J Sci Food Agric 1998, 77, 289È311

Starch Production and Industrial Use

Roger P Ellis,1* M Patricia Cochrane,2 M Finlay B Dale,1 Carol M Duþus,2
Andrew Lynn,3 Ian M Morrison,1 R Derek M Prentice,2 J Stuart Swanston1
and Sarah A Tiller1
1 Scottish Crop Research Institute, Invergowrie, Dundee, DD2 5DA, UK
2 Scottish Agricultural College, West Mains Road, Edinburgh, EH9 3JG, UK
3 Scottish Agricultural College, Auchincruive, Ayr, KA6 5HW, UK
(Received 19 July 1996 ; revised version received 30 June 1997 ; accepted 22 October 1997)

Abstract : This review of starch is concerned with its industrial uses, origins and
structure. The current demand for starch is met by a restricted range of crops,
the most important of which are potatoes, maize, wheat and tapioca. Improve-
ments in the properties of starches for industrial uses can be achieved through
chemical and physical modiÐcation of extracted starch and through the manipu-
lation of starch biosynthesis in the plant itself. We examine starch structure and
composition in relation to its use and exploitation by industry. The current
understanding of physiological and biochemical mechanisms inÑuencing starch
formation in higher plants is described. This information is set in the context of
the need to know the physical/chemical speciÐcation for each individual starch
and to understand the genetic control of these characteristics in order to identify
target genes for manipulation. ( 1998 SCI.

J Sci Food Agric 77, 289È311 (1998)

Key words : starch ; industrial uses ; structure ; composition ; biosynthesis ; physi-

cal properties ; genetic manipulation ; environment e†ects

INTRODUCTION as granules in the chloroplasts of green leaves and in the

Industrial, non-food, uses of starch are emphasised in
one of the deÐnitions of starch in the Shorter Oxford
Dictionary, “a constituent of Ñour used as a paste with
water to sti†en cloth, Ðnish textiles and size paperÏ. The
aim of this review is to assess recent research in the
biology, chemistry and industrial utilisation of starch
and to extend and complement earlier reviews such as
those of Morrison (1988, 1989, 1993, 1995), Lelievre and
Liu (1994), Albertsson and Karlsson (1995), Jane (1995),
Morell et al (1995), Nelson and Pan (1995) and Smith et
al (1995).
The two principal components of starch are polymers
of glucose : amylose, an essentially linear molecule, and
amylopectin, a highly branched molecule. Starch occurs
* To whom correspondence should be addressed.
Contract/grant sponsor : The Scottish Office Agriculture,
Environment and Fisheries Department.
289
( 1998 SCI. J Sci Food Agric 0022È5142/98/$17.50. Printed in Great Britain
amyloplasts of storage organs such as seeds and tubers.
There is rapid, diurnal turnover of starch in chloro-
plasts but, in contrast, amyloplast starch is deposited
over a period of days or even weeks, stored and then
re-mobilised during the germination of seeds and the
sprouting of tubers. This review concerns only storage
starch.
The world-wide market for industrial starches is
expanding (Batchelor et al 1996) and current demand is
met by a restricted range of crops, the most important
of which are potatoes, maize, wheat and tapioca. There
are considerable di†erences in the properties of the
starches from these species. These di†erences depend
not only on di†erences in the relative proportions of
amylose and amylopectin and the characteristics of
these molecules, but also on the di†erences in the non-
starch components of starch granules such as lipids,
proteins and phosphate groups. For example, waxy
maize starch is largely composed of amylopectin and
290 R P Ellis et al

TABLE 1 of genetic transformation. It is one objective of this

Industrial uses of starch review to indicate how starch structure relates to func-
tion in industrial use and hence to target the genetic
Industry Use of starch/modiÐed starch manipulations needed to produce new plant starches.
Adhesive Adhesive production
Agrochemical Mulches, pesticide delivery, seed
coatings STARCH PROPERTIES AND NON-FOOD USES
Cosmetics Face and talcum powders
Detergent Surfactants, builders, co-builders, Within the EU, the industrial starch market amounts to
bleaching agents and bleaching 2É4 ] 106 tonnes. The UK industrial use accounts for
activators. 2É2 ] 105 tonnes, which is 25% of the total starch used
Food Viscosity modiÐer, glazing agent in the UK. A recent report (Batchelor et al 1996) listed
Medical Plasma extender/replacers, transplant 10 diverse industrial markets and each with several dis-
organ preservation, absorbent sanitary tinct uses for starch.
products
The fact that native starches are diverse, and modiÐ-
Oil drilling Viscosity modiÐer
cations to starch are numerous (Tables 2 and 3), allows
Paper and board Binding, sizing, coating
Pharmaceuticals Diluent, binder, drug delivery di†erent starches to be used for di†erent applications.
Plastics Biodegradable Ðller Novel processing techniques and the current demand
PuriÐcation Flocculant for biodegradable and renewable resources means that
Textile Sizing, Ðnishing and printing, Ðre starch can command more versatile markets (Table 1).
resistance Additionally, once hydrolysed, starch is a feedstock
chemical for conversion into numerous Ðne chemicals.

contains little amylose or lipid. Potato starch granules
are much larger than those of cereal endosperms,
contain negligible lipid and have a relatively high phos-
phate content.
Starches for use in industry (Table 1) are extracted
from the raw plant material and yield a wide range of
products. While improvements in the properties of
starches for industrial use can be achieved through
chemical and physical modiÐcation, improvements may
also be achieved by the manipulation of starch biosyn-
thesis in the plant itself by breeding or through the use
Industrial uses in which the granular nature of starch is
signiÐcant
The physical dimensions of starch granules (see the
section entitled Starch in the Crop Plant, and Table 4),
their biodegradability and the amount and nature of
their non-carbohydrate components all a†ect industrial
uses. Attributes such as “textureÏ and the ability to act as
a lubricant are also signiÐcant. Granular starches have
been used as Ðllers in petrochemical-based materials to
produce “biodegradable plasticsÏ (Albertsson et al 1994).

TABLE 2
Properties of native starch granulesa

Granule properties Potato Maize W axy maize W heat T apioca

Diameter (lm)
Range 5È100 3È26 3È26 1È40 4È35
Mean 30 15 15 10 20
Lipid (%w/w) 0É05 0É60 0É15 0É80 0É10
Protein (%w/w) 0É06 0É35 0É25 0É40 0É10
Phosphorus (%w/w) 0É08 0É02 0É01 0É06 0É01
Amylopectin (%) 21 28 0 28 17
Degree of polymerisation
Amylose 3000 800 È 800 3000
Amylopectin ] 106 2 2 2 2 2
Pasting temp (¡C) 60È65 75È80 65È70 80È85 65È70
Peak viscosity 3000 600 800 300 1000
Swelling power at 95¡C 1153 24 64 21 71
Paste viscosity Very high Medium Medium-high Medium-low High
Paste and Ðlm clarity Very clear Opaque Fairly clear Cloudy Quite clear
Retrogradation rate Medium-low High Very low High Low

a Based on a table by Swinkels (1992).

Starch for industry 291

TABLE 3
ModiÐed starches and their usesa

ModiÐed starch T reatment Advantage over Examples of Examples of

(example) native starch food use industrial use
(non-food)

Pregelatinised Heat/moisture Cold-water soluble Pie Ðllings, “instant Oil drilling, mining
productsÏ, coatings
Acid-thinned Acid Low hot paste Gums, jellies Textiles, laundry
viscosity, high gel starch
viscosity
Oxidised Hypochlorite Increased clarity, Gravy, sauce Paper, textiles,
reduced set-back thickeners, jellies spray starch,
adhesives
Hydroxyalkyl ethers Propylene oxide Increased clarity Salad dressings, pie Paper and textile
increased stability Ðllings sizes
EsteriÐed Acetic anhydride Reduced set-back, “InstantÏ foods, Textiles, paper,
increased clarity, frozen foods packaging, Ðlm
forms Ðlms/Ðbres
Monophosphates Phosphoric acid Increased stability Frozen foods, infant Paper, textiles,
to freeze/thaw formulae metal reÐning
cycles
Cross-linked, eg Phosphorus Increased stability Wide range of Paper, metal
di-starch phosphate, oxychloride to heat, pH, thaw canned and frozen sequestrants
di-starch adipate adipic anhydride and freeze-thaw foods
cycles
Cationic starches Tertiary or Increased solubility Paper, mining.
quaternary amines and dispersability
in cold water, and
increased binding to
negatively charged
materials

a Based on Galliard (1987).

