J ournal of Cereal Science21(1995) 215230

Mini Review
Gas Cell Stabilisation and Gas Retention in Wheat
Bread Dough
Z. G an, P. R. Ellis and J. D. Schoeld
The University of Reading, Department of Food Science and Technology, W hiteknights, PO Box 226,
Reading RG 6 6AP, UK and KingsC ollege London, Division of Life Sciences, C ampden Hill Road,
London W 8 7AH, UK
Received 1 M ay 1994
Dedication
This paper is dedicated to thememory of Professor Terry Galliard, a good friend and
colleague, whomadeinvaluablecontributionstotheworkcarriedout inour laboratory,
which is described in this review, and to Cereal Sciencein general.
ABSTRACT
Gascell stabilisation and gasretention areof considerableinterest becauseof their technological sig-
nicanceinbreadmaking. Wereviewrecent studiesinrelationtothestabilisationof gascellsandthe
mechanismsof gasretention, anddiscusshowthesemaybeaectedbytheliquidphaseof dough. The
possibility isdiscussed of theinvolvement of surfaceactivematerials, such asproteinsand pentosans
dissolved in thedough aqueousphase, and, perhapsmoreimportantly, non-starch polar lipidsin the
formation andstabilisation of gascells. Thereisaccumulatingevidencefor thehypothesisthat liquid
lmsplayacritical roleinthemechanismsof gasretentionindough. Thehypothesisproposesthat two
closelyrelated, consecutivestagesareinvolvedindoughexpansion. Duringtherststage, theexpanding
gascellsremain discreteuntil discontinuitiesdevelopin thestarchprotein matrix, leavingareascon-
taining only a liquid lm. The timing and the degree to which such discontinuities occur is largely
dependent on gluten proteins. Thesecond stageinvolvesan increasein thesurfaceareaof theliquid
lmasdiscontinuitiesbecomeincreasingly frequent duringexpansion. Failureof thelamellar lmto
maintaintherateatwhichnewsurfaceareaisgeneratedleadstotheruptureofthislmand,consequently,
thelossof gasretention. Consideration isalso given to theroleof bakery fat in gasretention and to
additional factorsthat aect gasretentioninwholemeal doughs, inparticular thephysical disruptionof
thefoamstructureof suchdoughsbycomponentsof theouter layersof thegrain.
Keywords: gascells, gas retention, bread dough, liquid lm
INTRODUCTION
\nnnr\i\+ioxs tsrn: PoL=polar lipids; NPoL=non-
The unique position of wheat, compared with
polar lipids; NSL=non-starch lipids; ESP=equi-
other cereals, inbreadmakingisduetotheability
librium-spreading pressure; FA=free fatty acid;
of wheat our dough to retain gas on expansion.
PUFA=polyunsaturated fatty acid; LCL=liquid
To producea loaf of bread with a light and even
crystalline; SEM=scanning electron microscope;
crumb texture, the dough must be able to retain
CBP=Chorleywood Bread Process; SSL=sodium
stearoyl lactylate. the gases produced by yeast fermentation as dis-
Corresponding author: J . D. Schoeld, The University of
cretegascellsfor asucientlylongperiod
1,2
. The
Reading.
gas cells play a crucial role in gas retention, but
howtheyareformed, andespeciallywhat happens
to themduringbread dough processing, ispoorly
07335210/ 95/ 030215+16 $08.00/ 0 1995 Academic Press Limited
215
Z. Gan et al. 216
understood. Although work duringthelate1930s Under normal baking conditions, the initial loss
of gas is slow, but a sharp increase in the rate of and early 1940s produced much useful in-
formation
3,4,5,6
, and considerable eort has since lossoccurstowardstheend of oven spring
14,15
. The
slow initial loss of gas can be explained by its beendevotedtotheelucidationof gascell structure
and the mechanisms of gas retention, present diusion to the external surface of the dough fol-
lowed by evaporation, whereas the rapid loss has knowledgestill remains fragmentary.
Thefoamstructure of a fermenting dough has beenattributedtotheruptureof thestarchprotein
matrix surroundinggas cells. Thelatter is thought long been assumed to be a dispersion of discrete
gas cells in a continuous starchprotein matrix
2,3,4
. toresult intheinterconnectionof adjacent gascells,
convertingthefoamstructureof doughintoanopen The matrix ruptures during baking, leading to
the establishment of a continuous gas phase and, sponge and enabling direct escape of the gas (Fig.
1). The rupture of the matrix has, in turn, been consequently, the rapid loss of gas. Mechanisms
involvingdierent factorsthat control gasretention attributed to the sharp increase in dough viscosity
that occurs mainly as a result of starch gel- have been proposed on this basis, emphasising
largely the role of the gluten proteins
7,8
and non- atinisation
16
. The improving eects on loaf quality
of oxidisingimprovers, whichincreasetheresistance starch polar lipids
9,10
. Results fromthis laboratory,
however, have indicated the involvement of in- of dough to deformation
17,18
, and, of fat and water,
which decrease the resistance of dough to de- terfacial lms at the gas/ liquid interfaces in
fermenting bread dough
11,12
. The components re- formation
18,19,20
arediculttoreconcileonthisbasis,
however. Furthermore, proteinsareknown to play sponsible for stabilising the lms are not yet
known, but are envisaged to be non-starch polar a crucial role in determining the gas-holding cap-
acity of a dough, but the above description does lipids and/ or proteins and possibly pentosans
dissolved in the aqueous phase of dough. not explain whyproteinsaresoimportant in bread
making. The polymerisation of glutenin molecules Studies with specic systems or individual our
componenthavetemptedcereal scientistsfrequently as a result of sulphydryldisulphide interchange
21
maycontributetothesharpincreaseintensilestress to attributethecentral roleto a singlecomponent
and to discount the involvement of the others in that occurs in starchprotein matrix on heating,
thus leading to the rupture of the bulk phase and bread making. Thepresent reviewaimsto provide
a critical discussion of this area, makingparticular to thelossof gasretention.
