Processing Technology, and Utilization, Vol. 11. CRC Press,
Boca Raton, Florida.
Salunkhe, D.K., Sathe, S.K. and Deshpande, S.S. (1989) French Bean. In: Salunkhe,
D.K. and Kadam, S.S. (eds.) CRC Handbook of World Food Legumes: Nutritional
Chemistry, Processing Technology and Utilization, Vol. I. CRC Press, Boca Raton,
Florida, pp. 23–63.
Sambuceti, M.E. and Zuleta, A. (1996) Resistant starch in dietary fiber values mea-
sured by the AOAC method in different cereals. Cereal Chemistry 73, 759–761.
Sanago, M.H.M., Shattuck, V.I. and Strommer, J. (1996) Rapid plant regeneration
of pea using thidiazuron. Plant Cell Tissue and Organ Culture 45, 165–168.
Sandberg, A.-S., Andersson, H., Carlsson, N.-G. and Sandström, B. (1993) Degrada-
tion of soy bean oligosaccharides in the stomach and small intestine of human
ileostomy subjects. In: Schlemmer, U. (ed.) Proceedings of the International Confer-
ence on Bioavailability ‘93 – Nutritional, Chemical and Food Processing Implications
of Nutrient Availability. Bundesforschungsanstalt für Ernährung, Karlsruhe,
pp. 197–201.
Santangelo, E., Mosconi, P., Crino, P., Soressi, G.P. and Saccardo, F. (1997) In:
INTAGRES and ICARDA. New Perspectives for an Ancient Species. Aleppo, Syria,
pp. 73–83.
Saravitz, D.M., Pharr, D.M. and Carter, T.E. (1987) Galactinol synthase activity and
soluble sugars in developing seeds of four soybean genotypes. Plant Physiology
83, 185–189.
Sarko, A. and Wu, H.-C.H. (1978) The crystal structures of A-, B- and C-polymorphs
of amylose and starch. Starch/Stärke 30, 73–8.
Sasaki, K., Hicks, K.B. and Nagahashi, G. (1988) Separation of eight inositol isomers
by liquid chromatography under pressure using a calcium-form, cation-
exchange column. Carbohydrate Research 183, 1–9.
Sathe, S.K. and Salunkhe, D.K. (1981) Isolation and partial characterization of an
arabinogalactan from the Great Nortin Bean (Phaseolus vulgaris L.). Journal of
Food Science 46, 1276–1277.
312
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:47 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 297
Sator, C. (1990) Lupins (Lupinus spp.). In: Bajaj Y.P.S. (ed.) Legumes and Oilseed
Crops I, Biotechnology in Agriculture and Forestry, Vol. 10. Springer-Verlag,
Berlin-Heidelberg, pp. 288–311.
Saura-Calixto, F., Goni, I., Bravo, L. and Manas, E. (1993) Resistant starch in foods:
modified method for dietary fibre residues. Journal of Food Science 58, 642–643.
Savage, G.P. (1988) The composition and nutritive value of lentil (Lens culinaris).
Nutrition Abstract and Reviews (Series A) 5, 319–343.
Schaumann, A., Bruyant-Vannier, M.P., Goubet, F. and Morvan, C. (1993) Meta-
bolism in suspension cultured cells of Flax (Linum usitatissinum L.). Plant Cell
Physiology 34, 891–897.
Schlumbaum, A., Mauch, F., Vogeli, U. and Boller, T. (1986) Plant chitinases are
potent inhibitors of fungal growth. Nature 324, 365–367.
Schmitt, M.R., Hitz, W.D., Lin, W. and Giaquinta, R.T. (1984) Sugar transport into
protoplasts isolated from developing soybean cotyledons. II. Sucrose transport
kinetics, selectivity, and modelling studies. Plant Physiology 75, 941–946.
Schoch, T.J. and Maywald, E.C. (1968) Preparation and properties of various
legume starches. Cereal Chemistry 45, 564–571.
Scholda, R., Billek, G. and Hoffmann-Ostenhof, O. (1964) Biosynthesis of cyclitols.
VIII. Mechanism of conversion of myo-inositol to D-pinitol and D-chiro-inositol in
Trifolium incarnatum. Monatshefte für Chemie 95, 1311–1317.
Schroeder, H., Schotz, A., Wardley-Richardson, T., Spencer, D. and Higgins, T.
(1993) Transformation and regeneration of two cultivars of pea (Pisum sativum
L.). Plant Physiology 101, 751–757.
Schroeder, H.E., Gollasch, S., Moore, A., Tabe, L.M., Craig, S., Hardie, D.C.,
Chrispeels, M.J., Spencer, D. and Higgins, T.J.V. (1995) Bean (β-amylase
inhibitor confers resistance to the pea weevil (Bruchus pisorum) in transgenic
peas (Pisum sativum). Plant Physiology 107, 1233–1239.
Schrott, M. (1995) Selectable marker and reporter genes. In: Potrykus, I.
and Spangenberg, G. (eds) Gene Transfer to Plants. Springer-Verlag, Berlin,
pp. 325–336.
Schukla, T.P. (1996) Cereal grain and legume processing by extrusion. Cereal Foods
World 41, 35–36.
Schweizer, T.F., Horman, I. and Würsch, P. (1978) Low molecular weight carbohy-
drates from leguminous seeds; a new disaccharide: galactopinitol. Journal of the
Science of Food and Agriculture 29, 148–154.
Schweizer, T.F. and Horman, I. (1981) Purification and structure determination of
three α-D-galactopyranosylcyclitols from soya beans. Carbohydrate Research 95,
61–71.
Schweizer, T.F., Andersson, H., Langkilde, A.M., Reimann, S. and Torsdottir, I.
(1990) Nutrients excreted in ileostomy effluents after consumption of mixed
diets with beans or potatoes. II. Starch, dietary fibre and sugars. European
Journal of Clinical Nutrition 44, 567–575.