A major problem is that starches are naturally hydro-
philic, whereas plastics are generally hydrophobic
(Albertsson and Karlsson 1995). The interaction
between the components in these composites is there-
fore limited. Nevertheless, starch-containing plastic
Ðlms and bottles have been produced (Doane 1989 ;
Griffin 1994). When such plastics are exposed to micro-
organisms, the starch is broken down leaving the plastic
in a weakened state (Vallini et al 1994). In addition to
the loss of mechanical properties, there is an increase in
the permeability of the material, together with an
increase in the surface/volume ratio. These changes,
resulting from the biodegradation of starch granules,
facilitate further abiotic degradation processes
(Albertsson and Karlsson 1995). Whether the term “bio-
degradable plasticsÏ is appropriate for such composites
is a matter of debate.
The particle size of the starch (see the section entitled
Granule size and morphology) determines which
starches can be incorporated into composites. Particle
size is more critical in the production of thin Ðlms
(Lim et al 1992) than in the production of moulded
objects (Albertsson and Karlsson 1995) where up to
D50% (w/w) of the “plasticÏ can be granular starch
(Imam et al 1995). Although the diameter of the A-type
starch granules of wheat is greater than the thick-
ness of some plastic Ðlms, the lenticular shape of the
granules would allow them to orientate in the plane
of the Ðlm. However, since wheat starch granules are
often associated with signiÐcant amounts of protein and
this can result in Maillard-type reactions, which cause
discoloration, wheat starch has not been used in the
production of biodegradable plastic Ðlms (Griffin 1989).
Alternative sources of small-granule starch, such as
amaranth, could be used but these are still relatively
expensive. A method for the production of small-
particle starch from maize or other abundant starches
has been developed (Jane et al 1992). In addition to
incorporation into plastic Ðlms, the dimensions of small
starch granules make them suitable as a paper coating
(Wilhelm 1993). The smooth sensation they give when
rubbed on the skin has also found application in cos-
metic products such as face and “talcumÏ powders
(Mazza et al 1992). They are particularly used as
absorbents and in situations where allergic reactions are
produced by talc (magnesium silicate) powders.
 292
TABLE 4
The composition of starches from various sourcesa

Species V ariety/ Nitrogen (% w/w Protein Integral lipidb,c Amylose (%) Granule Granule
designation phosphorus) (%) (mg lipid per 100 g dry wt) shape diameter (lm)
(mg per 100 g starch) T otal Apparent

Barley Wild type LPL 587È1126M

A granules 0É23N 22É1N 10È25N
B granules 0É43N 19É0N 5N
Waxy LPL 158È460M
A granules 0É06N 3É6N 10È25N
B granules 0É15N 1É8N 1È5N
High amylose LPL 864È1360M
A granules 22É6L Disc-shapedB 15È32B
B granules 18É2L Lenticular 2È3
Steptoe Nitrogen 0É07H 0É40H 24É1H
Golf 28É0I
Glacier Nitrogen 0É08H 0É46H LPL 720, FFA 46M 27É7H, 30É0I
HA Nitrogen 0É11H 0É63H LPL 1137È1216M 34É8H, 37É4È47É9M
HA ] n 39É0I
HA ] N 40É0I
Compana LPL 888, FFA 36M 28É3M
wx Nitrogen 0É11H 0É63H LPL 457, FFA 28M 3É0H, 9É0I, 4É5M
wx ] n Nitrogen 0É16H 0É91H LPL 460, FFA 33M 1É9H, 5É3M
Maize Wild type Phosphorus 32K 0É73K LPL 162È344, FFA 297È529M 27É5G 21É4G Polyhedric 5È20B
0É2È0É4P 25É1K 20É9K
Waxy LPL 3È19, FFA 5È46M 7É0E, 0É4G 1É0E, 0G
High amylose LPL 262È602, FFA 375È671M
Dent 28É0E 28É0E Polygonal 5È18B
Amylomaize V 41É0E 47É0E Irregular 6È15B
Amylomaize-7 32P 66É5G 58É8G
Oh43 mutantsF
wild type 24F
wx 0F
ae 35F
Millet Cat-tail millet LPL 351È656, FFA 283È545M Polyhedric,
sphericalI 3È15B
Oat ChinensisA Nitrogen 0É05A LPL 977È1081, FFA 268È462M 19É4A 16É7A PolygonalB 2È15B
polyhedral to 6È10A

R P Ellis et al
irregularA
Potato Phosphorus 72K 0É14K 16G 25É9G 26É0G Smooth 15È75 long axis
0É05È0É2P 21É0K 21É0K ElipsoidalB 12È60 short axisB
Rice Wild type FFA 222È367M 21É9G 15É2G Irregular 3È8
19É5G 13É6G PolygonalB
 Starch for industry
Waxy FFA 15È60M
Rye Phosphorus 23È26K 0É20È0É47K 1É0K 20É3È25É8K 16É9È22É7K LenticularI 22È36
A granules Phosphorus 24 mgK 0É15 0É9 25É1K 22É0K
B granules Phosphorus 38 mgK 0É44 1É4 22É7K 17É5K
Triticale Phosphorus 45 mgK 0É23K 21É9K 18É3K Disc shapedB
A granules 22È36B
B granules 5B
Wheat Bread wheat Phosphorus 62 mgK 0É64K LPL 845M SphericalB
Durum wheat LPL 968M
426G 27É2G 20É4G
A granules 0É1È0É6P 929 (LPL 845)M 25
B granules \ 15D
Cappelle Desprez
A granules 30É4G 26É0G
B granules 28É4G 22É7G
Aotea 31É2G 25É7G
A granules 29É5G 22É9G
B granules
Saikai 173 1170C 27É6C
Kanto 107 1070C 26É0C
Bai Huo 1050C 28É4C
Wx-1 290C 1É6C
Wx-2 150C 2É0C
Wx-3 120C 1É2C

a Source : A, Hoover and Vasanthan (1992) ; B, Jane et al (1994) ; C, Yasui et al (1996) ; D, Shi and Seib (1989) ; E, Bradbury and Bello (1993) ; F, Inouchi et al (1991) ; G, Morrison
and Laignelet (1983) ; H, Lorenz (1995) ; I, Salomonsson and Sundberg (1994) ; J, Blanshard (1987) ; K, Radosta et al (1991) ; L, Stark and Yin (1986) ; M, Morrison (1988) ; N,
MacGregor and Morgan (1984).
b In barley and wheat the integral, or true, starch lipids are composed entirely of LPLs, while in oats, rice and maize, FFAs make up 30È60% of the integral lipids of the granule.
c Abbreviations : LPL, Lysophospholipids ; FFA, free fatty acids ; HA, high amylose, wx - waxy, n - naked ; N - covered.

293
294
Industrial applications in which the granular nature of
starch is not of primary signiÐcance
Many industrial processes and applications use starch
after some or all of the granule structure has been
destroyed. When this has occurred, the properties of,
and the relationships between, the components are of
increased importance. Starch amylose-complexing lipids
a†ect starch gelatinisation and swelling (Morrison
1995). Di†erences in the amount and type of lipid (see
the section entitled Starch lipids) may be one reason
why two starches with similar amylose : amylopectin
ratios can di†er in physical properties such as viscosity.
Gelatinisation is “. . . the collapse (disruption) of
molecular order within the starch granule manifested in
irreversible changes in properties such as granular
swelling, native crystallite melting, loss of birefringence
and starch solubilisationÏ (Atwell et al 1988). “ExcessÏ
water is essential for complete gelatinisation to occur.
The energy required for the disruption of the molecular
order di†ers between granules and so gelatinisation
occurs over a range of temperatures. Starch granules
may have a reduced gelatinisation temperature as a
consequence of prior removal of some molecular order
by, for example, mechanical damage (kinetic and, poss-
ibly, thermal energy input). On an industrial scale the
energy input necessary for starch gelatinisation is con-
siderable, and is a signiÐcant part of the processing
costs. This energy requirement may be increased if the
starch granule crystallites are allowed to anneal before
gelatinisation (Hoover and Vasanthan 1994).
The gelatinisation temperature is a qualitative index
of crystalline structure. A quantitative measure of crys-
tallite order can be determined by di†erential scanning
calorimetry (DSC), which measures the gelatinisation
enthalpy (*H) of the sample (Morrison 1995). The
thermal analysis of starch gelatinisation has recently
been reviewed (Lelievre and Liu 1994). Individual gran-
ules will gelatinise over a 1 or 2¡C range but there is
considerable variation between granules and so a starch
gelatinisation temperature range should always be
quoted. The gelatinisation temperature range is a char-
acteristic of the genotype of the plant in which the
starch is synthesised (Table 5) and is a†ected by the
environmental conditions, especially the temperature
during granule development (see the section entitled
E†ect of environment).
Gelatinisation is required for particular processes, eg
textile sizing and industrial starch hydrolysis, as it
a†ects the rheology and viscosity properties of the
system and makes the starch more accessible to enzymic
action. As the granules swell and components are
leached into solution, the properties of the system
change and there is a transition from a suspension of
granules to a paste. The source of starch, and any treat-
ment to it, can change pasting behaviour. Thus pastes
from di†erent starches vary in clarity (Table 2), homo-
R P Ellis et al
geneity, texture and fragility.
When undamaged granules are held in aqueous sus-
pension at 40È50¡C short-chain, low-molecular-weight
linear polyglucans are leached from the granules
(Prentice 1991). As leaching temperatures are increased,
higher-molecular-weight and more branched material is
leached from the granules. Some starches, eg oat
starches, are unusual in that both amylose and amylo-
pectin co-leach throughout the swelling and leaching
processes. Starch granules that have been mechanically
damaged during isolation or processing show a di†erent
pattern of leaching, with a predominance of amylopec-
tin leached into cold water. As the extraction tem-
perature of the water is raised, increasing amounts of
linear amylose-like components are released. The tem-
perature required for the release of linear material is
lower for damaged starch granules than for undamaged
granules (Lynn and Stark 1995).
Starch solutions are viscous. The ability of starch to
change the viscosity of other solutions and pastes is
well-known and is exploited in the food industry. This
property is also used in the oil exploration industry
where starch is used to adjust the viscosity of drilling
muds (Munck et al 1988). Starches from di†erent
sources di†er in their viscosity and pasting properties
(Table 2).
Obtaining a consistent viscosity of starch solutions is
a major aim in industrial processes that involve starch
pastes or “solutionsÏ being handled by mechanical
systems, eg paper, corrugating, textile and food
industries. In the corrugating paper industry, main-
tenance of a satisfactory viscosity is essential to allow
the desired levels of paper penetration, dewatering and
“slingingÏ (the splashing and throwing of starch adhe-
sive ; the degree of slinging a†ects the accuracy of the
placement of the starch adhesive) (McPherson 1991).
Most native starches do not maintain a stable viscosity
when their pastes are subjected to high shear rates and
heated for an extended period but chemically modiÐed
starches perform appropriately under these conditions.
Industrial applications of chemically modiÐed starch
It is possible to modify starches chemically in order to
change their properties (Table 3) and greatly extend the
range of starch applications (Table 1). The non-food
industries using starch expanded when the properties of
starch were improved through chemical modiÐcation.
Cationic starches are usually produced by reacting
starch with tertiary or quaternary ammonium com-
pounds. The chemical modiÐcation changes the proper-
ties of the original starch in a number of ways.
Solubility and dispersability in cold water are improved
and, in paper making, the starches have a stronger affin-
ity for the Ðbres (Formento et al 1994). The processing
makes modiÐed starches more expensive than native
 Starch for industry
TABLE 5
Swelling factors, gelatinisation temperatures and gelatinisation enthalpies of starches from various sources