Othershavesuggested that gluten contributesto reference to the role of surface active materials in
gasretention. gasretentionbyslowingthediusionof gasthrough
the dough phase
8,22
. CO
2
is produced by yeast
fermentationintheaqueousphaseof dough. It then
GAS CELLS ANDDOUGH EXPANSION
diusesthrough theaqueousphaseto thegascells,
where it evaporates to generate within them an The gas phase of a proving dough exists as a
dispersion of discretegascellsin asemi-solidbulk excess pressure that provides the driving force for
dough expansion. The CO
2
cannot diuse out of dough phasecomprisingstarch, gluten and other
minor components. The gas cell nuclei, which thegascellsbecausetheaqueousphasesurrounding
them is saturated. The saturation of the aqueous later expand as carbon dioxide (CO
2
) produced
by yeast fermentation is transported to them, are phaseismaintained becausetheyeast continuesto
producemoreCO
2
. A small proportion of thegas incorporatedbytheocclusion of air duringmixing
5,6
.
Yeast is incapable of producing new gas cells in does diuse to the surface of the dough piece and
evaporatesinto thesurroundingatmospheregiving the dough
5
, although subdivision of existing cells
occurs during punching and moulding operations. aslowreleaseof gasfromthedough. Towardsthe
endof baking, however, asharpincreaseintherate The technological signicance of the gas cells im-
mediately after mixing is that a suciently large of loss of gas occurs. Although some researchers
have attributed it to an increase in the rate of number, estimated
13
to be between 10
11
and 10
13
/
m
3
, isrequired. At advancedstagesof breaddough diusion throughthedoughaqueousphase
8,22
, oth-
ers have argued that this sharp increase is due to processing, thesurvival of intact gascellsisof great
importance in order to prevent the loss of gas, the rupture of the starchprotein matrix
1,2
. Bread
dough undergoes a structural transformation from thereby producing bread with a light and even
crumb texture (i.e. with large specic volume and
gascellsof relatively uniformsize).
Gas cell stabilisation and gas retention 217
Foam
Sponge
= Gas cel l = Starchprotei n matri x
Loss of gas
Rupture of the matri x
Starch gel ati ni sati on at above 60C
I ncrease i n tensi l e stress
i n starchprotei n matri x
END OF BAKI NG
END OF MI XI NG END OF PROVI NG
Figure1 Thestructural transformation of dough as explained by theconventional starchprotein matrix hypothesis. Soon
after mixing, the structure of dough is represented by small gas cells dispersed in a continuous starchprotein matrix. Each
discrete gas cell expands in response to CO
2
production during fermentation, and the foamstructure is maintained by thin
membranes separatingadjacent cells at theend of proof. Duringbaking, starch gelatinisation induces a dramatic increasein
dough viscosity, resultingin arapidincreasein tensilestrength in themembrane. Thisinitiatesmembranerupture, converting
thefoaminto a sponge.
a foam into an open sponge during baking as
indicated by previous studies
3
and conrmed by
energy required to produce a given deformation
and therupturestress, for example, wasshown to
recentobservations(Fig. 2)madeinour laboratory
11,12
.
decrease with increasing water level
20
. In fact, at
Although diusion controlled processes will cer-
verysmall deformations, therheological properties
tainly be important for gas retention during fer-
of anexperimental doughcanbereproducedfrom
mentation, it seems more likely that the physical
any our by thecontrol of water addition
24
.
ruptureof thebulkdoughphaseand/ or liquidlms
A wheat our doughcontainstypically0608g
surroundinggascellsismorerelevant tothesudden
water/ gof dry our, of which approximately half
lossof gasduringbakingthananincreaseddiusion
is thought to be bound or unfreezable
25,26
. The
ratethrough thedough phase.
presence of free or freezable water can be de-
tectedonlywhenthewater content exceedsabout
WATERANDTHE LIQUIDPHASE OF DOUGH 3035% by weight. Further addition of water
forms a second aqueous phase that may be sep-
Irrespective of the process involved, the primary
arated by centrifuging the dough in very high
event in bread makingistheaddition of water to
centrifugal elds
23,27
. Theaqueousphaseisneces-
a dry our at the onset of mixing, forming a
saryin dissolvingsolubleour componentsandin
cohesive, viscoelastic dough. The mechanical or
providing themediumfor reactions to takeplace rheological behaviour of thedough is profoundly
inuenced by the amount of water added
23
. The in thedough
28
.
Z. Gan et al. 218
Figure2 SEM micrographsof wheat bread dough and bread prepared by theChorleywood Bread Process(CBP), showing
(A) theapparent interconnectionsin bread doughsat advanced stagesof proving, and (B) theopen spongestructureof bread
crumb at theend of baking.
Benecial eectsonloaf volumeandcrustcolour used to furnish therequired water in bread mak-
ing
28
. It has been shown also that eectively no were observed when the liquid phase (dough li-
quor) centrifuged from wheat our doughs was gasisretainedbydough belowawater content of
Gas cell stabilisation and gas retention 219
about 35% of the total dough mass
27
, but gas extensibility. Thesecond stageinvolves primarily
an increase in the surface area of theliquid lm, retentionimprovesalmost linearlywhenthewater
content is increased abovethis valueup to about which maintains the integrity of the gas cells as
discontinuities in the starchprotein matrix be- 44%. Electrical conductivitydatafurther indicated
that theaqueous phase is continuous rather than comeincreasinglyfrequent duringexpansion. The
behaviour of thedoughat thisstageisdetermined comprisingdisperseddroplets(MacRitchie, 1976).
