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Meat Science 15 (1985) 15-30

The Relationship Between Water-Holding Capacity and


Cooked Meat Tenderness in Some Beef Muscles as
Influenced by Acidic Conditions Below the Ultimate pH

N. F. S. Gault

Department of Agricultural and Food Chemistry, The Queen's University of Belfast,


and Department of Agriculture for Northern Ireland, Newforge Lane,
Belfast BT9 5PX, Northern Ireland

(Received: 15 November, 1984)

SUMMARY

Previous studies have indicated the beneficial influence of higher than


normal ultimate p H (pHu) on the tenderness o[' cooked meat. Such
be.nefits have been indirectly linked to the influence of increasedpH on the
water-holding capacity (WHC) of meat above the iso-electric point (1EP)
of the myofibrillar proteins. In the present study, relationships between
W H C and the tenderness of some beef muscles were investigated under
p H conditions withil; and below the IEP of the myofibrillar proteins, such
that the maximum range of meat swelling was achieved.
It was found that increased WHC, as measured by swelling ratio in
both raw and cooked meat, markedly influenced cooked meat tenderness,
irrespective of the connective tissue content of the muscles. The results
fit~'ed a series of exponential decay equations relating swelling ratio to
cooked meat toughness. Additionally, strong linear decreases in
toughness were apparent over the p H range 4.6 to 4 l for the three muscle"
types studied.

INTRODUCTION

The pioneering work of Hamm and his colleagues in the 1950s (see Hamm,
1960) provided the basis of our present knowledge of the physico-
chemical properties of meat which influence its ability to retain water, be
15
Meat Science 0309-1740/85/$03.30 ~" Elsevier Applied Science Publishers Ltd, England,
1985. Printed in Great Britain
16 N. F. S. Gault

it endogenous or added during processing. In this work, pH was clearly


identified as a major factor influencing the water-holding capacity (WHC)
of homogenised meat systems, a minimal effect occurring at the iso-electric
point (IEP) of the major proteins in muscle (pH 5.0 to 5.5) and increasing
markedly with changing pH on either side of the IEP. Penny et al. (1963)
later demonstrated a similar phenomenon with purified beef myofibrils
over the pH range 4.5 to 7.5, which they attributed to changes in muscle
fibre diameter. Offer & Trinick (1983) have since observed an increased
swelling of individual myofibrils over the pH range 5.0 to 9.0 as part of a
more extensive study showing the beneficial influence of salt and
pyrophosphate solutions on myofibrillar swelling at the IEP. These
observations not only substantiated the earlier work of H a m m (1960) and
Penny et al. (1963) but prompted the first structural hypothesis of water-
holding in meat based on the swelling or shrinkage of myofibrils caused
by expansion or shrinkage of the myofilament lattices.
In the context of meat tenderness, however, the r61e of WHC has
received relatively little attention. Nevertheless, the importance of
intrafibre water to meat tenderness has recently been emphasised by
Currie & Wolfe (1980) who correlated changes in the mechanical
properties of uncooked beef muscle undergoing rigor mortis with changes
in the WHC of the muscle fibres as influenced by pH and the rate of fall of
pH. However, Miles & Lawrie (1970) were perhaps the first to identify a
relationship between pH and meat tenderness using cooked rabbit
muscle, tenderness increasing linearly with increasing pH over the range
5.5 to 7.1. More extensive studies on mutton (Bouton et al., 1971,
1972a, b) and beef (Bouton et al., t973a, b) not only confirmed this pH
effect on cooked meat tenderness, but also identified a linear increase in
WHC over the same pH range. In other studies, where deliberate
interference with post-mortem glycolysis was not utilized as a means of
varying ultimate pH (pH ~), meat of naturally high pHu has indirectly been
shown to be consistently more tender after cooking than normal pH u
meat (Khan & Lentz, 1973; Fredeen et al., 1974: Dransfield, 1977).
In the present study, relationships between WHC and cooked meat
tenderness in certain post-rigor beef muscles have been investigated under
acidic conditions below the pHu. The results obtained during this study
indicate that the swelling characteristics of intact beef muscle, as
influenced by muscle pH, bear a marked similarity to the influence of
acidic pH values on the WHC of homogenised beef as found by Hamm
(1960). Furthermore, the results obtained from the present study have
WHC and tenderness of beef muscles below the ultimate pH 17

been shown to fit a series of exponential decay equations which quantify


the ~'elationship between WHC and cooked meat toughness. The
importance of W H C to meat tenderness is discussed in the light of these
findings.