Starch V ariety/designation Procedure used to determine Gelatinisation temperatureb (¡C) Gelatinisation Swelling factor
gelatinisation parameters enthalpy (70¡C)
T T T (*H, J g~1)
o p f
Barley Not speciÐed DSCc (Starch : Water 1 : 2) 57È65E 8É3È13É3E
DSC (Starch : Water 1 : 1) 59 64 74F 13É3F
Steptoe Polarising Microscope/KoÑer Hot Stage 56 63 65D 6É98D
Glacier Polarising Microscope/KoÑer Hot Stage 56 61 65D 5É48D, 8É4A
Glacier (HA) Polarising Microscope/KoÑer Hot Stage 59 65 69D 4É09D
Compana
wx Polarising Microscope/KoÑer Hot Stage 58 65 68D 9É23D
wx ] n Polarising Microscope/KoÑer Hot Stage 58 64 67D 10É85D
Maize Oh43 mutantsG 5É38B
wildtype DSC (Starch : Water 1 : 2) 61 66 72G 3É6G
wx DSC (Starch : Water 1 : 2) 62 69 81G 4É6G
ae DSC (Starch : Water 1 : 2) 65 83 94G
Millet Foxtail DSC (Starch : Water 1 : 2) 62È75E 5É2È13É5E
Proso DSC (Starch : Water 1 : 2) 66È80E 6É4È11É4E
Oat ChinensisA DSC (Starch : Water 1 : 3) 61 66 73C 10É4C 12É14C
Commercial DSC (Starch : Water 1 : 1) 54 59 70F 10É5F
Commercial DSC (Starch : Water 1 : 2) 51 58 75K
Potato Not speciÐed DSC (Starch : Water 1 : 2) 54 62 75I 15É8I 18É40B
Rice Not speciÐed DSC (Starch : Water 1 : 2) 61È78E 7É5È15É6E
Commercial DSC (Starch : Water 3 : 4) 60 80 91H 14É5H
Rye Not speciÐed DSC (Starch : Water 1 : 2) 76È78E 7É5È15É6E 7É69B
Triticale Not speciÐed 6É92B
Wheat Not speciÐed DSC (Starch : Water 1 : 1) 53 61 82F 2É9L, 13É7F 10É00B
Not speciÐed DSC (Starch : Water 1 : 2) 59È64E 7É4È14É1E
Not speciÐed DSC (Starch : Water 1 : 2) 45 57 82 pk 12É4I
Waxy Not statedJ 59È61 64È66 68È73J 12É5È12É6J

a Source : A, Morrison et al (1986) ; B, Radosta et al (1991) ; C, Hoover and Vasanthan (1992) ; D, Lorenz (1995) ; E, Fujita et al (1992) ; F, Shamekh et al
(1994) ; G, Inouchi et al (1991) ; H, Seow and Teo (1993) ; I, Svensson and Eliasson (1995) ; J, Yasui et al (1996) ; K, Mua and Jackson (1995).
b Gelatinisation temperatures : T , onset ; T , peak ; T , Ðnal.
o p f
c DSC, di†erential scanning calorimetry.