Scowcroft, W.R., Brettell, R.I.S., Ryan, S.A., Davies, P.A. and Pallotta, M.A. (1987)
Somaclonal variation and genomic flux. In: Green, C.E. (ed.) Plant Tissue and
Cell Culture. Alan R. Liss, New York, pp. 275–286.
Selvendran, R.R. (1983) The chemistry of plant cell walls. In: Birch, G.G.
and Parker, K.J. (eds) Dietary Fibre. Applied Science Publishers, London,
pp. 95–147.
313
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:48 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
298 References
Selvendran, R.R. (1984) The plant cell wall as a source of dietary fibre: chemistry
and structure. American Journal of Clinical Nutrition 39, 320–337.
Selvendran, R.R. and King, S.E. (1989) Structural features of the cell-wall polysac-
charides of the parchment layers of the pods of mature runner beans. Carbohy-
drate Research 195, 87–99.
Selvendran, R.R. and O’Neill, M.A. (1987) Isolation and analysis of cell walls from
plant material, In: Glick, D. (ed.) Methods of Biochemical Analysis. John Wiley &
Sons, New York, pp. 25–154.
Selvendran, R.R. and Robertson, J.A. (1990) The chemistry of dietary fibre – an
holistic view of the cell wall matrix. In: Southgate, D.A.T., Waldron, K.,
Johnson, I.T. and Fenwick, G.R. (eds) Dietary Fibre: Chemical and Biological
Aspects. The Royal Society of Chemistry, Cambridge, pp. 27–43.
Selvendran, R.R., March, J.F. and Ring, S.G. (1979) Determination of aldoses and
uronic acid content of vegetable fiber. Analytical Biochemistry 96, 282.
Selvendran, R.R., Stevens, B.J.H. and O’Neill, M.A. (1985) Developments in the
isolation and analysis of cell walls from edible plants. In: Brett, C.T. and
Hillman, J.R. (eds) Biochemistry of Plant Cell Walls. Cambridge University Press,
Cambridge, pp. 39–78.
Senaratna, T., McKersie, B.D. and Bowley, S.R. (1985) Antioxidant levels in germi-
nating soybean axes in relation to free radical and dehydration tolerance. Plant
Physiology 78, 168–171.
Seve, P., Kerros, C., Lebreton, Y., Quemener, B., Gaborit, T. and Bouchez, P. (1989)
Effect of the extraction of α-galactosides from toasted or raw soybean meal on
dietary nitrogen and fat utilization in the young pig. In: Huisman, J., van der
Poel, T.F.B. and Liner, I.E. (eds) Recent Advances of Research in Antinutritional
Factors in Legumes Seeds. Pudoc, Wageningen, pp. 276–280.
Shade, R.E., Schroeder, H.E., Pueyo, J.J., Tabe, L.M., Murdock, L.L., Higgins, T.J.V.
and Chrispeels, M.J. (1994) Transgenic pea seeds expressing the α-amylase
inhibitor of the common bean are resistant to bruchid beetles. Bio-Technology
(New York) 12, 793–796.
Shamina, Z.B. and Butenko, R.G. (1976) Optimization of nutrient medium for
tissue culture of Vicia faba. Fiziologiya Rastenii (Plant Physiology) 23, 1264–1268.
Sharma, V.K. and Kothari, S.L. (1994) Genotypic differences of multiple
shoot-forming capacity in soybean. Plant Tissue Culture 4, 17–24.
Shehata, A., El-shimi, N.M. and Mesallam, A.S. (1985) Pectic substances of faba
beans and their relation to texture of cooked beans. Journal of Food Processing
and Preservation 9, 65–74.
Shekib, L.A. (1994) In-vitro digestibility and microscopic appearance of germinated
legume starches and their effect on dietary protein utilization. Food Chemistry
50, 59–63.
Shekib, L.A.E., Zouil, M.M., Youssef, M.M. and Mohammed, M.S. (1985). Effect of
cooking on chemical composition of lentil, rice and their blend (kloshary).
Food Chemistry 18, 163–168.
Shetty, K., Asano, Y. and Oosawa, K. (1992) Stimulation of in vitro shoot organo-
genesis in Glycine max Merrill by alantoin and amides. Plant Science 81, 245–251.
Shewmaker, C.K., Boyer, C.D., Wiesenborn, D.P., Thompson, D.B., Boersig, M.R.,
Oakes, J.V. and Stalker, D.M. (1994) Expression of E. coli glycogen synthase in
the tuber of transgenic potatoes (Solanum tuberosum) results in a highly
branched starch. Plant Physiology 104, 1159–1166.
314
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:48 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 299
Shoemaker, R.C. and Hammond, E.G. (1988) Fatty acid composition of soybean
[Glycine max (L.) Merrill] somatic embryos. In Vitro Cellular and Developmental
Biology – Plant 24, 829–832.
Shoemaker, R.C., Amberger, L.A., Palmer, R.G., Oglesby, L. and Ranch, J.P. (1991)
The effect of 2,4-dichlorophenoxyacetic acid concentration on somatic
embryogenesis and heritable variation in soybean [Glycine max (L.) Merr.]. In
Vitro Cellular and Developmental Biology – Plant 27, 84–88.
Shukla, K.S. (1987) Quantitative determination of oligosaccharides in defatted
soybean products by high speed liquid chromatography. Fat Science Technology
89, 75–79.
Sidduraju, P., Bravo, L. and Saura-Calixto, F. (1998) Chemical composition and
carbohydrate digestibility of common and underexploited legumes. In: AEP
(ed.) Improving Production and Utilisation of Grain Legumes. Proceedings of 3rd
European Conference on Grain Legumes. AEP, Paris, pp. 380–381.
Sievert, D. and Pomeranz, Y. (1989) Enzyme-resistant starch. I. Characterisation and
evaluation by enzymatic, thermoanalytical and microscopic methods. Cereal
Chemistry 66, 342–347.
Sievert, D. and Pomeranz, Y. (1990) Enzyme-resistant starch. II. Differential
scanning calorimetry studies on heat-treated starches and enzyme-resistant
starch residues. Cereal Chemistry 67, 217–221.