MacRitchie
27
envisaged a structure of dough in primarilybythestabilityof theliquidlm. Surface
active materials, such as endogenous our polar which gas cells are embedded in a continuous
starchprotein matrix and are enveloped by a lipids, andproteinsandpentosansdissolvedinthe
doughaqueousphasemaycontributepositivelyto continuousthin liquid lm. Although it isdicult
toassessthetechnological signicanceof theliquid gas retention by stabilising thelmso that it can
expandtoalarger surfaceareawithout rupturing. lm in gas retention in relation to that of the
mechanically strong solid dough phase, the hy- It istheruptureof theliquidlm, rather thanthat
of the starchproteinmatrix, that isthought tolead pothesisoersanewwayof thinkinginthesearch
for a better understanding of the mechanisms of totherapidlossof gasandtotheendof ovenspring
11,12
.
Gluten proteins undoubtedly play a major role gas retention.
in determining the baking performance of wheat
ours through their eects on dough rheological
properties, though theimportant roleof theliquid
THE LIQUIDFILM HYPOTHESIS
lmingascell stabilisationandgasretentionshould
Recent studiesusingscanningelectronmicroscopy
notbeoverlooked. Thechangeinfreesurfaceenergy
(SEM) havehighlighted thesignicanceof a con-
has been calculated to account for about 80%
tinuouslamellar lmingascell stabilisationandgas
of the elastic energy involved in the deformation
retention
11,12
. It hasbeen shown that asubstantial
(expansion) of dough
29
. Although this calculation
number of gas cells are incompletely enclosed
has been disputed
13
, theconclusion remains that a
by a continuous starchprotein matrix in bread
greater pressurein gascellsisneeded to overcome
doughs at advanced stages of proving; rather,
surfacetensionthanviscousresistanceof thedough.
therearediscontinuitiesinthematrix, withholes
Nevertheless, thesignicanceingascell stabilisation
ostensibly forming inter-connections between ad-
oftheliquidlmrelativetothatofthestarchprotein
jacent cells(Fig. 2A)
11
. It isclear that thestructures
matrix during bread making may vary amongst
observed cannot bepresent as such in thedough
ours. A stableliquid lmcould compensatefor a
in situ since that would imply an open sponge
starchproteinmatrixofrelativelypoor extensibility,
typical of that of the bread crumb (Fig. 2B), and
asmayoccur whensurfactantsareaddedtoimprove
incapableof retaininggas. Hypothetically, thegas gasretention, or viceversa, asmayoccur whengluten
cells remain separated by a fragile liquid lamella is added to improve baking performance. If both
stretching across the holes (Fig. 3)
11,12
. This lm, phases are unsatisfactory, a dough with poor gas
stabilised bysurfaceactivematerials, issupported retention will result.
initially by a continuous starchprotein matrix, Cryo-stage SEM and SEM of doughs prepared
but that matrix fails to enclosethegas cells com- bygradual freezinghaveprovidedindirect evidence
pletely at advanced stages of expansion, leaving for a liquid lm partly separating contiguous gas
areas between themcontaining only the lamellar cells
11
. In particular, abrillar structurewasnoted,
lm
11,12
. which resembled hydrated protein in excesswater,
Thus, two consecutive stages are envisaged as and it was concluded that thepresenceof a liquid
occurring during dough expansion. Firstly, the lm between adjacent gas cells may be necessary
expandinggascellsareembeddedinacontinuous for theformation of theobserved brils
11
.
starchprotein matrix, which develops into thin, Furthermore, cuttinga dough rapidly acrossthe
continuous membranes between adjacent cells surface using a sharp razor blade at the end of
until its tensile stress increases to the point that proving(50min) had littleeect on its volume, i.e.
ruptureoccursor until thereissimplynot enough thedough did not collapseaswould beexpected if
material to maintain the continuity of the mem- gases, which support the structure of dough, are
branes. Thebehaviour of thedough prior to this retainedonlybyatopskin. Interestingly, however,
is determined primarily by the rheological prop- doughscollapserapidly, losinganestimated2530%
in height within 23s, when the tins containing erties of the bulk dough phase, particularly its
Z. Gan et al. 220
= Starch granul es
Li qui d l amel l ae
End of oven spri ng or baki ng
Earl y stages of fermentati on Advanced stages of fermentati on
to
earl y stages of baki ng
= Starchprotei n matri x
= Gas cel l l i ned wi th a l i qui d fi l m
Figure3 A revised model of dough expansion. Soon after mixing, thedough consists of discretegas cells lined with liquid
lms and embedded in a continuous starchprotein matrix. The matrix fails to enclose the gas cells completely at advanced
stages of fermentation, leaving areas that contain only a thin liquid lamella. Baking increases the rateof expansion until the
lamellar lmisincapableof meetingthedemandfor newsurfaceareageneration, thusconvertingthefoamstructureof dough
into an open sponge. The loss of gas retention is caused, therefore, by the rupture of the liquid lm, not that of the
starchprotein matrix.
themarestruck sharply on oneend (L. S. Besford, our, andstarchlysophospholipids, whichdonot
32
.