295
296
starches. However, in paper production, this cost
penalty is outweighed because, compared with native
starches, less cationic starch is required to achieve an
equal level of Ðbre coating. The use of cationic starch
also reduces the amount of starch in the factory effluent,
thereby reducing the cost of disposal (Laleg and Pikulik
1993). Cationic starches which contain aldehyde groups
increase the dry strength of paper and also increase the
strength of never-dried paper wet webs (loc cit).
Chemical modiÐcation of starches can be performed
using methods described in a large number of patents.
Usually, modiÐcation is carried out in an aqueous
medium, necessitating the use of high concentrations of
reagents and extensive post-reaction washing. ModiÐ-
cation and washing processes are costly and there is
interest in either performing chemical modiÐcations
using “dryÏ methods (Hellwig et al 1992) or “activatingÏ
the starch prior to modiÐcation (Vasanthan et al 1995).
However, “activationÏ treatments may not produce the
same substitution patterns as those achieved by the wet
methods (loc cit). Hamunen (1995) found that cationisa-
tion seemed to proceed faster in the inner parts of
potato starch granules than on their surface, but that
the distribution of the cationic groups was the same
whether the granules were cationised by the “wetÏ or by
the “dryÏ method.
Industrial uses of starch that exploit its biodegradability
The stability of a very wide range of products from the
petrochemical industry causes disposal problems
(Glover 1993). The use of biodegradable polymeric
materials is one of many possible approaches to this
problem (Ellis 1995). The production, properties and
applications of such materials have recently been
reviewed (Chiellini and Solaro 1996).
For some applications, biodegradable materials may
have advantages (Tatarka 1996) but, in most cases, bio-
degradable alternatives do not achieve the economic or
functional performance of their traditional counterparts
(Jacobsen and Fritz 1996). Additionally, a material that
biodegrades rapidly is not a panacea, since, if degrada-
tion is rapid, the anaerobic conditions produced will
destroy the natural balance that ensures an optimal
recycling of elements (Albertsson 1993).
Granular starches have been used as Ðllers in biode-
gradable polymers, in an attempt to make these new
polymers price-competitive with petrochemical plastics
(Jacobsen and Fritz 1996). Not all starch-containing
plastics use granular starches. A number of formula-
tions use gelatinised or “destructuredÏ starches
(Verhoogt et al 1995). In these formulations starch is
chemically bonded to thermoplastic side chains in order
to strengthen the interaction between the starch and the
plastic and thereby overcome some of the problems
associated with composite materials (Doane, 1992 ;
Shogren et al 1993 ; Damodar and Fanta 1995).
R P Ellis et al
Biodegradable plastic production is still in its infancy
when compared to petrochemical plastic production.
Many di†erent approaches are being pursued in order
to produce materials with the desired properties. Starch
can play, and is playing, an important role in these
developments. Starch-based materials have already cap-
tured about 25% of the loose-Ðll packaging market
(“packaging peanutsÏ) from the expanded polystyrene-
type materials in the US (Alexander 1996).
Versatile biodegradable materials have been produc-
ed by microbial fermentations that use starch as a
carbon source (Chiellini and Solaro 1996), and Koch et
al (1993) concluded that it is possible to produce a new
generation of detergents in which the surfactants,
builders and co-builders and bleaching activators could
all be derived from starch. They calculated that 50È60%
of the chemicals in current powder formulations and
65È75% in liquid formulations could be replaced by
starch-derived products.
Hydrolysis
The hydrolysis of starch is outside the scope of this
review although it is a signiÐcant use of starch. For a
review of the enzymic hydrolysis of starch, see Guzman-
Maldonado and Paredes-Lopez (1995). The hydrolysis
products of starch can be further processed by micro-
bial fermentations into a very wide range of products
that include, organic acids, alcohols, ketones, polyols,
amino acids, nucleotides, biopolymers, lipids, proteins,
vitamins, antibiotics, and hormones (Doane 1989).
INDUSTRIAL EXTRACTION OF STARCH
FROM CROP PLANTS
For details of the industrial extraction of starch the
reader is directed towards Galliard (1987), Guzman-
Maldonado and Paredes-Lopez (1995) and Batchelor et
al (1996).
How the process of extraction may alter starch
properties
Mechanical damage
The mechanical processes necessary for the industrial
isolation of starch granules cause damage to the granule
structure. As a result, the rate and extent of the swelling
of the granules in water and their susceptibility to
enzymic digestion are increased relative to those of
“undamagedÏ starch granules (Tester and Morrison
1994a). The type and amount of carbohydrate that can
be leached or extracted from starch granules is also
dependent on whether they are damaged (Tester and
Morrison 1994b). Changes associated with damage to
the starch granules, eg swelling and enzymic suscepti-
bility, may alter the suitability of the starch for indus-
trial use. Damage levels can be measured using many
Starch for industry
methods including iodometric titration, enzymic
digestion and dye adsorption (Morgan and Williams
1995).
Drying
Drying is achieved by hot air dryers. High temperatures
give a greater throughput but, if the temperature is too
high, there is a reduction in the quality of the starch.
Economics dictate that a balance must be struck
between starch throughput, capital equipment costs and
starch quality. This balance may be more difficult to
achieve in the potato starch industry as a result of its
seasonal nature.
Process water
The viscosity of potato starch pastes and solutions is
a†ected by cations present in the process water
(Batchelor et al 1996) because the cations interact with
the phosphate groups in potato starch. Process water
can be conditioned to regulate its ion content. The
process water can also cause discoloration of isolated
starch if it contains an excessive amount of iron. Dis-
coloration is avoided by using chalk to precipitate Fe3`
ions.
STARCH IN THE CROP PLANT
Granule size and morphology
Starch granules range in size (Table 4) from those of
canna, with a maximum long axis of 100 lm, to those of
amaranth, with a long axis of 0É5È2É0 lm (Jane et al
1994). Granule surface morphology ranges from smooth
in most tuber starches, to indented in many legume
starches, sharp-edged in wild-type and waxy rice and
maize, or pock-marked in wheat, barley and triticale.
Starch granule shape can be characteristic of a genus
and species. A survey of starch granules from the genus
Solanum (Schittenhelm and Menge-Hartmann 1992)
showed that most granules were oval/round in shape,
but some had a length : breadth ratio [2 and others
had a very irregular shape. The mean length of starch
granules in 84 accessions from 57 species of Solanum,
determined by scanning electron microscopy, ranged
from 19É3 to 37É9 lm. For the same set of tubers, the
mean granule diameters, estimated by Coulter counter,
ranged from 18É2 to 35É2 lm. The large A-type starch
granules of wheat, barley and rye are lenticular, but
the smaller B-type starch granules are spherical or
polyhedral. The starch granules of rice, oats and
maize are irregular and polyhedral in shape, those of
rice being comparable in size to the B-type starch
granules of wheat and barley, while those of maize are
larger (Table 4).
Katz et al (1993) surveyed the granule morphology of
starches extracted from genotypes of maize which con-
tained mutant genes such as waxy (wx), dull (du),
297
amylose-extender (ae), Ñoury ( Ñ) and horny (h), and found
that each genotype had a distinct granule morphology.
In rice, the starch granules of high amylose mutants are
irregular and rounded, while those of the wild-type are
polyhedral (Yano et al 1985). In contrast, the starch of a
high amylose mutant in barley, though lacking the
largest A-type granules, does not di†er in granule mor-
phology from the starch of the wild-type (McDonald et
al 1991).
Granule size distribution
Microscopy can give information about granule shape
as well as granule size, but when a weight count is
made, statistical considerations require that granules
are measured in a large number of Ðelds of view. Image
analysis techniques have made this feasible, but the
most difficult problem remainsÈthat of preparing a
microscope slide containing a representative sample of
the starch granules (Allen 1990). Microscopy measures
surface area so, when these results are used to obtain
volume or mass distributions, errors are multiplied.
However, the method does allow granule shape to be
considered. In their analysis of bimodal populations of
barley starch granules using image analysis, Oliveira et
al (1994) calculated the volumes of the lenticular A-type
starch granules separately from those of the spherical
B-type granules.
Two methods of determining granule size distribution
are based on the response of an instrument to particle
volume. In the Coulter counter, the electrical sensing
zone method is used to determine the number and size
of a known weight of particles suspended in a known
volume of an electrolyte as the particles pass through an
oriÐce. Low-angle laser light scattering instruments
(LALLS) determine the size distribution of particles sus-
pended in a bath of liquid. In both instruments the size
is initially recorded as a volume and values for particle
diameters are derived using the assumption that the
particle is a sphere.
LALLS instruments (Cornell et al 1994 ; Blumenthal
et al 1996 ; Haase and Plate 1996) are easy to operate
and yield reproducible data (Allen 1990). However, the
Coulter counter determines both particle number and
particle size distribution, and has a much greater
number of class sizes than LALLS instruments. It has
been widely used (Chojecki et al 1986 ; McDonald et al
1991 ; Tester et al 1991 ; Shi et al 1994) and may well
continue to be the instrument of choice in studies of the
physiology of starch granule production in cereal
grains.
Sedimentation Ðeld Ñow fractionation (SFFF) has
been used for the size analysis of starch granules (Moon
and Giddings 1993). This system is rapid, gives high
resolution and has the added advantage that size frac-
tions can be collected.
298
Granule development
The size distribution of the starch granules in the
storage organ of a plant changes during its development
(Chojecki et al 1986) and the composition of starch
granules and their properties change during granule
development (Shannon and Garwood 1984). Morrison
and Gadan (1987) found that the proportions of
amylose and of lysophospholipids rose in both A-type
and B-type starch granules during grain development in
wheat. McDonald et al (1991), in a study of the com-
position of starch granules during grain development in
barley, observed similar changes in the proportions of
amylose and lysophospholipids.
Morrison (1993) suggested that the degree to which
amylose is complexed with lysophospholipids is a func-
tion of the stage of development of the endosperm, at
which the amylose is formed. B-type granules are initi-
ated later in grain development than A-type granules.
Thus mature A-type starch granules di†er in composi-
tion from mature B-type starch granules because the
Ðrst formed parts of the A-type starch granules contain
more lipid-free amylose than the corresponding parts of
the B-type starch granules. According to this hypothe-
sis, there is no need to envisage di†erent mechanisms
for the control of starch synthesis in the two types of
granule. However, there is still much to be understood
about the synthesis of starch in either type of granule.
The gelatinisation temperature of starch from mature
grains is lower than that of starch extracted from endo-
sperms of wheat, rye and barley at an early stage in
their development (Karlsson et al 1983). Viscosity
properties of starch change during grain development in
wheat (Kulp 1973) and during tuber development in
potato (Madsen and Christensen 1996).
E†ect of environment
Historical observations, summarised by Shannon and
Garwood (1984), indicate that growing conditions a†ect
the polysaccharide composition of starch, but that the
e†ects observed are not as large as those associated with
genotype or organ maturity. A comprehensive review of
the e†ect of the ambient temperature during the growth
of plants on the physicochemical properties of their
starches was published by Hizukuri (1969). He present-
ed evidence that the pasting temperature, a-amylase sus-
ceptibility and, in potato starch, the granule shape and
phosphorus content, all showed considerable depen-
dence on the temperature of the environment of the
mother organ. He also reported that temperature
during tuber growth had no e†ect on the iodine affinity
of potato starch. In contrast, Cottrell et al (1995) found
that starch from potatoes grown at a higher tem-
perature had a higher amylose content, higher gelati-
nisation temperature and greater resistance to bacterial
R P Ellis et al
a-amylase, than starch from tubers grown at lower tem-
peratures. Analyses of starch extracted from mature
tubers of a range of genotypes grown on several sites in
two successive years, indicate that environmental condi-