MATERIALS AND METHODS

Preliminary treatment of muscles

The muscles chosen for this study were the M. longissimus lumborum
(LD), M. triceps brachii caput longum (TB) and the M. injhaspinatus (IS).
Thes,e muscles were selected on the basis of their collagen content and
their relative size and shape which facilitated the sampling procedures
used in this work. The LD and TB muscles have a relatively low and
similar collagen content (2'3~o dry weight) while the IS has an
exceptionally high collagen content (13.0 ~o dry weight) with characteris-
tic sheets of connective tissues running throughout its length (Bendall,
1967). Four of each of these muscles were individually taken from the
carcases of twelve young steers of average slaughter weight and age.
Muscles were obtained, when required, three days after slaughter from
the same commercial source. Each muscle was cut transversely to the
muscle fibre direction on a gravity feed slicer into steaks approximately
1.0cm thick, from which discs of meat 3.0cm in diameter (wt 7.5 _+ 1.0g)
were prepared using a cork borer. To facilitate slicing and the preparation
of discs, transverse blocks of muscle and steaks were tempered
indivJLdually at - 2 5 °C until the surfaces were rigid.

Adjustment of pH within muscle samples


Acetic acid solutions of the following molarities were used to adjust the
pH of the meat discs to cover the range pH 5.2 to 3.0:* 0-0025,+ 0.005,+
0.01,0.025, 0-05, 0.075, 0.10, 0.25, 0.50, 0.75, 1.00, 1.25,++ 1.50, + 2.00 and
3-00.
* The results obtained from the LD experiments showed that pH's close to the IE P could
not be obtained with 0.01macetic acid solution, the weakest used. In subsequent work on
the TB and IS muscles, therefore, weaker acetic acid solutions were used to achieve the
desired pH effect.
t TB and IS only.
++ LD only.
18 N. F. S. Gault

Each meat disc was accurately weighed and plgced in a 200 ml capacity
screw-cap polystyrene jar to which 50 ml chilled (~ °C) acetic acid solution
was added. Penetration of acid into the meat was aided by continuous
swirling at 120 rpm in a Gallenkamp Cooled Orbital Incubator for 48 h at
4°C. Untreated meat discs were used as controls. Each treatment was
replicated four times for each individual muscle used.

Swelling ratio and tenderness measurements

After equilibration, meat discs were surface dried with paper towelling
and reweighed. Two discs from each treatment were individually cooked
in sealed polythene bags in a water bath at 80°C for 20min and
immediately chilled in an ice-water bath. They were then surface dried as
before and reweighed. The weight of each meat disc, before and after
cooking, was divided by the corresponding raw meat weight to express
change in weight as swelling ratios which served as measures of WHC in
the raw and cooked states.
Cores of meat, 0-9 cm in diameter, were cut from each meat disc parallel
with the muscle fibre direction (where possible) for tenderness assessment
on an Instron Model 1122 Universal Testing Instrument fitted with a
Warner-Bratzler shear device. The average tenderness value for each meat
disc was calculated as the mean of either four (control samples and
those equilibrated in < 0"05M acetic acid) or six (samples equilibrated in
>0"075M acetic acid) measurements of peak shear force.

Chemical analyses

The remaining two meat discs from each treatment were individually
homogenised and made up to 100 ml with distilled water. Measurements
of pH were carried out on these solutions using a digital pH meter fitted
with a combination glass electrode. Titratable acidity was determined on
a 25-ml aliquot of each solution against 001M or 0' 10M NaOH using a
phenolphthalein indicator. Total nitrogen was determined on a 10-ml
aliquot of each solution by the Kjeldahl method (AOAC, 1980) modified
by incorporating a selenium, rather than mercury-based, catalyst. All the
used acetic acid solutions were assessed for total nitrogen (25-ml aliquot),
titratable acidity (5-ml aliquot) and pH by the methods described above.
Titratable acidity values were used to calculate the quantity of acetic
acid (g) in the various equilibrating solutions and meat discs. Values
WHC and tenderness of beef muscles below the ultimate pH 19

obtained from the nitrogen analyses were used to calculate the per cent
total nitrogen content of the raw meat discs leached into the equilibrating
solutions.