RHMResearchandEngineeringLtd, personal com- Using reconstitution methods, a general, al-
munication). Further mechanical shocksproduceno though unusual, relationship was found between
further reductions in dough height. The internal the content of the natural our lipid and loaf
structure of the dough, therefore, does not allow volume measured in a baking test, in which exo-
freemovement of gasover extended distances, but genous lipid was omitted fromthe formulation
33
.
the structure is mechanically fragile as for most Theloaf volumewas high for a our fromwhich
liquid foams. theNSL had been removed (Fig. 4). As this lipid
wasaddedback, volumedecreasedtoaminimum
at a lipid content intermediate between the de-
THE ROLE OF SURFACE ACTIVE LIPIDS
fatted and untreated ours. After this point, loaf
volume increased, approaching a constant value
The lipid reserves of wheat are triacylglycerols.
at lipid contents higher than that of the original
Small quantities of diacylglycerols, monoacyl-
our. TheNPoL fraction produced a progressive
glycerols and free fatty acids (FA) may be in-
decline in loaf volume, however, and the FA of
termediates in triacylglycerol biosynthesis
30
, but
the NPoL fraction have been shown to be those
they may also bedegradation products produced
which are mainly responsible for the depression
by lipase action
31
. These lipids are collectively
in loaf volume
34
. Loaf volumeswerealso found to
termednon-polar (apolar)lipids(NPoL), although,
behighly correlated with parameterssuch asPoL in fact, some are moderately polar. The other
content, PoL/ NPoL ratioandgalactolipidcontent lipids in wheat are structural lipids in various
of petroleumether-extracted (free) lipid, but not membranesand organelles, comprisingnumerous
with the water-saturated butanol-extracted (free glycolipidsand phospholipids, collectively termed
and bound) lipid
35,36
. Similar relationships have polar lipids (PoL). The endospermlipids are also
beenreportedbetweenloaf volumeandfreelipid
37
. convenientlydividedintonon-starch lipids(NSL),
which aect the baking performance of wheat Although theresults fromtheabovestudies have
Gas cell stabilisation and gas retention 221
Wheat lipids, which are found to be associated
closelywiththeliquidphaseofdough(about03%)
27,28
,
probably represent the most signicantly surface-
active components in wheat our dough. These
lipids, and particularly the polar fraction, are
thought to assist in the foamability of dough by
forming a lipid monolayer at the gas/ liquid in-
terface
10,40
. Becausedoughsprepared fromdefatted
ourscanexpand
33
, it must beassumedthat surface
activelipids arenot essential for gas retention and
that proteins and/ or pentosans dissolved in the
dough aqueous phase can take the place of lipids
in defatted our. Thereisno doubt, however, that
endogenous PoL lipids (e.g. phospho- and galacto-
1000
200
Added l i pi d (mg)
L
o
a
f

v
o
l
u
m
e

(
m
l
)
800
180
160
140
0 200 400 600
NPoL
PoL + NPoL
PoL
lipids), aswell assurfaceactivesyntheticemulsiers,
such asDATA esters, improvegasretention.
Figure 4 Loaf volume (ml) as a function of lipid content.
Theresponsesto small changesin lipid content, PoL, polar lipids; NPoL, non-polar lipids. Dashed line rep-
resentsvolumeat end of proving. Additionsweremadeto a
which do not aect the rheological properties of
dry our weight of 302g(reproduced with permission from
dough, can be rationalised on the basis that the
MacRitchieand Gras
33
).
lipid exerts its eects at the gas/ liquid interface,
thereby inuencing the formation and stability of
thegascells. Wheat lipidsarecapableof orienting
all agreed in showing the PoL fraction to be themselves at the gas/ liquid interface to form a
favourable and the NPoL fraction to be detri- lipidmonolayer. Thespreadingpressureof thelm
mental inbreadmaking, anumber of experiments provides a force that counteracts the interfacial
havebeenunsuccessful inestablishingcorrelationsbe- tensionbetweenthegasandtheliquidphases, and,
tweenloaf volumeandfreelipid
38,39
. thus, stabilises the gas cells. Unlike the hexagonal
A characteristic feature of lipids is their poly- phase, which behaves like a plastic fat and is not
morphicbehaviour. PoL mayformvarioustypesof spontaneously dispersible, the LCL phase spon-
phasewhenwheat our ishydratedtoformdough, taneouslyformssmall, bilayer aggregates(liposomes)
dependingonfactorssuchastheproportionsof the in an aqueous environment on mixing. The rapid
PoL and water and thestructureof thePoL class. formation of condensed lipid monolayers at in-
The phase relationships, corresponding to the terfaces, suchasgas/ liquidinterfacesinwheatbread
thermodynamic equilibrium between various pro- doughs during fermentation and the early stages
portions of PoL and NPoL in water, are usually of baking, requires lipid depots to be available
describedbyso-calledternaryphasediagrams
40
. The throughout theprocessinthedoughstructure. The
lamellar, liquidcrystalline(LCL) phaseconsistsof
LCL phase in the formof liposomes is ideal for
lipid bilayers with the hydrocarbon chains in a
thispurpose. A liposomal dispersion of lecithin, for
liquid-likedisorder formingthecoreand thepolar
example, when added (about 2%, w/ w) to a 1:1
head groupsformingthetwo outsidesurfaces. PoL
mixtureof a wheat and riceour, produced bread
in theLCL formcan transformspontaneously in
of the same volume as that produced when wheat
an aqueous environment on mixing into lamellar
our alonewasused
40
. A lecithin sludge obtained
dispersions, comprising small, bilayer aggregates
asaby-product of soyaoil reningproducedsimilar
(liposomes). When such a transformation occurs,
resultsin a wheat/ sorghumour blend
42
.
virtually all PoL and most NPoL are likely to be
detectedintheboundform
41
. Inafermentingdough
the PoL may provide surface-active molecules at
THE ROLE OF SURFACE ACTIVE PROTEINS
thegas/ liquidinterfacetostabilisethedispersedgas
Proteins, being amphipathic in nature, may also phase
10
. The lack of appreciation of the dierent
exert a surface eect at air/ water interfaces in physical states of lipids may well have contributed
doughbyformingacontinuouslm. Proteinsdier totheoftencontradictoryconclusionsthatarefound
substantially in their surface activities, and these in the literature regarding the eects of lipids on
bakingperformance. dierencesarenotattributablesimplytounspecic
Z. Gan et al. 222
variation in amphipathicity, since most proteins process of transport and adsorption of proteins
exhibit similar distributions of hydrophobic and fromthe bulk phase, they may have limited sig-
hydrophilicresidues
43
. Thevariationisbelievedto nicance in terms of understanding the mech-
arisefromdierencesinconformational structure, anisms and molecular processes involved in gas
stability and/ or exibility of the conformation, cell stabilisation in wheat bread dough. Many
symmetricor asymmetricdistributionof polar and
functional properties of proteins, including their
apolar patchesonthesurfaceof themolecule, and
ability to formgelsand to stabiliseemulsionsand
in molecular sizeand shape
43
.