Sievert, D., Czujachowska, Z. and Pomeranz, Y. (1991) Enzyme-resistant starch. III.
X-ray diffraction of autoclaved amylomaize. VII. Starch and enzyme starch
residues. Cereal Chemistry 68, 86–91.
Siljeström, M. and Björck, I. (1990) Digestible and undigestible carbohydrates in
autoclaved legumes, potato, and corn. Food Chemistry 38, 145–152.
Siljeström, M., Eliasson, A.C. and Björck, I. (1989) Characterisation of resistant
starch from autoclaved wheat starch. Starch/Stärke 41, 147–151.
Silvester, K.I., Englyst, H.N. and Cummings, J.H. (1995) Ileal recovery of starch
from whole diets containing resistant starch measured in vitro and fermenta-
tion of ileal effluent. American Journal of Clinical Nutrition 62, 403–411.
Simmonds, D.H. (1992) Plant cell wall removal: cause for microtubule instability
and division abnormalities in protoplast cultures. Physiologia Plantarum 85,
387–390.
Simonenko, Y.V., Gleba, Y.Y. and Kuchuk, N.V. (1999) Double transformation:
producing transgenic phosphinotricin-resistant plants of commercial pea
lines. Russian Journal of Plant Physiology 46, 804–807.
Simoni, A., Mugnai, M., Storti, E., Bittini, P., Schipani, C., Simeti, C., Angelini, P.
and Buiatti, M. (1995) Biochemical markers for early screening of tolerant
genotypes in the system Glycine max – Diaporthe phaseolorum var. Caulivora.
Journal of Genetics and Breeding 49, 169–178.
Sims, I.M., Munro, S.L., Currie, G., Craik, D. and Bacie, A. (1996) Structural charac-
terization of xyloglucan secreted by suspension-cultured cells of Nicotiana
plumbaginifolia. Carbohydrate Research 293, 147–172.
Singh, K.B. (1997) Chickpea. Field Crops Research 53, 161–170.
Sinnaeve, G., Jones, A., Merghem, R., Jay, M., Dardenne, P. and Biston, R. (1995)
Non destructive single seed analysis of peas by near infrared spectroscopy.
In: AEP (ed.) Proceedings of 2nd European Conference on Grain Legumes: Improving
Production and Utilisation of Grain Legumes. AEP, Paris, France, pp. 370–371.
315
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:49 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
300 References
Sivak, M.N. and Preiss, J. (1995) Starch synthesis in seeds. In: Kigel, J. and Galli, G.
(eds) Seed Development and Germination. Marcel Dekker, New York, pp. 139–168.
Skrabanja, V., Liljeberg, H.G.M., Hedley, C.L., Kreft, I. and Bjorck, I.M.E. (1999)
The influence of genotype and processing on the in vitro rate of starch hydroly-
sis and resistant starch formation in peas (Pisum sativum L.). Journal of Agricul-
tural and Food Chemistry 47, 2033–2039.
Smietana, Z., Szpendowski, J. and Soral-Smietana, M. (1996) Modification of potato
starch by extrusion. Acta Academica Agricultura, Tech. Olstyn Technologia
Alimentorum 29, 3–13.
Smiley, K., Hensley, D. and Gasdorf, H. (1976) Alpha-galactosidase production and
use in a hollow-fiber reactor. Applied Environmental Microbiology 31, 615–617.
Smith, A.M. (1988) Major differences in isoforms of starch-branching enzyme
between developing embryos of round- and wrinkled-seeded peas (Pisum
sativum L.). Planta 175, 270–279.
Smith, A.M. and Denyer, K. (1992) Tansley Review No. 39. Starch synthesis in
developing pea embryos. New Phytologist 122, 21–33.
Smith, A.M., Quinton-Tulloch, J. and Denyer, K. (1990) Characteristics of plastids
responsible for starch synthesis in developing pea embryos. Planta 180,
517–523.
Smith, M.K. and McComb, J.A. (1981) Effect of NaCl on the growth of whole plants
and their corresponding callus cultures. Australian Journal of Plant Physiology 8,
267–275.
Smith, P.T., Kuo, T.M. and Crawford, C.G. (1991) Purification and characterization
of galactinol synthase from mature zucchini squash leaves. Plant Physiology 96,
693–698.
Smythe, B.M. (1967) Sucrose crystal growth. Australian Journal of Chemistry 20,
1097–1114.
Snedecor, G.W. and Cochran, W.G. (1973) Statistical Methods. Iowa State University
Press, Ames, Iowa.
Snoad, B. (1974) A preliminary assessment of ‘leafless peas’. Euphytica 23, 257–265.
Somiari, R. and Balogh, E. (1995) Properties of an extracellular glycosidase of
Aspergillis niger suitable for removal of oligosaccharides from cowpea meal.
Enzymatic and Microbial Technology 17, 311–316.
Song, H.S., Lim, S.M. and Widholm, J.M. (1994) Selection and regeneration of
soybean resistant to the pathotoxic culture filtrates of Septoria glycines.
Phytopathology 84, 948–951.
Sonnewald, U. (1992) Expression of E. coli pyrophosphatase in transgenic plants
alters photoassimilate partitioning. Plant Journal 2, 571–581.
Sonnewald, U., Hajiraezaei, M.R., Kossman, J., Heyer, A., Trethewey, R.N. and
Willmitzer, L. (1997) Expression of a yeast invertase in the apoplast of potato
tubers increases tuber size. Nature Biotechnology 15, 794–797.
Soral-Smietana, M. (1995) Faba bean monostarch phosphates. In: AEP (ed.) Proceed-
ings of the 2nd AEP Conference. Improving Production and Utilisation of Grain
Legumes. AEP. Paris, pp. 364–365.
Soral-Smietana, M. and Dziuba, Z. (1995) Changes in fraction structure of
faba bean starch. In: AEP (ed.) Proceedings of the 2nd AEP conference. Improving
Production and Utilisation of Grain Legumes. Copenhagen. AEP, Paris, pp. 402–403.
