CEREAL CHEMISTRY 294
Synergistic and Additive Effects of Three High Molecular Weight Glutenin
Subunit Loci. I. Effects on Wheat Dough Rheology
S. Uthayakumaran,
1,2
H. L. Beasley,
3
F. L. Stoddard,
4
M. Keentok,
1,2
N. Phan-Thien,
1,2
R. I. Tanner,
1,2
and F. Békés
1,3,5
ABSTRACT Cereal Chem. 79(2):294–300
The high molecular weight glutenin subunits (HMW-GS) play an
important role in governing the functional properties of wheat dough. To
understand the role of HMW-GS in defining the basic and applied rheo-
logical parameters and end-use quality of wheat dough, it is essential to
conduct a systematic study where the effect of different HMW-GS are
determined. This study focuses on the effect of HMW-GS on basic
rheological properties. Eight wheat lines derived from cvs. Olympic and
Gabo were used in this study. One line contained HMW-GS coded by all
three loci, three lines were each null at one of the loci, three lines were
null at two of the loci and one line null at all three loci. The flour protein
level of all samples was adjusted to a constant 9% by adding starch. In
another set of experiments, in addition to the flour protein content being
held at 9%, the glutenin-to-gliadin ratio was maintained at 0.62 by adding
gliadin. Rheological properties such as elongational, dynamic, and shear
viscometric properties were determined. The presence of Glu-D1
subunits (5+10) made a significantly larger contribution to dough prop-
erties than those encoded by Glu-B1 (17+18), while subunit 1, encoded
by Glu-A1, made the least contribution to functionality. Results also
confirmed that HMW-GS contributed to strength and stability of dough.
Understanding the role of the various constituents of wheat
flour in determining the functional properties of dough has been a
major endeavor throughout the 20th century. With this knowledge,
better wheat cultivars can be bred and more economical manufac-
turing technologies can be developed. The protein in wheat is mainly
responsible for inherent differences in baking quality (Finney 1943),
with both quantity and quality of the protein being important (Finney
and Barmore 1948). It has been well established that two classes
of proteins, glutenins and gliadins, are the key components in deter-
mining the rheological properties of dough as they interact to form
the gluten matrix during dough mixing. Glutenins are polymers of high
molecular weight glutenin subunits (HMW-GS) and low molecular
weight glutenin subunits (LMW-GS), which differ in their mobility
during SDS-PAGE. The HMW-GS have the largest role in defining
dough strength (MacRitchie 1992) and possibly stability (Payne 1987).
Bread wheat (Triticum aestivum L. emend Thell.) is a hexaploid
species with three genomes designated A, B, and D. The Glu-1
loci on the long arms of the homoeologous Group 1 chromosomes
code for the HMW-GS, and are designated Glu-A1, Glu-B1 and
Glu-D1, respectively. Each locus generally codes for two subunits,
“x” and “y”, although the Glu-A1 y-type gene is usually not
expressed. Each locus also has several alleles, differences between
which account for a considerable proportion of the variation in dough
strength (Payne et al 1987). Null alleles, where no subunits are
produced at all, have been developed at each of the three Glu-1
loci (Lawrence et al 1988). This set of lines provides a valuable
resource for identifying the effect of each Glu-1 locus on overall
rheology and end-product quality. The set has already been used
for studies on polymeric protein formation (Gao and Bushuk 1993;
Gupta et al 1995), but without correction to a constant protein content.
For these reasons, basic rheological tests (elongational, shear
viscometry, and dynamic oscillatory tests) and end-use quality tests
(breadmaking) were conducted. Fundamental or basic rheological
tests are considered to be valuable tools in determining dough
functionality and unlike the empirical dough measurement tests
(mixograph, farinograph, extensigraph, and alveograph), are founded
on parameters of basic physics (Levine 1987). The rheological prop-
erties of dough change considerably during every baking phase
and the viscoelastic properties of dough have a profound effect on
the finished product (Walker and Hazelton 1996). The basic rheo-
logical measurements can be used for constitutive modeling of
bread dough and this may lead to predictions of dough processing
behavior.
MATERIALS AND METHODS
The eight flour samples differed by the activity (designated +) or
inactivity (designated – or null) of each of the three glutenin loci
(Table I). The ash content was determined using Approved Method
08-01 (AACC 2000). The nitrogen content of the flours was deter-
mined by the Dumas total combustion method using an elemental
analyzer (CHN–1000, Leco Inc., St Joseph, MI). The protein
content (%) was determined as N × 5.7. The moisture content of the
flours was determined by a near infrared spectrometer with sample
transport (Foss NIRSystems 6500, Silver Spring, MD). Flour
protein level of all samples was adjusted to a constant 9% by
adding commercial wheat starch (1.6% protein). The ratio of glutenin
to gliadin, ratio of high molecular weight glutenin subunits to low
molecular weight glutenin subunits (HMW-GS/LMW-GS ratio) and
the unextractable polymeric protein percentage (% UPP) (Table I)
of the samples were determined using reversed-phase HPLC
(Marchylo et al 1989) and size-exclusion HPLC (Singh et al 1990;
Batey et al 1991; Gupta et al 1993), respectively.
The effect of maintaining the glutenin-to-gliadin ratio constant
was investigated in a subset of the flours. The choice of materials
was limited to four by the quantity of flour available. Gliadin (85%
protein) was extracted from flour of the line null for all three HMW-
GS loci using the mild pH extraction method of MacRitchie
(1985). Lyophilized gliadin was added to + + +, + – + and – – +
flours to achieve a uniform glutenin-to-gliadin ratio of 0.62, as
found in the – – – sample. Commercial starch was also added to
maintain the constant protein content of 9%.
Mixing was conducted at a water absorption of 57%. Blended
flour and water were mixed in a 10-g mixograph to peak dough
development and rheological measurements were then made. Mixing
was done in triplicate for each sample and the mean peak dough
development time was calculated from the mixing curve (Gras et
al 1990).
1
Quality Wheat Cooperative Research Centre Ltd., Locked Bag 1345, North Ryde,
NSW 1670, Australia.
2
Department of Mechanical and Mechatronic Engineering, University of Sydney,
NSW 2006, Australia.
3
CSIRO Plant Industry, Grain Quality Research Laboratory, PO Box 7 North
Ryde, NSW 1670, Australia.
4
School of Applied Sciences, University of Wolverhampton, Wulfruna St.,
Wolverhampton WV1 1SB, UK.
5
CSIRO Plant Industry, GPO Box 1600 Canberra, ACT 2601, Australia. Corres-
ponding author: Phone (Sydney Office): +61 2 9490 8437. Fax: +61 2 9490
8419. E-mail: F.Bekes@pi.csiro.au
Publication no. C-2002-0211-02R.
© 2002 American Association of Cereal Chemists, Inc.