foams, dependontheir solubility
52
. Glutenproteins
Alternative pathways for folding protein
are well known for their insolubility in salt so-
moleculestominimisefreeenergybecomepossible
lutions, and yet the calculated salt concentration
when proteins are adsorbed at an air/ water in-
in the aqueous phase of a bread dough is about
terface. In addition to foldinginto theinterior of
05x, assumingthat all thesalt added asadough
aglobuleintheaqueousphase, apolar sidechains
ingredient remains in the aqueous phase. How
can be located in the air phase
44
. The rate of
the gluten proteins might be transported to and
adsorption is diusion-controlled in the initial
adsorbed into the interface in dough under such
stages of adsorption (i.e. at lowsurfacecoverage),
conditions is dicult to envisage. Moreover, im-
whereas there is an energy barrier to adsorption
munolocationstudieshavefailedtoshowthepres-
at high surface coverages
45
. Although the exact
ence of gliadin proteins at gas cell surfaces in
natureof theenergybarrier isnotfullyunderstood,
bread, althoughtheseproteinswerereadilylocated
it may be related to A, where is the surface
at thecut surfaces of thebulk bread crumb
53
.
pressure and A is the molecular area required to
Proteinsrepresent about 23%byweight of the
be cleared for the molecule to adsorb at the
liquid phase of dough
27,28
. If they do have a role
surface
45,46
. Thebasicpremiseisthat, for aprotein
to play in interfacial lm stabilisation, it is the
molecule to clear and occupy an area A at the
water andsalt extractablefractions, whichaccount
surfaceagainst thesurfacepressure, it shouldpos-
fof 1520%of total our protein, that seemlikely
sess a molecular energy equal to or greater than
to full thisrole. Theseproteinshavebeen shown
A. Theenergybarrier mayactuallyberelatedto
to be benecial in bread making
5456
. The ad-
surfacedenaturationoftheprotein, whichinturnis
sorptionof suchproteinsatthegas/ liquidinterface
dependent onsurfacewater activity
47
. Thesurface
could besignicant in this respect.
denaturation of proteins is thought to be caused
Also of interest in relation to the possible role
by the reaction of high-energy surface water
of protein in stabilising interfacial lms in bread
molecules with the internal polar groups of the
doughs is the recent discovery of an amphiphilic
proteins, which facilitates their unfolding at the
protein, puroindoline, in wheat our
57,58
. Puro-
surface(Fig. 5).
indolinewasisolatedfromthedetergent richphase
In addition to loweringinterfacial tension, pro-
of a 4% (w/ v) Triton X-114 extract of our after
tein molecules can forma continuous lmat the
phase partitioning by condensation of the de-
interfaceviacomplex intermolecular interactions.
tergent at 35C. Friabilin, a protein identied
These are thus able to impart structural rigidity
previously as a starch granule surface associated
totheinterface
48
. Thedevelopment of suchmech-
protein
59
, is also extracted from our by Triton
anical strengthmaynot bepossibleinthecaseof a
X-114 and partitionsin thedetergent rich phase;
simplesurfactant lm, inwhichtheintermolecular
it shows extensive homology with puroindoline
57,58,60
interactions arerelatively weak.
(also D. Marion, INRA, Nantes, France and
Much of the earlier work on the surface be-
P. Greenwell, FMBRA, Chorleywood, UK, un-
haviour of wheatproteinswaslimitedtoexamining
published results). The phase partitioning be-
eectsof proteinmonolayersspreadat thesurfaces
haviour of puroindoline suggests that it may have
of aqueous media
4951
. Both gliadin and glutenin
strong polar lipid binding properties and that its
fractionshavebeen shown to besurfaceactiveas
amphiphilic nature may make it strongly surface
determined by surface balance techniques. The
active. Indeed, evidence has been presented that
gliadin fraction spreads more quickly at the gas/
puroindoline is surface active and has the ability liqud interface, achieving much higher surface
to stabilise interfacial lms/ foams
61
. Dening the pressure per unit area concentration than the
role of puroindoline and related proteins in gas glutenin fraction
51
.
Sincesuchstudiesdonot represent thedynamic retention will bean important goal in thefuture.
Gas cell stabilisation and gas retention 223
= Hi gh energy water mol ecul es
Gas
= Low energy water mol ecul es
Li qui d
Trai n
Loop
Tai l
(a)
(c)
(b)
Gas
Li qui d
Figure5 Schematic representation of protein adsorption at a gas/ liquid interfacein dough. (a) nativeprotein immersed in
a liquid phase; (b) protein in contact with high energy water moleculesat theinterface; (c) adsorbed, unfolded, and hydrated
protein moleculeswith trainsin direct contact with theinterface, loopsbetween thetrainsprojectinginto thetwo bulk phases,
and tails at the N- and C-terminal ends of the polypeptide. The tails are expected to be present in the aqueous phase since
the N- and C-terminals of a polypeptide are charged at around neutral pH (adapted and reproduced with permission from
Ter-Minassian-Saraga
47
).
terfaces (Fig. 6). Interesting, puroindoline, which
FILM STABILITY WITH CO-EXISTING LIPIDS,
is thought to be a lipid binding protein, can act
PROTEINS
cooperatively with polar lipids in interfacial lms
Although the mechanical stability of a lm with
under some circumstances resulting in increased
two or more components is believed to be some-
foamstability
61
.
what lower than for any of the highly condensed
states of the single components, especially if the
individual component molecules do not interact
THE ROLE OF PENTOSANS
with each other
48
, lipidsand proteinscan co-exist
in interfacial lms and aregenerally envisaged as Pentosans are the major non-starch poly-
saccharides of wheat our. They originate from actinginacompetitivemanner. Thismayexplain
the unusual relationship observed between PoL the endosperm cell walls of wheat grains, and
comprise mainly arabinoxylans and arbino- content and loaf volumewhen PoL isadded back
incrementallyto defatted our (Fig. 4)
33
. Gascells galactans, whicharepartlyextractablewithwater.