tions a†ect granule size distribution, phosphorus
content and viscosity properties but have little e†ect on
amylose content (Haase and Plate 1996).
Asaoka et al (1984) showed that an e†ect on the
amylose content of rice starch could be induced by a
relatively small di†erence in temperature (25¡C day/
19¡C night to 30¡C day/24¡C night), provided it was
experienced by the plants at 5È15 days after heading.
The starch synthesised in the higher temperature regime
had a lower amylose content and had amylopectin with
a higher amount of long B chains and a lower amount
of short B chains, than starch synthesised in the lower
temperature regime. Rice plants which experienced high
temperatures early in grain development produced
starch that had a higher gelatinisation temperature than
the starch from plants grown at lower temperatures.
Tester et al (1991) also used controlled environments in
their investigation of the e†ects of ambient temperature
during grain-Ðlling on the composition and properties
of the starch from four barley genotypes. They found
that the gelatinisation temperatures of the starches pro-
duced at higher temperatures were higher than those of
the starches formed in plants grown at lower tem-
peratures. They did not Ðnd di†erences in amylose
content, or in the Ðne structure of the amylopectin, but
did Ðnd temperature-related di†erences in the lipid
content of the starches. In a study of the variability of
starch properties in two cultivars of wheat in relation to
weather during grain-Ðlling, Morrison (1989) demon-
strated a signiÐcant positive correlation between lyso-
phospholipid content and accumulated temperature,
solar radiation and sunshine hours. Shi et al (1994)
investigated the e†ect of temperature in controlled
environments during grain-Ðlling on starch from six cul-
tivars of wheat. They reported similar e†ects of tem-
perature on the lipid content of starch and on
gelatinisation temperature and starch swelling. They
found that the proportion of shorter length B chains of
amylopectin was higher in starch from plants grown at
40¡C than in starch from plants grown at 25¡C. In a
recent investigation on the e†ects of elevated growth
temperatures and carbon dioxide levels on physico-
chemical properties of wheat starch using plants
grown in a double-skinned polythene tunnel, Tester et al
(1995) found that starch gelatinisation temperature
increased with plant growth temperature, whereas gela-
tinisation enthalpy was not a†ected by plant growth
temperature. Carbon dioxide concentration had no con-
sistent e†ects on any of the parameters measured. In
similar experiments, Blumenthal et al (1996) also failed
to detect any e†ect of elevated levels of carbon dioxide
during grain growth on the amylose/amylopectin ratio
in wheat starch.
Starch for industry
The starch from potatoes grown at higher tem-
peratures had fewer large granules than starch from
potatoes grown at lower temperatures (Cottrell et al
1995). Wheat plants which were subjected to high tem-
peratures during grain-Ðlling produced starch with
fewer B-type starch granules than that produced by
plants grown at lower temperatures (Bhullar and Jenner
1985). High temperature has been shown to cause a
reduction in the size of both A-type and B-type starch
granules in both wheat and barley (Tester et al 1991 ;
Shi et al 1994). Recent evidence indicates that, on a
volume basis, the proportion of A-type granules in
starch extracted from wheat grains grown at an elevated
concentration of carbon dioxide is greater than that in
starch extracted from wheat grains grown in ambient
levels of carbon dioxide (Blumenthal et al 1996).
The extent of genotype with environment interaction
in relation to starch properties has not been fully
explored in any crop species. The degree to which this
natural variation in the properties of starches a†ects
their suitability for industrial processing is still
unknown.
Enzymology of starch biosynthesis
The source of carbon for starch synthesis in storage
organs such as cereal endosperms and potato tubers is
generally believed to be sucrose (Du†us 1993). Sucrose
is converted to starch by the series of enzyme-catalysed
reactions shown in Fig 1. The Ðnal stages, which take
place within the amyloplast, are thought to be catalysed
by ADP-glucose pyrophosphorylase (EC 2.7.7.27),
starch synthase (EC 2.4.1.21) and starch branching
enzyme (EC 2.4.1.18). The factors regulating the amount
and structure of starch in storage organs have been
recently reviewed by Smith et al (1995).
ADP-glucose pyrophosphorylase is often referred to
as the Ðrst committed enzyme of starch biosynthesis.
ADP-glucose pyrophosphorylase has been identiÐed as
a key regulatory enzyme in photosynthetic tissues, being
strongly inhibited by inorganic phosphate and activated
by 3-phosphoglycerate. Whether or not this is the case
in starch storage tissue is less clear (Kleczkowski et al
1993) and other factors such as changes in speciÐc
signal compounds, turgor or compartmentation may be
involved (Geigenberger et al 1994).
To test the possible role of ADP-glucose pyrophos-
phorylase in starch synthesis during development in
potato tubers, MuŽller-RoŽber et al (1992) generated
transgenic plants in which the expression of ADP-
glucose pyrophosphorylase was inhibited. These were
obtained by introducing a chimeric gene containing the
coding region of one of the subunits of the ADP-glucose
pyrophosphorylase linked in the antisense orientation
to the CaMV 35S promoter. ADP-glucose pyrophos-
phorylase activity was almost completely inhibited in
299
Fig 1. Hypothetical scheme for the conversion of sucrose to
starch in developing amyloplasts. (1) Sucrose synthase ; (2)
invertase ; (3) hexokinase ; (4) phosphohexoisomerase ; (5) phos-
phoglucomutase ; (6) ADP-glucose and UDP-glucose pyrop-
hosphorylase ; (7) starch synthase ; (8) branching enzyme ; (9)
pyrophosphate/adenine nucleotide exchanger ; (10) nucleoside
diphosphate kinase. Abbreviations : Glc-1-P, glucose-1-phos-
phate ; Fru-6-P, fructose-6-phosphate ; PPi, pyrophosphate ;
Pi, inorganic phosphate. ADP-glucose transporter not shown.
tubers and the starch content of the tubers from trans-
formed plants was reduced to amounts between 4% and
35% of the levels in control plants. The dramatic
reduction in starch content observed as a direct result of
ADP-glucose pyrophosphorylase inhibition was taken
as further support for the hypothesis that ADP-glucose
is the major precursor for starch synthesis in higher
plants. In contrast, by expressing regulatory variants of
ADP-glucose pyrophosphorylase in potato tubers, the
amount of tuber starch may be increased (Stark et al
1992). In this case, the Escherichia coli ADP-glucose
pyrophosphorylase (fused to the plastid targeting
sequence) was fused to the tuber-speciÐc patatin pro-
moter. On average, the transgenic lines produced 35%
more starch than the control lines. However, it is
important to point out that, since the E coli and plant
enzymes have di†erent allosteric properties, it is not
possible to draw any conclusions from these data about
the role of allosteric regulation of the higher plant
enzyme in starch synthesis (MuŽller-RoŽber and
Komann 1994).
Recent work suggests that, in developing maize
(Denyer et al 1996) and barley (ThorbjÔrnsen et al 1996)
endosperm, both plastidial and cytosolic forms of ADP-
glucose pyrophosphorylase contribute to the synthesis
of ADP-glucose for starch synthesis. If this is the case,
then a proportion of the ADP-glucose formed from
300
sucrose (Fig 1) would be synthesised in the cytosol. This
may then be subsequently translocated to the amylo-
plast where it could be used directly for starch synthesis.
For this hypothesis to be tenable, a transporter would
be required to transfer ADP-glucose into the developing
amyloplast. Some evidence for such a transporter in the
plastidial membranes of maize endosperm has been put
forward (Cao et al 1995). Nevertheless we are still very
far from a full understanding of the intermediates,
enzymes, kinetics and cellular organisation of the con-
version of sucrose to ADP-glucose within the
developing endosperm cell.
Starch synthase catalyses the step-wise addition of
glucose residues to the non-reducing termini of malto-
dextrin primers with the formation of linear chains of
a-(1-4)-linked D-glucose residues. In developing barley
and wheat endosperms, activity has been observed with
both UDP-glucose and ADP-glucose (Baxter and
Du†us 1973 ; Caley et al 1990), but the greater part of
starch synthesis is catalysed by ADP-glucose-dependent
starch synthase. The starch synthases can be divided
into two classes, the granule-bound (GBSS I and II) and
the soluble starch synthases (SSS I and II). Within
classes, a number of isoforms have been reported in
several plant species. These include the primer-
dependent and the primer-independent starch synthases.
The enzyme responsible for the synthesis of amylose
is thought to be ADP-glucose-dependent GBSS (Preiss
et al 1991). ConÐrmation has been obtained in studies
of amylose synthesis in Agrobacterium rhizogenes-
transformed potato plants where expression of the
GBSS gene is inhibited via antisense RNA (Kuipers et
al 1994). In every case there was a reduction in the
amount of amylose synthesized. Similar observations
have been made by Kull et al (1995). In developing
cereal endosperms, the involvement of GBSS in amylose
synthesis is conÐrmed by the existence of waxy mutants
where both the amylose content and the GBSS activity
are low (Preiss et al 1991).
Starch branching enzyme (SBE) catalyses the forma-
tion of amylopectin by the introduction of a-(1-6)-link-
ages. The potato branching enzyme (Q-enzyme) has
been puriÐed and partially characterised (Blennow and
Johansson 1991). One form of the enzyme was found to
have a molecular mass of 103 kDa and it was suggested
that the apparent polymorphism might be due to
protein hydrolysis during extraction. Further difficulties
in the study of SBE arise from the need for an assay
procedure which allows for low levels of amylase con-
tamination, combined with sufficient sensitivity and
speciÐcity. A novel assay procedure has been developed
by Takeda et al (1993b) which uses NaBH -reduced
4
starch as a substrate. Following debranching with iso-
amylase, the number of introduced branch points is
determined by a reducing sugar assay. This method
improves both sensitivity and speciÐcity but is still sen-
sitive to the presence of amylase activity.
R P Ellis et al
Boyer and Preiss (1981) have explored the relation-
ship between branching enzymes and starch synthases
using mutant endosperms of maize. For example, in
amylose-extender (ae) mutants, which have starches with
an amylose content of around 66% compared to 26% in
the wild-type, there is an 80% decrease in one of the
isoforms of SBE. A starch synthase preparation, free
from branching enzyme, was isolated from the mutant.
When branching enzyme from wild-type maize was
added back to this fraction, starch synthesis was found
to increase 3É5È4 times. This indicates that the forma-
tion of amylopectin may occur by the combined and
simultaneous action of both starch synthase and
branching enzyme. That is, branching enzyme may
catalyse the formation of an increasing number of non-
reducing chain ends in the growing glucan polymer
which are then able to accept glycosyl residues from
ADP-glucose. Since it was shown (loc cit) that activities
of starch synthase in the mutant were unchanged in
comparison with the wild-type, the increased amylose/
amylopectin ratio of the ae mutant starch appears to be
due to the deceased action of branching enzyme.
Di†erences between the isoforms of maize SBE in
relation to activity and temperature optima have been
reported by Takeda et al (1993b). Di†erences were also
observed in the structure of the starches formed. In ae
maize mutants, SBE I preferentially transfers chains
which are longer than those transferred by SBE II. The
gene encoding for mature SBE I has been expressed in
E coli using the T7 promoter (Guan et al 1994). The
expressed SBE I was puriÐed to remove amylolytic
activity and the results conÐrmed earlier reports that
SBE I was able to branch amylose at a higher rate and
amylopectin at a much lower rate than SBE II. It may
be that SBE I and SBE II play di†erent roles in the
synthesis of amylopectin in vivo, ie SBE I could be
involved mainly in the synthesis of the B chains of
amylopectin, whereas SBE II could play a major role
in the synthesis of the A chains.
The joint contributions of ADP-glucose pyrophosp-
horylase and SBE to the control of the rate of starch
synthesis in developing pea embryos have been investi-
gated (Denyer et al 1995). Since the values for the Ñux
coefficients for both enzymes were found to be low, it
was suggested that neither enzyme was of overriding
importance in the control of starch synthesis in peas.
Experiments have been undertaken to suppress SBE
activity in transgenic potato plants through the expres-
sion of a cDNA for SBE of potato (Komann et al
1991) in antisense orientation (MuŽller-RoŽber and
Komann 1994). Surprisingly, neither the amylose nor
the total starch content was altered. Possible expla-
nations advanced (MuŽller-RoŽber and Komann 1994)
are that either the degree of down regulation in the
plants selected was not high enough or that another
enzyme capable of branching linear glucans is present in
potato tubers. If the latter is correct, the di†erent func-
Starch for industry 301