Soral-Smietana, M., Fornal, J. and Wronkowska, M. (1998) Microstructure and func-
tional properties of wheat and potato resistant starch preparates. Proceedings
316
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:50 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 301
317
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:50 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
302 References
Stephens, P.A., Barwale-Sehr, U., Nickell, C.D. and Widholm, J.M. (1991) A cyto-
plasmatically inherited, wrinkled-leaf mutant in soybean. Journal of Heredity 82,
71–73.
Steup, M., Robeneck, H. and Melkonian, M. (1983) In-vitro degradation of starch
granules isolated from spinach chloroplasts. Planta 158, 428–436.
Stewart, C.N. Jr., Adang, M.J., All, J.N., Boerma, H.R., Cardineau, G., Tucker, D. and
Parrott, W.A. (1996) Genetic transformation, recovery, and characterization of
fertile soybean transgenic for a synthetic Bacillus thuringiensis crylAc gene. Plant
Physiology 112, 121–129.
Stikova, O., Sekavova, H., Mrhalkova, I. and Fronek, P. (1996) Food consumption
and prediction of future food demand (in Czech). Výzkumný Ústav Zemedelské
Ekonomiky, Research Reviews 34, 1–40.
Stikova, O., Sekavova, H., Mrhalkova, I. and Baudisova, H. (1997) Food consump-
tion and estimation of food demand (in Czech). Výzkumný Ústav Zemedelské
Ekonomiky, Research Reviews 46, 1–60.
Stinard, P.S., Robertson, D.S. and Schnable, P.S. (1993) Genetic isolation, cloning,
and analysis of a Mutator-dominant antimorph of the maize amylose extender1
locus. Plant Cell 5, 1555–1566.
Stitt, M. (1996) Plasmodesmata play an essential role in sucrose export from leaves:
a step towards an integration of metabolic biochemistry and cell biology. Plant
Cell 8, 565–571.
Stute, R. (1990a) Eigenschaften und Anwendungsmöglichkeiten von Erbsenstärken.
Teil 1. Starch/Stärke 42, 178–184.
Stute, R. (1990b) Eigenschaften und Anwendungsmöglichkeiten von Erbsenstärken.
Teil 2. Starch/Stärke 42, 207–212.
Sugimoto, H. and Buren, J.P. (1970) Removal of oligosaccharides from soy milk by
an enzyme from Aspergillus saitoi. Journal of Food Science 35, 655–660.
Suhasini, K., Sagara, A.P. and Krishnamurthy, K.V. (1994) Direct somatic embryo-
genesis from mature embryo axes in chickpea (Cicer arietinum L.). Plant Science
103, 189–194.
Sun, W.Q. (1997) Glassy state and seed stability: the WLF kinetics of seed viability
loss at T > Tg and the plasticization effect of water on storage stability. Annals of
Botany 79, 291–297.
Sun, W.Q. and Leopold, A.C. (1993) The glassy state and accelerated aging of
soybeans. Physiologia Plantarum 89, 767–774.
Sun, W.Q. and Leopold, A.C. (1995) The Maillard reaction and oxidative stress
during aging of soybean seeds. Physiologia Plantarum 94, 94–104.
Sun, W.Q., Leopold, A.C., Crowe, L.M. and Crowe, J.H. (1996) Stability of dry
liposomes in sugar glasses. Biophysical Journal 70, 1769–1776.
Sun, Z., Duke, S.H. and Henson, C.A. (1995) The role of pea chloroplast
α-glucosidase in transitory starch degradation. Plant Physiology 108, 211–217.
Suzuki, H., Ozawa, Y. and Tanabe, O. (1966) Studies on the decomposition of
raffinose by α-galactosidase of Actinomycetes. Agricultural Biological Chemistry 30,
1039–1046.
Švábová, L. and Griga, M. (1997) Utilization of some Fusarium filtrates in resistance
breeding programme of peas. Cereal Research Communications 25, 847–848.
Švábová, L. and Griga, M. (1998a) Fusarium spp. on peas – several in vitro techniques
used in resistance breeding. In: AEP (ed.) Opportunities for High Quality, Healthy
318
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:51 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 303
319
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:52 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
304 References
320
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:52 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 305
Tovar, J., Granfeldt, Y. and Björck, I. (1992b) Effect of processing on blood glucose
and insulin responses to starch in legumes. Journal of Agricultural Food Chemistry
40, 1846–1851.
Trethewey, R.N., Geigenberger, P., Riedel, K., Hajirezaei, M.R., Sonnewald, U.,
Stitt, M., Riesmeier, J.W. and Willmitzer, L. (1998) Combined expression of
glucokinase and invertase in potato tubers leads to a dramatic reduction in
starch accumulation and a stimulation of glycolysis. Plant Journal 15, 109–118.
Trick, H.N. and Finer, J.J. (1998) Sonication assisted Agrobacterium-mediated
transformation of soybean [Glycine max (L.) Merrill] embryogenic suspension
culture tissue. Plant Cell Reports 17, 482–488.
Tricoli, D.M., Hein, M.B. and Carnes, M.G. (1986) Culture of soybean mesophyll
protoplasts in alginate beads. Plant Cell Reports 5, 334–337.
Trijatmiko, K.R. and Harjosudarmo, J. (1996) Plant regeneration of soybean
through somatic embryogenesis. Jurnal Bioteknologi Pertanian 1, 53–59.
Troszynska, A., Honke, J., Waszczuk, K. and Kozlowska, H. (1995) Oligosaccharide
content in legume seeds and their changes during sterilization. In: AEP (ed.)
Improving Production and Utilisation of Grain Legumes. Proceedings of the 2nd
European Conference on Grain Legumes. AEP, Paris, p. 288.
Trowell, H. (1972) Ischemic heart disease and dietary fibre. American Journal of
Clinical Nutrition 25, 926–932.
Trowell, H. (1976) Definition of dietary fibre and hypotheses that it is a protective
factor in certain diseases. American Journal of Clinical Nutrition 29, 417–427.
Trugo, L.C., Almeida, D.C.F. and Gross, R. (1988) Oligosaccharide contents in the
seeds of cultivated lupins. Journal of the Science of Food and Agriculture 45, 21–24.