Thehigh water-bindingcapacityof pentosans
62,63
, are probably stabilised by surface active proteins
when NSL have been removed, resulting in the together withtheir highsolutionviscosities
6466
and
their ability to undergo oxidativegelation
67
, have initial high volume. As PoL is added back, the
protein lms are destabilised, reaching minimum been considered potentially important in in-
uencing our water absorption, dough mixing stability at approximately 100mg PoL, probably
by formation of a mixed lm, with reduced in- and dough rheological properties
62,63,66,6870
.
Despite considerable research, however, the termolecular interactions of proteins at the in-
terface. Further additions of PoL may result in functional role of pentosans in bread making is
still not well dened. Thisisduelargely to a lack theformation of PoL-dominated lmsby gradual
displacement of proteins from the gas/ liquid in- of insightuntil relativelyrecentlyintothestructural
Z. Gan et al. 224
= Protei n
0
Added pol ar l i pi d (mg)
(c)
= PoL
= Protei n-PoL
compl ex
(a)
(b)
400
L
o
a
f

v
o
l
u
m
e

(
m
l
)
aqueous phase
Figure 6 Speculative mechanisms for liquid lm stabilisation as PoL is added incrementally to defatted our. (a) lm
stabilisation by surface active proteins, resulting in the initial high loaf volume when NSL have been removed; (b) as PoL is
added, theprotein lmsaredestabilised, probably by formation of a mixed lm, with reduced intermolecular interactions; (c)
further additions of PoL gradually displace the proteins fromthe gas/ liquid interfaces, forming a stable, single component
PoL lm.
properties and heterogeneity of arabinoxylans, ilarly, whereasdetrimental eectsonbreadmaking
performance of adding water-unextractable ara- thuslimitingthedesignof meaningful experiments
and the interpretation of experimental results. binoxylans were noted by some
69,78
, others ob-
served positiveeects
79,80
. It isdicult to besure, Studiesto examinetheroleof pentosansin bread
making have been of three types in the main however, that thefunctional eect of acomponent
added to adough will bethesameasthat present reconstitution, correlation, and enzymic modi-
cation andtheresultsof suchstudieshavebeen endogenouslyinaour. Indeed, dierent methods
of additionof pentosanstodoughhavebeenshown to someextent contradictory.
Benecial eectsonloaf volumeof addingwheat to producedierent functional eects
81
.
Correlation studies showed a positive re- and/ or ryewater-extractablepentosanshavebeen
observed in a number of studies
68,7175
using re- lationshipbetweenthewater-extractablepentosan
content of 58 hard wheats and bread textural constitution methods. However, other groupshad
observed that addition of extra pentosan did not characteristics
82
. On theother hand, a slight neg-
ativeeect of water-extractablepentosanson loaf increase loaf volume
76,77
, although water-ex-
tractable pentosans were found to be responsible volumewasalso observed
83
. Analysisof six Euro-
pean wheat ours with varying bread making for the reduction in loaf volume when water-
solubles were removed from wheat our
76
. Sim- potentials indicated an inverse relationship be-
Gas cell stabilisation and gas retention 225
tween thehandlingpropertiesof adough and the viscositylmsarelessstable. It islikely, therefore,
that the increase in the amount of high M
r
ara- our water-extractable pentosan content
84
. Pre-
dictivemodellingstudies of thebakingproperties binoxylan, and, consequently, an increase in the
viscosity of the dough aqueous phase as a result of wheat
85
showed that inclusion of non-starch
polysaccharide variables helped to explain some of the limited solubilisation of arabioxylan from
water-unextractable pentosans at the optimum of the variation in dough rheological properties
and contributed signicantly to variation in our level of enzyme addition
90
, improved the foam
stability of the dough. This is consistent with the water absorption, but did not explain variation in
loaf volume. ndingthat arabinoxylanof highintrinsicviscosity
weremoreeectivein stabilisingan aqueouspro- Although earlier studies with puried en-
doxylanaseenzymesshowedadverseeectsonloaf tein foam during thermal expansion
91
. The ad-
dition of arabinoxylan wasalso shown to produce properties
70
, more recent work has shown that
substantial improvements in bread making per- a substantial reduction in the surface tension of
water, and that, at a concentration of 05%, all formance can be produced by such enzymes
86,87
.
The use of an endoxylanase preparation in a pentosan preparations fromeight wheat cultivars
weresurfaceactive
91
. Thus, apart fromplaying a Chorleywood Bread Process (CBP) formulation
resulted in prolonged expansion of dough during positiverolein dough rheology, water-extractable
the early stages of baking
87
indicating that gas pentosans may contribute to the stabilisation of
retention bythedough wasimproved. Themech- gascellsby improvingthemechanical strength of
anism of this eect has not been dened, but theliquid lms in thedough. Overall, thestudies
analysis of fractionated polysaccharides from summarised above indicate a positive eect of
doughs at dierent stages of mixing and proof water-extractablearabinoxylanson gasretention.
indicated that the enzyme caused a rapid and Firm conclusions about the functional role(s) of
specic degradation of water-unextractable ara- pentosans in bread making will only be possible
binoxylans and thus a substantial increase in the when the structure and properties of wheat our
water-extractablefraction
88
. Kulp
89
hadalsoshown arabinoxylansandtheir variation, especiallythose
earlier that thebread makingpropertiesof water- of the water-extractable fraction, are more fully
unextractable pentosans were improved by di- understood.
gestion with a crudefungal pentosanase.