Statistical analyses

Analyses of variance were computed to examine the differences between


equilibrating treatments for each parameter measured within each muscle
type studied. Results are expressed as mean values with the effective
standard error of the means either included in the Tables or, for clarity of
presentation, in the legends to Figures.
Relationships between swelling ratios were examined by regression
analysis. Relationships between peak shear force values and swelling
ratios were also examined by regression analysis as outlined in the
'Results' section.

RESULTS

Efficiency of acetic acid penetration into the meat discs

From preliminary studies carried out with a range of acetic acid solutions
of varying strength (N. F. S. Gault, unpublished data), it was found that
optimum swelling of meat discs had occurred well within a 48-h period
under the conditions described in the 'Materials and Methods' section.
Consequently, this time was considered adequate to ensure that diffusion
of acetic acid into the meat discs had reached a state of equilibrium with
the ex~Lernal solution.
Table 1 shows that there was an expected decrease in the pH of the acid-
treated meat discs, the extent of which was related to the acetic acid
concentration of the equilibrating solutions. However, the results
obtained with the weakest acetic acid solutions were peculiar in that their
corresponding titratable acidity values were less than those of the
untreated controls of normal pH, (Table 1). This was particularly
noticeable with those discs equilibrated in < 0.025M acetic acid solution,
the effect becoming obscure at acetic acid concentrations > 0.05~. These
effects were markedly consistent, not only within each muscle group, but
also between the three muscle types studied, and highlight the complex
nature and uniqueness of each equilibrating treatment.
TABLE 1
Influence o f Acetic Acid C o n c e n t r a t i o n o n its D i s t r i b u t i o n * Between M e a t a n d E q u i l i b r a t i n g S o l u t i o n , a n d o n their p H tO
Characteristics (Acetic Acid C o n c e n t r a t i o n (M) o f F r e s h E q u i l i b r a t i n g S o l u t i o n s ( 8 t o r 165 o b s e r v a t i o n s per m e a n ) )

Musch, Control 0.0025 0.005 0.01 0.025 0.05 0.075 0.10 0.25 (I.50 0.75 1.00 1.25 1.50 2.00 3.00 SEM

{TA ~ 0.03 0.08 0.15 0-23 0.30 0.75 1.50 2.25 3.00 3.75 4.50 6-00 9-00 0.000
:[:TAu 0-06 0.08 0.14 0.20 0.25 0-57 1 . 1 2 1-66 2 . 2 1 2.79 3.36 4.45 6-81 0.035
t T A m 0.06 -- 0.04 0.05 0.07 0.09 fill 0.22 0.39 0.60 0.81 0.97 1.12 1.45 1:97 0.029
%A ~ 0-06 0.15 0-30 0.45 0-60 1 . 5 0 3.00 4.50 6.00 7.50 9.00 12.00 18-00 0.000
LD~
%A -- 0.11 0.17 0.29 0.42 0.55 1.30 2.62 3.89 5.24 6.57 7-89 10.40 15.60 0-055
t%A m 0.81 0.48 0.53 0.67 0.77 0.92 1.62 2-60 3-99 5-11 6.30 7.48 9.82 14.10 0.085
}pH ~ -- 4-41 4.42 3-99 3.90 3.80 3.53 3.29 3.16 3.06 2-97 2.91 2.83 2 - 7 1 0-017
tpH m 5.33 -- - 4.61 4.43 4.31 4.19 4.12 3-84 3.65 3.50 3.44 3.34 3.29 3.22 3.15 0.017
3~TAI -- 0-01 0-02 0.03 0.08 0.15 0-23 0-30 0.75 1,50 2.25 3.00 - 6.00 9.00 0.000
{TA ~ -- 0.04 0.04 0.05 0.09 0-14 0.20 0.25 0.58 1 . 1 3 1.65 2.28 4.50 7-02 0.023
tTA" 0.05 0.01 0.02 0.03 0-05 0.07 0.09 0.12 0.23 0.42 0-62 0.76 -- - 1.50 1.98 0.023
TB { ToA1 - - 0-02 0.03 0.06 0.15 0-30 0.45 0.60 1 . 5 0 3-00 4.50 6.00 12.00 18-00 0.000
++%A" -- 0.08 0.09 0 - 1 1 0.18 0.30 0.43 0.56 1.32 2.64 3.87 5.33 10.58 16-06 0-065
t % A m 067 022 0.27 0.41 0.48 0.58 0.74 0.90 1.59 2.76 3-88 4.95 9.49 13.73 0.080
{pH ~ -- 5.16 4.89 4.66 4.37 4.14 4.01 3 - 9 1 3.60 3.36 3.20 3.10 2.83 2.66 0-012
tpH ~ 5.53 5.21 4.75 4-49 4.36 4.22 4.11 4.00 3.76 3.56 3.45 3.36 3.13 3.02 0.024
{TA 1 001 0.02 0.03 0.08 0.15 0.23 0.30 0.75 1.50 2.25 3-00 6.00 9-00 0-000
{TA u 0.03 0-04 0.05 0.08 0.13 0.19 0.25 0-60 1 . 1 4 1 . 6 6 2.16 4.31 6-55 0.026
t T A " 0.05 0.03 0.03 0.03 0.06 0.08 010 0 - 1 1 0.21 0.42 0.63 0.84 -- - 1.62 2:05 0-038
{ %A t 0"02 0.03 0-06 0-15 0.30 0.45 0.60 1 . 5 0 3-00 4-50 6-00 12.00 18.00 0.000
IS
{ YoA" 0.06 0.07 0.09 0.16 0.29 0.41 0-54 1-35 2.66 3.97 5.22 -- 10.56 15.67 0-027
t %A" 0.57 0.30 0.30 0-36 0.52 0.62 0.78 0.85 1.63 2.78 3.87 4.92 9.22 12-31 0-108
:[:pH~ 5,23 4.95 4.71 4.39 4-14 400 3.92 3.58 3.35 3.20 3.09 2.81 2.67 0.007
t p H m 5.80 5.24 4.88 4-63 4.39 4.26 4.12 4.04 3.77 3.58 3.43 3.32 --- 313 3-02 0-025