Trugo, L.C., Ramos, L.A., Trugo, N.M.F. and Souza, M.C.P. (1990) Oligosaccharide
composition and trypsin inhibitor activity of P. vulgaris and the effect of germi-
nation on the α-galactoside composition and fermentation in the human
colon. Food Chemistry 36, 53–61.
Tsukamoto, C., Kikuchi, A., Harada, K., Kitamura, K. and Okubo, K. (1993) Genetic
and chemical polymorphisms of saponins in soybean seed. Phytochemistry 34,
1351–1356.
Tuan, Y.-H. and Phillips, R.D. (1991) Effect of the hard-to-cook defect and process-
ing on protein and starch digestibility of cowpeas. Cereal Chemistry 68, 413–418.
Turova, A.P. and Gladkych, A.S. (1964) Experimental and clinical pharmacology of
saponins. Farmakologia and Toxikologia 27, 242–249.
Tyler, R.T. and Panchuk, B.D. (1982) Effect of seed moisture content on the air
classification of field peas and faba beans. Cereal Chemistry 59, 31–33.
Tyson, R.H. and ap Rees, T. (1988) Starch synthesis by isolated amyloplasts from
wheat endosperm. Planta 175, 33–38.
Uchiyama, T., Numata, M., Terada, S. and Hosino, T. (1993) Production and
composition of extracellular polysaccharide from cell suspension cultures of
Mentha. Plant Cell Tissue and Organ Culture 32, 153–159.
Ueno, Y., Hasegawa, A. and Tsachiya, T. (1973) Isolation of O-methyl-scyllo-inositol
from mung bean seeds. Carbohydrate Research 29, 520–521.
Upadhyay, J.K. and Garcia, V.V. (1988) Effect of soaking and cooking on reduction
of oligosaccharides of cowpea (Vigna unguiculata (L.) Walp.). Philippine Journal
of Food Science and Technology 12, 21–28.
Urbano, G., Lopez-Jurado, M., Hernandez, J., Fernandez, M., Moreu, M.C., Frias, J.,
Diaz-Pollan, C., Prodanov, M. and Vidal-Valverde, C. (1995) Nutritional
321
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:53 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
306 References
322
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:54 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 307
323
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:54 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
308 References
324
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:55 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 309
Wang, G., Robertson, J., Parpia, B., Chen, J. and Campbell, T.C. (1991) Dietary fibre
composition of selected foods in the People’s Republic China. Journal of Food
Composition and Analyses 4, 293–303.
Wang, H. and Holl, F.B. (1988) In vitro culture and the incidence of somaclonal
variation in regenerated plants of Trifolium pratense L. Plant Science 55, 159–167.
Wang, H., Cutler, A.J. and Fowke, L.C. (1989) High frequences of preprophase
bands in soybean protoplast cultures. Journal of Cell Science 92, 575–580.
Wang, H.L., Patrick, J.W. and Offler, C.E. (1995) Cellular pathway of photosynthate
transfer in the developing wheat grain. III. A structural analysis and physiologi-
cal studies of the pathway from the endosperm cavity to the starchy endosperm.
Plant, Cell and Environment 18, 389–407.
Wang, T.L. and Hedley, C.L. (1991) Seed development in peas: knowing your three
‘r’s’ (or four or five). Seed Science Research 1, 3–14.
Wang, T.L. and Hedley, C.L. (1993) Genetic and developmental analysis of the
seed. In: Casey, R. and Davies, D.R. (eds) Peas, Genetics, Molecular Biology and
Biotechnology. CAB International, Wallingford, pp. 83–120.
Wang, T.L., Cook, S.K., Francis, R.J., Ambrose, M.J. and Hedley, C.L. (1987a) An
analysis of seed development in Pisum sativum. VI. Abscisic acid production.
Journal of Experimental Botany 38, 1921–1932.
Wang, T.L., Smith, C.M., Cook, S.K., Ambrose, M.J. and Hedley, C.L. (1987b) An
analysis of seed development in Pisum sativum, III. The relationship between
the r locus, the water content and the osmotic potential of seed tissues in vivo
and in vitro. Annals of Botany 59, 73–80.
Wang, T.L., Hadavizideh, A., Harwood, A., Welham, T.J., Harwood, W.A., Faulks, R.
and Hedley, C.L. (1990) An analysis of seed development in Pisum sativum L.
XIII. The chemical induction of storage product mutants. Plant Breeding 105,
311–320.
Wang, T.L., Bogracheva, T.Y. and Hedley, C.L. (1998) Starch: as simple as A, B, C?
Journal of Experimental Biology 49, 481–502.
Wardlaw, I.F. (1990) The control of carbon partitioning in plants. Tansley review
No. 27. New Phytologist 116, 341–381.
Warkentin, T.D. and McHughen, A. (1993) Regeneration from lentil (Lens
culinaris) cotyledonary nodes and potential of this explant for transformation
by Agrobacterim tumifaciens. LENS-Newsletter (ICARDA), Lentil Experimental News
Service 20, 26–28.
Weber, H., Heim, U., Borisjuk, L. and Wobus, U. (1995a) Cell-type specific, coordi-
nate expression of two ADP-glucose pyrophosphorylase genes in relation to
starch biosynthesis during seed development of Vicia faba L. Planta 195,
352–361.
Weber, H., Borisjuk, L., Heim, U., Buchner, P. and Wobus, U. (1995b) Seed coat-
associated invertases of faba bean control both unloading and storage
functions, Cloning of cDNAs and cell type-specific expression. The Plant Cell 7,
1835–1846.
Weber, H., Borisjuk, L. and Wobus, U. (1997) Sugar import and metabolism during
seed development. Trends in Plant Science 2, 169–174.
Weber, H., Heim, U., Golombek, S., Borisjuk, L. and Wobus, U. (1998) Metabolic
control of legume seed development: altering carbohydrate metabolism in
transgenic Vicia narbonensis. In: AEP (ed.) Opportunities for High Quality, Healthy
325
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:56 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
310 References
and Added-Value Crops to Meet European Demands. Proceedings 3rd European Confer-
ence on Grain Legumes, Valladolid. AEP, Paris, pp. 374–375.