The release of arabinoxylan from water-un-
extractable pentosans by a crude pentosanase-
THE INVOLVEMENTOF BAKERY FAT
containingenzymepreparationincreasedthespe-
It is outside the scope of this review to consider
cic viscosities of aqueous extracts of dough,
indetail theroleof addedbakeryfat or shortening though the increase was not proportional to the
in gas retention in bread doughs. Nevertheless, extent of solubilisation of the unextractable pen-
some mention should be made of the possible tosans
90
. An improvement in dough quality was
involvement at thegas/ liquidinterfaceof fat crys- evident when a greater viscosity was achieved at
tals and their inuenceon gas retention. theoptimumlevel of enzymeaddition, whereasat
Fat is an optional ingredient for traditional excessive enzyme levels, viscosity decreased and
longor bulkfermentationbreadmakingprocesses. doughqualitydeteriorated
90
. Thereisaclear need
For notime mechanical doughdevelopment pro- to dene the mechanism of bread improvement
cess, suchastheCBP, however, it isanobligatory by xylanases more precisely, and this may throw
ingredient for production of bread of acceptable further light on the roleof pentosans themselves.
quality. Furthermore, not only must fat be in- Aqueous solutions of water-extractablepentosans
cluded, but a proportion (5%) of that fat must be are highly viscous and it is possible for such
solid(i.e. incrystallineform) at proof temperature. solutions to be converted into gels via oxidative
Fat crystals per seare not surface active, and covalent cross-linking
67,91
; the arabinoxylan frac-
therehasbeen no direct demonstration to dateof tion has been identied as the major component
the involvement at the gas/ liquid interface of fat responsiblefor thehigh viscosity
91
.
crystals and their inuence on gas retention in The stability of a lamellar lm, and thus the
bread doughs. Theassociation of fat crystalswith stabilityof afoamitself, dependsonseveral factors,
the surfaces of gas cells in cake batters has been such as lmviscosity, shear resistance and elast-
icity. Lowelasticity, lowshear resistanceand low described recently, however, and isthought to be
Z. Gan et al. 226
a secondary process
92
. Initially, the gas cells in ner textured loaf produced by using improvers,
emulsiers and added wheat gluten. The non- cakebattersarestabilised by an interfacial lmof
egg proteins. Fat globules containing both liquid endospermcomponentsoccurringnaturallyinthe
wheat caryopsis are known to be responsible for fat and fat crystals are also surrounded by a
proteinaceous interfacial lm, and, indeed, ejec- producing the low specic volume and dense
crumb structure of the traditional wholemeal tion of fat crystals from liquid fat may be aided
by thissurroundinglm. Asmixingproceeds, gas bread
32,94
. This eect cannot be explained solely
by these components diluting the gluten-forming cells and fat crystals come into contact, and the
proteinlayersfusesoastoformasinglecontinuous proteins.
The interactions of white our fractions with lm surrounding both the gas cells and the fat
crystals. It hasbeenshownthat fat crystalsbecome thedegradationproductsof thelipidsof germand
bran contribute to the depression in loaf volume aligned tangentially at thegascell surfacein cake
battersandthat thisimprovesgasretention
92,93
. In of bread made from stored wholemeal our
95,96
.
Lipid hydrolysis, due mainly to enzymic activity general, small fat crystals of the polymorph
haveagreater abilitytostabilisegascellsthanthe in the bran, liberates free fatty acid, the poly-
unsaturated fraction (PUFA) of which (ca. 60% much larger crystals.
It has been speculated that similar events may of the total) is oxidised by lipoxygenase activity
concentratedmainlyinthegerm
95
. Thisoxidation, occur in bread doughs containing fat and may
help to explain the role of fat in improving gas which occurs when the dough is formed as well
as in the our itself, competes with the yeast for retention in bread doughs
92
. Such fat crystals are
also thought to be important in another respect, availabledissolved oxygen in thedough and with
other oxygen-requiringreactions, such as theox- i.e. in gas retention during the baking stage. As
thetemperatureof thedough or batter increases, idation of ascorbic acid, which is a necessary
rst step in its bread improving reaction. This is the fat crystals melt and the liquid oil then ows
over the inner surface of the gas cells to forma particularlyimportantinwholemeal breaddoughs,
in which ascorbic acid is added as thesolebread hybrid interface comprising the oil layer in ad-
dition to the protein and/ or polar lipid layer
92
. improver incountriessuchastheU.K.
94
. Thefree
fatty acids and other lipid degradation products The layer of oil helps to maintain the continuity
of thegas/ liquidinterfaceintheexpandingdough mayalsoact asfoamdestabilisersanddepressloaf
volume. Theeectof enzymicdegradationof lipids piece and thus aids gas retention during oven
spring. A further advantage of the fat crystal onbakingperformanceismuchmoresignicant in
wholemeal than whiteoursbecauseof themuch polymorph over the polymorph in this respect
may bethat theformer has a lower meltingtem- greater rate at which it occurs in wholemeal
ours
96
. perature and thus may be more readily available
for contributingtogascell surfaceintegrityduring The addition to wheat our of small amounts
(12%) of pearlings, obtained by dry abrasion bakingthan thelatter.