TABLE 6
Starch properties of three mutant genotypes compared to their parent cultivars

Mutanta Parenta Mutanta Parenta Mutantb Parentb

waxy Oderbrucker high amylose Glacier shrunken Bomi

Gene symbol wx amo1 shx

Chromosome 1 5 È
Starch content (mg per grain) 17É6 18É7 24É8 29É5 9É6 30É9
Amylose (% of starch) 5É8 28É3 43É4 28É5 28É0 26É0
A-type granules
Number ( ] 106)c 36 26 21 28 35d 12
Mean diameter (lm) 9É6 11É0 11É1 12É3 5É2d 13É1
Mean volume (lm3) 604 873 787 1242 205d 1663
B-type granules
Number ( ] 106) 166 140 222 323 È 89
Mean diameter (lm) 2É34 2É60 4É30 2É45 È 3É70
Mean volume (lm3) 8É7 13É3 72É0 10É9 È 37É7
Proportion of A-type granules
(% of Total) By number 18 15 8 8 È 12
By volume 94 92 50 91 È 85

a McDonald et al (1991).
b Schulman et al (1994).
c Number of granules per grain.
d Intermediate granules, no bi-modal distribution of A-type and B-type granules observed.

tions of the two enzymes in potato require to be eluci-
dated.
It has long been known that debranching enzymes
are present in developing cereal endosperms. However
their role (if any) in starch synthesis has not been
explained. One possibility is that they may be required
to produce primers for the starch synthase reaction
(Du†us 1993). More recently, Ball et al (1996) have pro-
posed a model to explain the processing mechanisms
which are thought to precede the formation of amylo-
pectin from “pre-amylopectinÏ. The model depends on
the observation that tissues synthesising “phytoglyco-
genÏÈa more branched form of amylopectinÈare
associated with reduced or absent debranching enzyme
activity. In the Chlamydomonas species used, it was
found that no other enzyme activities appeared to be
deÐcient. Hence it was concluded that debranching or
“glucan trimmingÏ is mandatory for starch biosynthesis.
Thus the model describes how pre-amylopectin trim-
ming may proceed through the selective action of
debranching enzymes. The debranched oligosaccharides
generated may then be used by starch synthases to
form amylopectin, provided sufficient SBE activity is
present. It seem likely that debranching enzyme activ-
ity may prevent the accumulation of phytoglycogen.
MUTATIONS AFFECTING STARCH
SYNTHESIS
Enzymes of starch synthesis can be a†ected by naturally
occurring or induced mutations. Much work has been
done in barley, where mutations are readily observed
due to the diploid nature of the species. Although there
is some interest in growing barley for starch in areas
where maize has to be imported (Vasanthan and Bhatty
1996), an understanding of barley starch mutations is
likely to be most relevant in its application to other
crops. The synteny between graminaceous genomes
means that rice/maize/barley/wheat can all be com-
pared (Korzun and KuŽnzel 1997). In addition, barley
genes, once identiÐed and isolated, could be used to
modify more readily manipulated species such as wheat
or potato. Thus, it is appropriate to use barley as a
model in reviewing current knowledge of genetic varia-
tion in starch composition.
Variation in starch synthetic enzymes
The waxy (wx) mutation is associated with a change at
a single locus on the short arm of chromosome 1 (7H).
It reduces the level of GBSS. Domon (1996) suggested
that this was due to partial suppression of gene expres-
sion, as the wx allele preserves the structural gene for
the enzyme intact. Waxy barleys have amylose contents
of 2È10%, higher than those of other waxy cereals
(Tester and Morrison 1992). However, Ishikawa et al
(1994) have identiÐed lines from a mutation programme
in which GBSS appeared to be completely absent.
These lines were characterised as amylose-free. Further
research is necessary to determine whether this muta-
tion has a di†erent mechanism from wx, which most
closely resembles the waxy mutation in other crops.
 302
TABLE 7
Starch granules characteristics associated with RisÔ mutants of barley
(a) Starch granule size distribution

Mutant Gene Chromosome Starch content A-type granules B-type granules

numbera symbol
mg per grain % grain wt Number Mean diameter Mean volume Number Mean diameter Mean volume
(]106)b (mm) (mm3) (]106)b (mm) (mm3)

8B lys4d 5 (1 H) 24É18 48É27 17 13É0 1357 29 3É51 36É3

16B È 1 (7 H) 17É09 34É55 17 11É3 901 36 3É44 33É6
17B È 1 (7 H) 18É51 38É49 No apparent bi-modal distribution of granules
29C lys5g 6 (6 H) 22É67 47É40 11 14É6 1882 28 4É39 72É3
56C Hor2ca 5 (1 H) 29É12 54É42 19 13É5 1561 30 3É44 29É6
527B lys6i 6 (6 H) 24É29 45É30 19 12É7 1231 22 4É25 58É0
1508B lys3a 7 (5 H) 29É73 50É84 19 13É5 1515 49 3É93 45É8
Bomi 39É07 53É96 15 15É3 2152 247 3É54 33É7

(b) Associated e†ects on starch granule characteristics

Mutant L ysophospholipid Gelatinisation Swelling

numbera content (L PL ) temperature factor
(mg per 100 g) (T ) (¡C) at 80¡C
p
8B 695 57É1 10É1
16B 1164 58É9 8É5
17B 1746 55É1 7É2
29C 986 57É6 8É7
56C 1002 54É9 9É0
527B 469 54É4 10É4
1508B 760 58É2 12É8
Bomi 1002 57É4 9É2