Weightman, R.M., Renard, C.M.G.C. and Thibault, J.F. (1994) Structure and
properties of the polysaccharides from pea hulls. Part 1: Chemical extraction
and fractionation of the polysaccharides. Carbohydrate Polymers 24, 139–148.
Wen, Q.B., Lorenz, K.J., Martin, D.J., Stewart, B.G. and Sampson, D.A. (1996)
Carbohydrate digestibility and resistant starch of steamed bread. Starch/Stärke
40, 180–185.
Wettlaufer, S.H. and Leopold, A.C. (1991) Relevance of Amadori and Maillard
products to seed deterioration. Plant Physiology 97, 165–169.
Wheeler, P.G., Menzies, I.S. and Creamer, B. (1978) Effect of hypersmolar stimuli
coeliac disease on the permeability of the human gastrointestinal tract. Clinical
Science and Molecular Medicine 54, 495–501.
Wiege, B., Bergthaller, W. and Lindhauer, M.G. (1995) Separation of starch,
protein, fibre, and water soluble components from wrinkle pea flour in a pilot
plant and processing of wrinkle pea starch by extrusion. Proceedings of 2nd
European Conference on Grain Legumes, Copenhagen. AEP, Paris, pp. 352–353.
Wiggins, H.S. (1984) Nutritional value of sugars and related compounds undigested
in the small intestine. Proceedings of the Nutrition Society 43, 69–75.
Wilkie, K.C.B. (1985) Chapter 1: New perspectives on non-cellulosic cell-wall
polysaccharides (hemicelluloses and pectic substances) of land plants. In:
Brett, C.T. and Hillman, J.R. (eds) Biochemistry of Plant Cell Walls. Cambridge
University Press, Cambridge, London, pp. 1–37.
Williams, R.J. and Leopold, A.C. (1989) The glassy state in corn embryos. Plant
Physiology 89, 977–981.
Williamson, F.A., Fowke, L.C., Constable, F.C. and Gamborg, O.L. (1976) Labelling
of concavaline-A sites on the plasma membrane of soybean protoplasts.
Protoplasma 89, 305–308.
Williamson, J.D. and Quatrano, R.S. (1988) ABA-regulation of two classes of
embryo-specific sequences in mature wheat embryos. Plant Physiology 86,
208–215.
Willison, J.H.M. (1985) Isolated protoplasts as laboratory tools in the study of cell
wall deposition. In: Fowke, L.C. and Constabel, F. (eds) Plant Protoplasts. CRC
Press, Boca Raton, Florida, pp. 77–89.
Willmot, D.B., Nickell, C.D., Widholm, J.M. and Gray, L.E. (1989) Evaluation of soy-
bean resistance to Phialophora gregata culture filtrate in tissue culture. Theoretical
and Applied Genetics 77, 227–232.
Wink, M. (1984) Evidence for an extracellular lytic compartment of plant cell
suspension cultures: the cell culture medium. Naturwissenseshaften 17, 635–637.
Wink, M. (1994) The cell culture medium – a functional extracellular compartment
of suspension-cultured cell. Plant Cell Tissue and Organ Culture 38, 307–319.
Woitke, H.P., Kayser, J.P. and Hiller, L.M. (1970a) Fortschritte in der Erfotschung
der Treiterpenesaponine. Pharmazie 25, 133–143.
Woitke, H.P., Kayser, J.P. and Hiller, L.M. (1970b) Fortschritte in der Erfotschung
der Treiterpenesaponine. Pharmazie 25, 213–241.
Wolever, T.M.S. (1995) Dietary fibre and lipid metabolism in humans. In: Cherbut,
C., Barry, J.L., Lairon, D. and Durand, M. (eds) Dietary Fibre: Mechanism of Action
in Human Physiology and Metabolism. John Libbey Eurotext, Paris, pp. 69–81.
326
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:56 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 311
Wolever, T.M.S. and Brand-Miller, J. (1995) Sugars and blood glucose control. Clini-
cal Nutrition 62 (Suppl.), 212–221.
Wolever, T.M.S., Cohen, Z., Thompson, L.U., Thorne, M.J., Jenkins, M.J.A.,
Prokipchuk, E.L. and Jenkins D.J.A. (1986) Ileal loss of available carbohydrate
in man: comparison of a breath hydrogen methods with direct measurement
using a human ileostomy model. American Journal of Gastroenterology 81,
115–122.
Wrather, J.A. and Freytag, A.H. (1991) Selection of atrazine tolerant soybean calli
and expression of that tolerance in regenerated plants. Plant Cell Reports 10,
44–47.
Wright, D.J., Bumstead, M.R., Coxon, D.T., Ellis, H.S., DuPont, S. and Chan,
H.W.-S. (1984) Air classification of pea flour – analytical studies. Journal of
Science of Food and Agriculture 35, 531–542.
Würsch, P., Del Vedovo, S. and Koellreutter, B. (1986) Cell-structure and starch
nature as key determinations of the digestion rate of starch in legume. American
Journal of Clinical Nutrition 43, 25–29.
Xu, X.L., Li, X.H., Liu, W.H., Li, J.L. and Wang, Y. (1996) Study of transferring the
soybean mosaic virus coat protein (SMV-CP) gene into soyabeans using the
Ri-plasmid. Soybean Science 15, 279–288.
Xu, Z.H., Davey, M.R. and Cocking, E.C. (1982) Callus formation from root proto-
plasts of Glycine max (soybean). Plant Science 24, 111–115.
Xu, Z. and Yang C. (1993) Influential factors of somatic embryogenesis of Vicia faba,
Pisum sativum and Phaseolus vulgaris. Southwest China Journal of Agricultural
Sciences 6, 42–47.
Yadav, S. and Khetarpaul, N. (1994) Indigenous legume fermentation: effect on
some antinutrients and in-vitro digestibility of starch and protein. Food Chemistry
50, 403–406.