In addition to its eect on the rheological be- (pearling) of theouter pericarplayersof thewheat
caryopsis, hasbeenshowntohaveamarkeddetri- haviour of the bulk dough phase, the possibility
mental eect on loaf volume
97
. Since lipase is that bakeryfat/ shorteningalso participatesin gas
concentrated in the bran fraction
98
, it is possible retentionthrougheventsatthegas/ liquidinterface
that the eect of pearlings on loaf volume could addsanother level of complexity. It will beneces-
bedue, inpart, toincreasedlevelsof FAcontaining sary in futurework to determinetherelativecon-
PUFA. ThedierencesinFA levelsbetweenwhite tributions of the dierent factors that have been
our, wholemeal andour frompearledgrainare proposed as being of importance to gas cell sta-
small
99
, however, and are unlikely to account for bility, gas retention and nal bread quality.
theobserved dierences in bakingperformance.
Moreover, heattreatment(autoclaving)doesnot
alleviate the adverse eect of theoutermost bran
GAS RETENTION IN WHOLEMEALBREAD
fraction on loaf volume
99
, indicating that the ob-
DOUGHS
served eect is due largely to a physical rather
Wholemeal bread is produced in various forms, than a biochemical mechanism. In other words,
ranging from products with low specic volume the result is inconsistent with the possibility that
heat-sensitive components (e.g. enzymes) are re- andadensecrumbtexturetoamoreexpandedand
Gas cell stabilisation and gas retention 227
sponsiblefor theadverseeect of thisfraction on expansion, theprecisearchitectureof thegascells
is still incompletely understood. Nevertheless, loaf volume. It is quite plausible that particulate
components, of thekind that havebeen identied recent studies have provided accumulating evid-
encefor theinvolvement of interfacial phenomena inwholemeal bread, couldcreateareasof weakness
in an expanding wholemeal dough
97,99
. SEM mi- in gas cell stabilisation and gas retention. Hypo-
thetically, the dough consists of discrete gas cells crographs revealed that the bran materials are
incorporated into the gas cell walls of doughs lined with liquid lms stabilised by surfaceactive
materials and embedded in a continuous starch produced fromoursof both control and pearled
wheat
97,99
. These materials, notably the epicarp protein matrix soon after mixing. As the volume
fraction of the gas increases, the expanding gas hairs, werefoundtoimpedethenormal formation
anddevelopment of thegascell structureof whole- cells become polyhedral, with discontinuities de-
velopingin thematrix, leavingareasthat contain meal dough, and perhaps more signicantly, to
restrict andforcegascellstoexpandinaparticular only a thin, lamellar lm between neighbouring
cells. The degree to which such discontinuities dimension. Thisdistortsgreatly thegascell struc-
ture, and may contribute to the resultant crumb occur is dependent on the extensibility of the
starchproteinmatrix, which, inturn, isdependent morphology (e.g. the size, the shape and the dis-
tribution of the gas cells), which is an important largely on the gluten proteins. The surface area
of thelamellar lmincreases as discontinuities in element of crumb texture
99
.
the starchprotein matrix develop further during The outer branny materials may also provide
the second stage of expansion. Failure of the anextremelyeectivecapillarysystem, whichmay
lamellar lmto maintain the rate of newsurface removewater fromthedough aqueousphaseand
area generation towards the end of oven spring thusreduceitsavailabilityfor formationof lamellar
leadstotheruptureof thislmand, consequently, lms. Thiswasindicated in an experiment where
thelossof gas. Thus, theend of dough expansion 2%additional water wasfoundtohaveabenecial
ismarked bytheruptureof thelm, and not that eect on the volume of loaves made fromwhite
of the starchprotein matrix. Additional factors, our to which bran had been added
100
. Further
such asthepresenceof theouter brannylayersof additionof water producedamarkedimprovement
the grain, which may alter the continuity and only in the presence of sodiumstearoyl lactylate
rheological propertiesof thestarchproteinmatrix (SSL); loaves with 6% additional water and 2%
and the availability of water for lamellar lm SSL had volumes almost equal to that of the
formation, are involved in gas cell stabilisation control without bran
100
. The latter observation
and gas retention in wholemeal doughs. may be explained by the combined contribution
Further research is needed to determine the of water and surfactants to the formation and
validityof theliquid lmhypothesisand to dene stabilisation of lamellar lms and thus to gas cell
the molecular species involved in surface lm stabilisation and gas retention.
stabilisationinwheatbreaddoughs. Theindividual Thetechnological signicanceof glutenproteins
and joint contributions of the starchprotein
in bread makingdependson their contribution to
matrix and of theliquid lmto gasretention also
theextensibility and cohesivestrength of thebulk
need to beclaried. Oncethespecic rolesof the
dough phase. The addition of vital wheat gluten
starchprotein matrix and the lamellar lms are
has benecial eects on the microstructural in-
understood, dierencesbetweencultivarscouldbe
tegrity of wholemeal bread
97
. The gas cell walls
investigatedinorder toimproveecienciesinthe
appear to be strengthened by the extra gluten
use of materials and ingredients, and to facilitate
in the starchprotein matrix. This may result in
greater control over processingperformancesand
improved mechanical properties of the starch
product quality.
protein matrix so that itsruptureisdelayed, lead-
ingto improved gas retention.
Acknowledgements
The research described here that was carried out in
CONCLUSIONS
our laboratory was funded in part by a research stu-
Although eortshavebeen madeover decadesto
dentship to ZG fromRHM Research and Engineering
investigate the mechanism(s) of gas retention in
Ltd, TheLordRankCentre, HighWycombe, UK and
by an Overseas Research Student Award also to ZG dough and the events that occur during dough
Z. Gan et al. 228
time, temperature, and water absorption. RheologicaActa
fromtheCommitteeof Vice-ChancellorsandPrincipals
9(1970) 223238.
of UK Universities, to both of whom thanks are ex-
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pressed. Financial support fromtheBiotechnologyand
J.B.M. A rheological investigation of theroleof water in
Biological Sciences Research Council, formerly the
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