* Titratable acidity calculated as acetic acid in all samples, including meat controls.
TA = total content of acetic acid (g): %A = percentage acetic acid present on wt/wt (meat) or wt/vol (solution) basis: Superscriptsy~ u
and rn refer to fresh equilibrating solution, used equilibrating solution and meat, respectively.
WHC and tenderness of beefmuscles below the ultimate pH 21

Loss of nitrogenous material from meat discs to equilibrating solutions

The loss of nitrogenous material from the meat discs as a function of the
pH of the used equilibrating solutions is illustrated in Fig. 1, where it can
be seen that the greatest loss was found with the IS (30 ~ ) followed by the
TB (27 ~o) and the LD (24 ~). Such losses occurred within the pH range
3.5 to 4.0 which corresponded to original acetic acid concentrations of
0-25~, 0.10M and 0.075r~ for the IS, TB and LD samples, respectively.
There was also a progressive decrease in the loss of nitrogenous material
with decreasing pH to a common value of around 15 ~ for each muscle
type. ]However, at higher pH values, the nitrogen solubility behaviour of
the three muscle types was quite different. While the values for the LD and
TB samples remained between 20 ~o and 25 ~o, there was a progressive
decrease with the IS samples which reached a minimum value of 12 ~o
around pH 5.2.

30
(A)
25
~N
20

15

30

25
~N
20

15

i t t t | | t |

25
~N
2C

15

| t i t n n ,

2.5 3,0 3.5 4.0 4.5 5.0 5.5


pH

Fig. 1. Means plots showing the relationship between pH of used equilibrating solution
and per cent nitrogen solubilized from discs of LD (A), TB (B) and IS (C) muscles. (Eight
observations per mean.) SEM (per cent N solubilized) = 0'56 CA), 0.80 (B) and 0'99 (C).
22 N. F.S. Gault