Yaklich, R.W. (1985) Effect of ageing on soluble oligosaccharide content in soybean
seeds. Crop Science 25, 701–704.
Yamaguchi, F., Ota, Y. and Hatanaka, C. (1996a) Extraction and purification of
pectic polysaccharides from soybean okara and enzymatic analysis of their
structures. Carbohydrate Polymers 30, 265–273.
Yamaguchi, F., Kojima, H. and Muramelo, M. (1996b) Effects of hexameta-
phosphate on soybean pectic polysaccharide extraction. Bioscience, Biotechnology
and Biochemistry 60, 2028–2031.
Yamaguchi, M., Kainuma, K. and French, D. (1979) Electron-microscopic observa-
tions of maize starch. Journal of Ultrastructure Research 69, 249–261.
Yamane, Y. (1975) Chromosomal variation in calluses induced in Vicia faba and
Allium cepa. Japanese Journal of Genetics 50, 353–355.
Yamasaki, Y. and Konno, H. (1989) α-Glucosidases of suspension-cultured
sugar-beet cells. Phytochemistry 28, 2583–2585.
Yang, L.J., Barratt, D.H.P., Domoney, C., Hedley, C.L. and Wang, T.L. (1990) An
analysis of seed development in Pisum sativum. X. Expression of storage protein
genes in cultured embryos. Journal of Experimental Botany 41, 283–288.
Yasui, T. (1980) Identification of a new galactosyl cyclitol from seeds of Vigna
angularis Ohwi et Ohashi (Adzuki bean). Agricultural and Biological Chemistry 44,
2253–2255.
327
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:57 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
312 References
Yasui, T., Tateishi, Y. and Ohashi, H. (1985) Distribution of low molecular weight
carbohydrates in the subgenus Ceratotropis of the genus Vigna (Leguminosae).
Botanical Magazine Tokyo 98, 75–87.
Yasui, T., Endo, Y. and Ohashi, H. (1987) Infrageneric variation of the low
molecular weight carbohydrate composition of the seeds of the genus Vicia
(Leguminosae). Botanical Magazine Tokyo 100, 255–272.
Yazdi-Samadi, B., Rinne, R.W. and Seif, R.D. (1977) Components of developing
soybean seeds: oil, protein, sugars, starch, organic acids and amino acids.
Agronomy Journal 69, 481–486.
Yellowless, D. (1980) Purification and characterization of limit dextrinase from
Pisum sativum L. Carbohydrates Research 83, 109–118.
Yook, C., Sosulski, F. and Bhirud, P.R. (1994) Effects of cationization on functional
properties of pea and corn starches. Starch 46, 393–399.
Young, D.H. and Kauss, H. (1983) Release of calcium from suspension-cultured
Glycine max cells by chitosan, other polycations and polyamines in relation to
effects on membrane permiability. Plant Physiology 73, 698–702.
Young, D.H., Kohle, H. and Kauss, H. (1982) Effect of chitosan on membrane per-
meability of suspension-cultured Glycine max and Phaseolus vulgaris cells. Plant
Physiology 70, 1449–1454.
Yue, S.X., Fu, J.H., Li, L.C., Yuan, H.L., Zhai, W.X., Chen, H., Su, W.Q. and Wang,
Y.L. (1996) Atrazine-resistant gene introduced into Heilongjiang soyabean
varieties and its expression and heredity. Scientia Agricultura Sinica 29, 78–83.
Yuste, P., Longstaff, M.A., McNab, J.M. and McCorquodale, C. (1991) The digest-
ibility of semipurified starches from wheat, cassava, pea, bean and potato by
adult cockerels and young chicks. Animal Feed Science and Technology 35,
289–300.
Zagorska, N.A. (1995) Induced adrogenesis and somaclonal variation in economi-
cally important species. DSc thesis, Bulgarian Academy of Sciences, Sofia.
Zagorska, N.A., Shamina, Z.B. and Butenko, R.G. (1974) The relationship of
morphogenetic potency of tobacco tissue culture and its cytogenetic features.
Biologia Plantarum 16, 262–274.
Zalewski, K. and Lahuta, L.B. (1998) The metabolism of ageing seeds. Changes in
the raffinose family oligosaccharides during storage of field bean (Vicia faba
var. minor Harz) seeds. Acta Societatis Botanicorum Poloniae 67, 193–196.
Zamora, F.A. and Fields, M.L. (1979) Nutritive quality of fermented cowpeas (Vigna
sinensis) and chickpeas (Cicer arietinum). Journal of Food Science 44, 234–236.
Zamski, E. (1995) Transport and accumulation of carbohydrates in developing
seeds: The seed as a sink. In: Kigel, J. and Galli, G. (eds) Seed Development and
Germination. Marcel Dekker, New York, pp. 25–44.
Zdunczyk, Z., Juskiewicz, J., Frejnagel, S., Wroblewska, M., Krefft, B. and Gulewicz,
K. (1999) Influence of oligosaccharides from lupin seeds and fructooligo-
saccharides on utilisation of protein by rats and absorption of nutrients in the
small intestine. Proceedings of 9th International Lupin Conference, 20–24 June 1999.
Klink Müritz, Germany.
Zdunczyk, Z., Juskiewicz, J., Frejnagel, S. and Gulewicz, K. (1998) Influence of
alkaloid and oligosaccharides from white lupin seeds on utilization of diets
by rats and absorption of nutrients in the small intestine. Journal of Animal and
Feed Sciences 72, 143–154.