R P Ellis et al
a B mutation in the cultivar Bomi ; C mutation in the cultivar Carlsberg II.
b Number of granules per grain.
Starch for industry
This will greatly assist in understanding how amylose
synthesis is partially or completely blocked at the
molecular level.
In barley the amylose content of endosperm starch
can be raised to around 40È45%. This is associated
with the single recessive gene, amo1, on chromosome 5
(1H) (Schondelmaier et al 1992), but changes in speciÐc
enzymes have not yet been attributed to this mutation.
From crosses between waxy and high amylose (amo1)
parents, it is possible to identify segregants expressing
both mutant genes. By using iodine staining, Schondel-
maier et al (1992) suggested that these segregants had
around 15% amylose, but this seems unlikely given the
e†ect of the wx gene in reducing GBSS expression.
Alterations in the structure of amylopectin could inÑu-
ence the extent of iodine binding, so iodine staining may
not be the most suitable determinant of amylose
content in this instance. It is necessary to elucidate the
e†ect of combining waxy and high amylose mutations
on molecular structure. The starches produced may
have substantial changes in chain lengths thus altering
the properties of the starches and making them more
suited for some industrial processes.
A spontaneous recessive mutant, shx (Schulman and
Ahokas 1990), has a characteristically shrunken appear-
ance of the endosperm due to a 70% reduction in starch
content (Table 6). Several starch synthases have lower
activities in this mutant, but the site of the mutation is
in the gene for the primer-independent form of soluble
starch synthase SSS (Schulman and Ahokas 1990).
GBSS activity does not appear to be a†ected by the
mutation (Schulman et al 1994).
Variation in granule structure and composition
Waxy starch genotypes showed overall reductions of
about 10% in A-type granule size and 1.6% in both
A-type and B-type granule numbers per grain compared
to non-waxy types (Table 6) so the ratio of A-type to
B-type granules was una†ected. By contrast, the high
amylose type was associated with a reduction in A-type
granule volume in comparison to the wild type, but a
signiÐcant increase in the volume of the B-type granules
which changed from less than 10% to around 50%
(McDonald et al 1991) of the total granule volume.
Lines with recessive alleles of both waxy and high
amylose genes were characterised by starch granules
which were much reduced in size and heavily embedded
into the surrounding protein matrix (Swanston 1994),
but no data on relative proportions of A-type and
B-type granules have been published.
The shx mutant showed a reduction, compared to the
wild type, both in granule number per endosperm and
in the size of the A-type granules. The usual bimodal
size distribution of A-type and B-type granules was not
observed (Schulman et al 1994).
303
Other barley mutations a†ecting starch composition
E†ects on starch granule size and number have been
demonstrated for mutations apparently unrelated to
starch synthesis. Examples are the high lysine mutations
induced at a number of loci on chromosomes 1 (7H), 5
(1H), 6 (6H) and 7 (5H) in the cultivars Bomi and Carls-
berg II (Table 7) in which Tester et al (1993) demon-
strated variation in granule dimensions, gelatinisation
proÐles and lipid contents. All the mutants showed a
reduction in total starch content, with the volume of
A-type granules lower and the number of B-type gran-
ules per endosperm greater than the wild type. Starch
from RisÔ 17 lacked a bimodal granule size distribution.
It was composed of a large number of intermediate
sized granules with a mean volume about three times
that of the B-type granules in the wild type. RisÔ 17 also
had a higher starch lipid content than the wild type,
while starch lipid content of RisÔ 527 was greatly
reduced. Starches from both these mutants, however,
had lower gelatinisation temperatures than those of
wild-type starches (Tester et al 1993) and this factor
may be of importance in energy considerations during
industrial processing.
STARCH CHEMISTRY
Composition of starch
The principle components of starch are the polymers of
glucose, amylose (MW 105È106) and amylopectin (MW
107È109). Iodine complexes with amylose to generate a
strong blue colour (Yu et al 1996), which forms the basis
of many methods for determining amylose contents
(Martinez and Prodolliet 1996). Improvements in the
Ñexibility and accuracy of such assays can be achieved
by recording absorbance measurements at more wave-
lengths (Jarvis and Walker 1993), but colorimetric
methods su†er interference from the limited iodine
binding ability of amylopectin (Davis and Khan 1994).
Enthalpies of melting of starchÈlipid complexes have
been used to obtain amylose contents for various
starches (Sievert and Holm 1993), but the efficiency of
this method is inÑuenced by the solubility of the
amylose in each sample. Amylose contents can be deter-
mined with greater accuracy by gel permeation chroma-
tography (GPC) (Salomonsson and Sundberg 1994) but
the method is not suitable for routine investigation.
High pressure size exclusion chromatography has also
been used to measure the amylose content of starch
(Bradbury and Bello 1993). This method is more rapid
than the time-consuming GPC. A novel procedure for
the determination of amylose after complexing the
amylopectin with concanavalin A can be used on a
routine basis (Gibson et al 1997).
304
Amylose
Debranching (Takeda et al 1992) and chain length
studies (Shibanuma et al 1994) on amylose have demon-
strated that this a-(1È4) linked glucose polymer is
actually a mixture of unbranched and randomly, lim-
itedly branched polymers. The proportions of branched
to unbranched amylose vary both within and between
species (Takeda et al 1987 ; Shibanuma et al 1994),
whilst starches from di†erent botanical sources di†er in
the average number of chains their amylose molecules
possess (Takeda et al 1987). Methylation (Cura and
Krisman 1990 ; Cura et al 1995), debranching (Praznik
et al 1994), and NMR studies (Falk et al 1996) have
respectively revealed the percentage of glucose residues
involved in branching for the amyloses of waxy, normal
and high amylose maize as 1É4%, 1É0È2É0% and
0É256%.
Amylose can be extracted from starch by leaching in
hot (50¡C) water. As the resulting gel or paste cools,
aggregation (retrogradation) occurs. This involves the
alignment of linear segments of amylose chains which
associate into a more thermodynamically stable form
through hydrogen bonding. Thus, the apparent molecu-
lar size of the amylose increases and its solubility
decreases, with insoluble particles eventually being
formed. The type of aggregation displayed by amylose is
dependent on chain length, shorter chains favouring
precipitation and longer ones favouring gel formation
(Gidley and Bulpin 1989). The degree of retrogradation
of amylose is governed by its moisture content and, at
high temperature, the crystallinity of the amylose is
enhanced (Hoover 1995). Retrograded amylose is more
resistant to degradation by amylolytic enzymes than the
native form, but amyloseÈlipid complexes (see the
section entitled Starch lipids) restrict retrogradation and
the starch remains accessible to such enzymes
(Czuchajowska et al 1991). However, the solubility of
these complexes is less than that of lipid-free amylose
owing to the hydrophobicity of the lipids. The extent of
retrogradation is of signiÐcance in applications where a
clear solution or product is required as well as in those
which depend on the viscosity of the product.
Amylopectin
Amylopectin consists of short a-(1È4) linked chains
which are connected by a-(1È6) bonds. Debranching
studies on starches from a range of maize genotypes
revealed maize amylopectin to have a bimodal chain
length distribution. Amylopectin from starch with the
ae genotype has a higher average chain length and
lower degree of branching than wild type or waxy
starches (Wang et al 1993). Amylopectin from the ae/wx
genotype shows a bimodal proÐle whilst those from the
du/wx genotype (Yuan et al 1993) and amylomaize
(Takeda et al 1993a) are trimodal. Amylopectins from
high amylose starches have lower degrees of branching
than those from waxy and normal starches, with the
R P Ellis et al
highest level of branching being that of waxy maize
(Salomonsson and Sundberg 1994 ; Falk et al 1996). The
chain length in high amylose starch amylopectin is
greater than that in normal or waxy starches (Wang et
al 1993). Amylopectins from oat starches have a tri-
modal chain length distribution and, in starches which
have a higher lipid and amylose content, there is less
multiple branching in the amylopectin (Wang and
White 1994). Debranched amylopectins from normal
wheat, investigated by HPLC, have a tetramodal
proÐle, the pattern of chain length distribution varying
with genotype (Shibanuma et al 1994). The proÐle of
waxy wheat amylopectin chain length distribution,
analysed by high performance anion exchange chroma-
tography is trimodal (Yasui et al 1996). NMR studies
showed that the proportion of glucose units which are
a-(1È6) linked in barley, potato and waxy maize
starches are 4É2È5É4%, 6É0% and 3É0, respectively
(McIntyre et al 1990 ; Salomonsson and Sundberg 1994 ;
Falk et al 1996).
Intermediate fractions
Using 13C-NMR, a component with a similar structure
to amylopectin but with shorter side branches has been
found in wheat starch (Dais and Perlin 1982). In maize,
intermediate material considered to be low-molecular-
weight amylopectin has been found at high levels in
mutants with the ae genotype (Wang et al 1993). An
intermediate component with longer branches than
amylopectin was reported in oats by Wang and White
(1994) but Tester and Karkalas (1996), using a contin-
uous quantiÐcation procedure which eliminates the
need for starch fractionation, did not detect this
material and suggested it was an artefact of the Wang
and White (1994) fractionation procedure.
Starch lipids
Lipids extracted from starch granules may be integral
components of the granules or have originated else-
where in the tissues and become associated with the
granules during the starch extraction process. The lipid
composition of starches from a variety of sources has
been described by Vasanthan and Hoover (1992), whilst
the lipids in cereal starches have been extensively
reviewed by Morrison (1988, 1993, 1995). Starch gran-
ules from cereal endosperms appear to be unique in
having lipids as integral components. Granules isolated
from potato tubers, legume seeds or from cereal tissues
other than the endosperm do not contain integral lipids
(Galliard and Bowler 1987).
Three categories of lipid are associated with cereal
starch granules. The non-starch lipids originate in the
membranes and spherosomes of the endosperm. These
are loosely associated with the granules and consist of
triacylglycerides, diacylglycolipids and phospholipids.
During storage of the grain, some lipolysis of these
compounds takes place, resulting in free fatty acids and
monoacylglycerides being formed. Even slight swelling
Starch for industry
of the granules during the starch extraction process
allows these free fatty acids and monoacyl lipids to be
absorbed into the surface layers of the granules. These
are described as starch surface lipids. The true or inte-
gral starch lipids are found within the intact granules.
In barley and wheat, these lipids are composed entirely
of lysophospholipids, while in oats, rice and maize, free
fatty acids make up 30È60% of the integral lipids of the
granule (Morrison 1993). The starches from waxy
mutants of maize, barley, wheat and rice have negligible
or very low amylose contents and much lower lipid con-
tents than the normal genotypes, and the starches of
high amylose maize, barley and rice have higher lipid
contents than the starches of normal genotypes
(Morrison 1995 ; Yasui et al 1996). Oat starches have
higher lipid, amylose and phosphorus contents than
other cereal starches (Sowa and White 1992). No matter
the source, the amount of amylose present is more than
sufficient to accommodate all the lipid within the
amylose helix in six- or seven-membered inclusion com-