328
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:58 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
References 313
329
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:01:58 A3867: AMA: Hedley: First Proof:30-Oct-00 References
Color profile: Disabled
Composite Default screen
Index
Index
Index
315
331
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
14 November 2000 14:32:47 A3867: AMA: Hedley: First Proof:14-Nov-00 Index
Color profile: Disabled
Composite Default screen
316 Index
available carbohydrates 79, 84–85 carbohydrates 5, 12, 22, 26, 28, 30–31,
classification 79 34, 37–43, 45, 49, 53–54, 56,
nutritional properties 84–85 120
glycaemic index 84 accumulation 122
see also mono- and disaccharides, biosynthesis 125–127
starch chemistry 15
extraction procedure 34
GC determination 35
bean 8 physiological role 131–132,
adzuki 19–20, 32 134–138
broad 26, 122 unloading of 118
brown 26 cell wall 25–28, 47, 49–50, 52, 54
butter 30 preparation 50
common 3, 123 cell wall components 127, 202
dry 20 biosynthesis 159–160, 202–203
faba 3, 8, 17, 122, 130 oligosaccharides 203–204
field 30 cellulose 25, 50, 52, 55
garden 20 content in seeds 65
green 29 chemical analysis 31
haricot 29 chickpea 2, 8, 19–21, 30, 32, 122
jack 3 D-chiro-inositol 18–19, 23, 32, 133
kidney 21, 29–30 Cicer arietinum 2, 8, 20–21, 30, 32,
lima 29, 123 214
mung 2, 20, 29–30, 32, 123 ciceritol 20–22, 32, 126, 236
navy 29 content in seeds 63–64
pinto 29 chickpea 63–64
runner 29 lentil 63–64
D-bornesitol 19 common bean 3
breeding 209–232 consumption of grain legumes 7–11
contradiction of goals 210 cooking of legume starch 112–113
goals 209–211, 235 extrusion 113
pedigree 211 high pressure 112
physical selection 222 low pressure 112
selection methods 220–222 cowpea 3, 19–20, 122, 123
Brabender viscograph 100, 103 crude fibre 48
broad bean 26, 123 crystallinity 94, 96–97, 100–101
brown bean 26 biaxial crystalline polymers 94
butter bean 30 cyclitols 31–32, 127, 135–137, 143
occurrence of 32
Cajanus cajan 2, 20
calculation and statistical analysis 37, debranching enzymes 140
40, 43 delignification 54, 57
Canavalia ensiformis 3 desiccation 131
capillary zone electrophoresis (CZE) 42 injury 143
advantages 43 stress 124, 132
disadvantages 43 tolerance 131, 141
recommended method 43 developmental stages 120
332
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
14 November 2000 14:33:40 A3867: AMA: Hedley: First Proof:14-Nov-00 Index
Color profile: Disabled
Composite Default screen
Index 317
333
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:02:01 A3867: AMA: Hedley: First Proof:30-Oct-00 Index
Color profile: Disabled
Composite Default screen
318 Index
334
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:02:01 A3867: AMA: Hedley: First Proof:30-Oct-00 Index
Color profile: Disabled
Composite Default screen
Index 319
muco-inositol 18 mutant 17
mung bean 2, 20, 29–30, 32, 123 production 209
myo-inositol 18, 20–21, 31–32, 36, 125, peanut 30, 123
128 pectic acid 27
D-myo-inositol-1-phosphate 126 pectic substances 26–27, 55
occurrence of 29
pectin 16, 25–26, 28, 50, 52, 55
Nastar R 116 pectinic acid 27
Nastar R Instant 116 pentoses 16, 50
native starch 102, 104, 106, 110, 116 Phaseolus aureus 30
navy bean 29 Phaseolus coccineus 30
near-infrared (NI) spectroscopy 224 Phaseolus lunatus 30
neutral detergent fibre NDF 48 Phaseolus vulgaris 3, 6, 20, 26, 30, 214
non-food use 98 phenyl α-D-glucoside 31
non-starch polysaccharides 69, 76–78 photoassimilates, supply of 119
composition 69, 77 pigeon pea 2, 19–20, 122
content in seeds 69–70, 76 D-pinitol 19–20, 22, 31–32, 126
nutritional properties 76–78 pinto bean 29
digestibility of energy 78 Pisum sativum 3, 8, 20, 22, 26, 30,
eliminate of negative effect 214
76 plant genetic transformation, methods
energetic effect 76 181–195
gas production 78 Agrobacterium-mediated
ileal digestibility 76–77 transformation 183
intestinal transit 77 electroporation 184
nutrient absorption 77–78 microinjection 184
see also fibre, unavailable particle bombardment 183–184
carbohydrates plant regeneration 148–156
organogenesis 151–153
pea regeneration 149
oligomers 126 somatic embryogenesis 150–151
oligosaccharides 16, 25, 45, 71, 124 polymerization, degree of 103, 108
accumulation 121 polysaccharides 15–16, 22, 27, 45
degradation 140 cellulosic 25
α-galactosides 63, 68, 71–73, 80 non-cellulosic 25
rathinose 120, 123, 125, 132, 135, protopectin 27
137, 140, 143 protoplasts culture 156–158
RFO 125–126, 132–134, 136, 140
stachyose 120, 123, 125, 132, 135,
137, 140, 143 raffinose 16–19, 31–32, 36, 40–41,
verbascase 120, 123, 132, 137, 140 63–65, 68, 71
ononitol 126 antinutritional effect 71
D-ononitol 19–21, 32 content in seeds 63–64, 68
optical rotation 44–45 beans 63
chickpea 63–64
faba bean 63, 65, 68
pea 3, 8, 20, 26, 29–30, 122, 123 lentil 63–64, 68
breeding programmes 210, 211 lupin 68
hulls 29 pea 63, 68
335
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:02:02 A3867: AMA: Hedley: First Proof:30-Oct-00 Index
Color profile: Disabled
Composite Default screen
320 Index
336
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
14 November 2000 14:34:13 A3867: AMA: Hedley: First Proof:14-Nov-00 Index
Color profile: Disabled
Composite Default screen
Index 321
337
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
14 November 2000 14:34:44 A3867: AMA: Hedley: First Proof:14-Nov-00 Index
Color profile: Disabled
Composite Default screen
322 Index
338
Z:\Customer\CABI\A3867 - Hedley - Carbohydrates\A3933 - Hedley - Carbodhydrates #L.vp
30 October 2000 15:02:04 A3867: AMA: Hedley: First Proof:30-Oct-00 Index