plexes. There has been uncertainty about whether these
complexes exist in the intact starch granules or whether
they are formed when granules were swollen and
hydrated, but not necessarily gelatinised (Morrison
1988). Studies using 13C cross-polarisation/magic angle
spinning (CP/MAS) NMR spectroscopy and DSC have
conÐrmed that lipidÈamylose complexes and lipid-free
amylose are both present in intact barley starch gran-
ules (Morrison et al 1993).
Monoacyl lipids form complexes with hydrated
amylose in vitro. The molecular inclusion complexes are
heat stable and insoluble at neutral pH values
(Szczodrak and Pomeranz 1992). LipidÈamylose com-
plexes in the starch granule a†ect both the gelati-
nisation temperature and the swelling behaviour of the
granule (Morrison 1995). Eliasson (1994) demonstrated
that chemical modiÐcation of the starch a†ects the
thermal properties of the amyloseÈlipid complex. In-
direct evidence for the formation of an amylopectinÈ
lipid complex was also reported. Since lipids and amyl-
ose are positively related in cereal starches, the proper-
ties of these complexes a†ect starch properties (see the
section entitled Amylose).
Starch proteins
Proteins are also associated with starch granules and
the amounts vary between and within species (Table 4).
The protein has been classiÐed as either surface protein
(readily extracted at temperatures below the gelati-
nisation temperature) or as integral protein (extractable
at temperatures near or above the gelatinisation
temperature). About 10% of the protein extracted from
wheat starch granules was reported to be surface
protein (Galliard and Bowler 1987). One of the granule
surface proteins, friabilin, has been linked with kernel
hardness (Anjum and Walker 1991). Kernel hardness
a†ects the energy costs incurred with extraction of the
305
starch granules and the amount of friabilin present in a
cultivar may be a useful marker for softer genotypes.
Some of the enzymes catalyzing the synthesis of
starch polysaccharides are immobilized within the
structure of the starch granule as it forms while others
remain soluble and are retained by the amyloplast
membrane. In starches isolated from mature cereal
endosperms, it is not possible to detect the amyloplast
membrane by microscopy. Rahman et al (1995) reported
Ðve major protein bands of 60, 75, 85, 100 and 105 kDa,
respectively, in extracts from wheat starch granules
extracted at 100¡C in 50 mM Tris (pH 8) with 10% SDS
and separated by SDS-PAGE. The most prominent
band (60 kDa) was identiÐed as GBSS. This band is
absent from the starches of waxy barley, waxy maize,
waxy rice and waxy potato (Shure et al 1983 ; Preiss
1991 ; Sivak et al 1993). Rahman et al (1995) showed
cross reactivity between the 85 kDa protein band and
an antibody to SBE IIb of maize and the 75 kDa band
was tentatively identiÐed as a starch synthase based on
the similarity of amino acid sequence with SSS from rice
starch.
All starch granules contain detectable amounts of
protein. Due to the possibility of Maillard-type reac-
tions taking place with polysaccharides, the amount and
type will a†ect any new uses of the starch or its applica-
tion to existing uses such as wheat starch for paper
making.
Inorganic components
Starches contain a number of mineral components such
as Ca2`, K`, Mg2` and Zn2` ions as well as bound P.
In cereal starches, most, if not all, of the P is in the
lysophospholipid fraction (Morrison 1995) while, in
potato starch granules, the P is present as phosphate
which is bound to the amylopectin component through
phosphomonoester linkages (McIntyre et al 1990)
(Table 4). These phosphomonoester groups make the
starch cationic and this is one reason why potato starch
is favoured by papermakers.
The structure of starch granules
The composition and structure of individual starch
components determine many starch properties, espe-
cially in solution, but the arrangement of these com-
ponents within the granule determine its structure for
applications requiring intact or modiÐed granules.
Under polarised light, native starch granules display
birefringence in the form of a “Maltese crossÏ pattern.
This property reÑects the order of the granule. The crys-
talline nature of starch is evident from wide-range X-ray
studies, and starches from di†erent botanical sources
yield di†erent X-ray patterns (Cameron and Donald
1992). The sharpness and deÐnition of the crystallinity
306
pattern gives evidence of crystallite quality, whilst the
degree of crystallinity can be assessed by comparing
amorphous and highly crystalline (ie waxy) starch
(Koksel et al 1993). Crystallinity is derived from the
structure of amylopectin, with its radial arrangement in
the granule being at right angles to the granule surface
(Davis 1994). This radial arrangement is built from a
series of stacked amylopectin clusters, each consisting of
a region containing branch points and another region
containing ordered double helices comprised of short
chains of amylopectin (Jenkins et al 1993). From DSC
studies, it has been suggested that the enthalpy gener-
ated during gelatinisation may be related to the loss of
this double helical order (Cooke and Gidley 1992). The
di†erent cluster components can be related to growth
rings which have been identiÐed in starch granules by
electron microscopy (Galliard and Bowler 1987). The
cluster branch points and amylose are found in the
amorphous zones or lamellae of the granule, where they
are held together by hydrogen bonds (Trommsdor† and
Tomka 1995), whilst the double helices are located in
the crystalline regions (Jenkins et al 1993). Small angle
X-ray di†raction studies give repeat distance (d-spacing)
of 9È10 nm which is thought to represent the average
size of an amylopectin cluster and the consistency in
starches from a variety of species suggests similarities in
the mechanisms by which amylopectin is synthesised in
di†erent plants (Jenkins et al 1993).
When granular starch was treated with cross-linking
agents and the reaction products examined by GPC
and 31P NMR, it was found that amylose cross-linked
with amylopectin or formed substitution products
within the granule. Whilst amylose, after solubilisation
in DMSO, was capable of cross-linking, there was no
evidence of amyloseÈamylose cross-linking when granu-
lar starch was treated. It was concluded that amylose
was dispersed randomly throughout the granule rather
than located in bundles (Kasemsuwan and Jane 1994).
Leaching experiments on native and defatted starches
indicate that the degree of organisation of amylose in
the amorphous regions of starch varies with botanical
source (Hoover and Vasanthan 1992).
FUTURE PROSPECTS
Starch has an advantage over other materials with
similar properties in that it is a renewable and biode-
gradable resource which can be chemically modiÐed to
produce a diverse range of products. Industries will be
able to use starches more efficiently and for a wider
range of applications only if starches with the appropri-
ate properties become available. This can be achieved,
to some extent, if further progress is made in the chem-
istry of starch modiÐcation, but greater opportunities
R P Ellis et al
for generating starches with speciÐc properties will
come from our ability to extend the range of the chemi-
cal composition of the native starches produced in crop
plants.
There are some wide gaps in our knowledge of the
relationship between starch chemistry and starch
properties. This is partly because the architecture of the
starch granule is inherently complex and partly because
our methods of analysis have not been sufficiently sensi-
tive or sufficiently probing. Since 1992, a number of
techniques have been developed which have the poten-
tial to improve this situation. A procedure for both
chain length determinations and the calculation of
amylose/amylopectin ratios has been introduced which
is a modiÐcation of existing size-exclusion chromato-
graphic methods. Starch is dissolved in sodium hydrox-
ide and is then applied to a three-column system, using
a sodium chloride solution as elutant. This technique
represents an advance over existing procedures since it
uses less toxic materials and is completed in only
30 min. Furthermore, the use of sodium hydroxide
enables heat-treated (ie partially gelatinised) starches to
be analysed (Kennedy et al 1992). Jarvis and Walker
(1993) have proposed a method of measuring actual
amylopectin and amylose concentration by measuring
the absorbances of starchÈiodine complexes at six di†er-
ent wavelengths. Advantages claimed over previous
methods include rapidity of use and greater accuracy.
The use of electrophoretic methods involving Ñuoro-
chrome labelling (OÏShea and Morell 1996) promises to
increase both the speed and sensitivity of chain length
analysis and the range of oligosaccharide chain lengths
which can be quantiÐed accurately following isoamylase
digestion of starch.
Electron optical tomography is a technique which
can be used to build up the three dimensional structure
of starch granule fragments from a series of transmis-
sion electron microscope sections. This procedure has
the potential to reveal details of crystalline structure
(Oostergetel and van Bruggen 1993) and could be used
with X-ray di†raction (Jenkins et al 1993 ; Koksel et al
1993) and di†erential scanning calorimetry (Cooke and
Gidley 1992) to investigate the ordered structure of
starch. NMR is also being used in conjunction with
these techniques to increase our understanding of starch
structure (McIntyre et al 1990).
The chemistry and properties of starch granules are
under genetic control and are also signiÐcantly a†ected
by the environmental conditions experienced by the
plant during starch deposition. The research that has
been carried out on the e†ect of environmental factors
on starch composition and properties has concentrated
on only one factor, namely temperature. The complex
interaction of other factors remains to be explored.
Further investigation of the e†ects of environmental
factors will help us to understand and predict variations
in the chemistry and properties of native starches but,
Starch for industry
to produce starches of speciÐed and novel chemical
composition and properties, it will be necessary to alter
the genome of starch-producing crop plants. Progress
has been made using conventional breeding methods as,
for example, in the identiÐcation of a range of starch
mutants in barley and maize and in the identiÐcation of
the amylose-free potato mutant. Similar modiÐcations
of the genome of wheat will increase greatly the diver-
sity of native starches available for industrial uses.
The mutants available for the study of starch synthe-
sising enzymes have allowed the pathway of starch syn-
thesis to be worked out in detail, especially in maize
(Nelson and Pan 1995). In addition to gene cloning, the
development of new DNA marker technology has
allowed the creation of densely marked gene maps. In
barley, it has been shown that ADP-glucose pyrophosp-
horylase activity is controlled by at least four genes
(Kilian et al 1994). The large subunit expressed in the
endosperm is coded by a gene, Aga7, on the short arm
of chromosome 5 (1H) and the small subunit by Aga1,
on the long arm of chromosome 1 (7H). The ADP-
glucose pyrophosphorylase activity seen in the leaf is
also coded by two genes ; the large subunit by Aga6,
which was mapped to the log arm of chromosome 5
(1H) and the small subunit by Aga5 on the short arm of
chromosome 7 (5H). The complexity of genetic control
in di†erent tissues allows a Ñexible response to regula-
tion e†ectors.
Progress has already been made in the identiÐcation
of the genes which control the process of starch synthe-
sis in transgenic potatoes. From this will follow the
achievement of the goal of creating, in agronomically
acceptable crop plants, genomes which programme the
production of starches with properties appropriate to
speciÐc applications.
ACKNOWLEDGEMENTS
This review was funded by The Scottish Office Agricul-
ture, Environment and Fisheries Department.
The authors wish to thank all their colleagues who made
useful suggestions while the review was being written.
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