Influence of pH on swelling ratios (WHC) and cooked meat tenderness

The influence of the pH of the uncooked meat discs on swelling


characteristics and tenderness is illustrated for each muscle type in Fig. 2.
The overall swelling behaviour of the three muscle types was similar
throughout the pH range studied. Minimal swelling of the raw meat discs
occurred between the pHu of the control samples and pH 4.5. Below this
pH there was a marked increase in swelling to the maximum value
achieved around pH 3.4 (LD and TB) and pH 3.2 (IS), corresponding to
an approximate doubling of the volume of the meat discs. It is also worth
noting the decline in swelling with further decreases in the pH of the meat
discs.
Cooking resulted in a loss of weight from all meat discs, particularly
those with a pH > 4.3. However, swelling was substantially retained after
cooking in the more acidic samples, weight loss generally diminishing
with decreasing pH. The relationship between raw meat swelling and
cooked meat swelling for each muscle type was investigated in more detail
by regression analysis of the individual values obtained for cooked meat
swelling against the corresponding values for raw meat swelling. The

TABLE 2
Regression Analyses of Individual Values Obtained for Cooked Meat Swelling (y) Against
the C o r r e s p o n d i n g Values for Raw Meat Swelling (x) of Discs of LD, T B and IS muscles
(N = 112 per Muscle Type)

Musch' Regression equation Percentage t'ariance Let'el o[


accounted.lor sign(ficance

LD y = 1 . 2 5 8 x - 0.674 988 P < 0.001


TB y = 1256x - 0.692 970 P < 0.001
IS v = 1 . 1 6 8 x - 0601 950 P < 0.001

analyses summarised in Table 2 clearly emphasise the dependence of


cooked meat swelling on swelling in the raw state as influenced by the pH
ranges used in this study.
Maximum peak shear force values were achieved within the pH range
4.5 to 5.5, indicating toughness of the cooked meat. However, over the
relatively narrow pH range 4.6 to 4.1, there was a very sharp drop in peak
shear force values, indicating a pronounced increase in the tenderness of
the cooked meat. With a further lowering of pH, the rate of tenderness
WHC and tenderness of beef muscles below the ultimate pH 23

(A) ]9,0
1
8. O ~
2.0
7.0 u
.0 6.0

so
4.0
3.0 ~
o
2,0 ~
1,0
, a j ,i

(B) A~A
9.0
2.( 8.0 t

7.0 °~
.0
6.0

5.0
4. o
u') 3.0
2.0 ~
1.0 ~

(c) 9.0
2.( 8.0 t

7.0
o
~ 1.E 6.0
5.0
o
4.0
u'l
3,0 .~

0.E 2.0
1.0 0.
f,-.,-A- , ~ v ' A ' / . , , ,
3.0 3.5 4.0 4.5 5.0 5.5
pH

Fig. 2. Means plots showing the influence of pH of uncooked meat discs on raw meat
swelling (©), cooked meat swelling (O) and cooked meat peak shear force (A) for the LD
(A), TB (B) and IS (C) muscles. (Eight observations per mean for cooked samples, sixteen
observations per mean for raw samples.) SEM (raw meat swelling) = 0.032 (A), 0.025 (B)
and 0.034 (C): SEM (cooked meat swelling) = 0.030 (A), 0.027 (B) and 0.044 (C): SEM
(peak shear force)= 0348 (A), 0.269 (B) and 0-176 (C).
24 N. F. S. Gault

improvement decreased sharply, maximum tenderness being achieved


around pH 3.3, before increasing marginally at lower pH values.
It is clear that the general improvement in meat tenderness was not only
related to an increased swelling of the meat discs, but also that the major
tenderizing effects were achieved at little more than 50~o maximum
swelling.
Figures 3+ 4 and 5 illustrate in more detail the relationships between

10,0 (A) (B)

9.0

7.0 \o
JP
6.0
~ y = 106.70e-2"913x y : 22.76e-2.326x
5.0
q.o
3.0

2.0 • ~ e o • •

1.0 _"~ o 0

• ~,i~aiaall.,f~L. _. ~
0.6 1.0 1.4 1.8 2.2 0+6 1.0 1.q 1.8 2.2
Swelling ratio S ~ l l i ng ratio

Fig. 3. The relationships between cooked meat peak shear force and swellingratio in
discs of beef M. longissimus lumborum. (A) raw meat swelling,(B) cooked meat swelling.

peak shear force and swelling ratios of the meat discs from the LD, TB
and IS muscles, respectively. Since all of these Figures indicated an
inverse curvilinear relationship between peak shear force and swelling
ratio, it was considered that the data might best be described by a series of
exponential decay equations of the type:
Y=ae -bx

which transforms to the linear equation:


In y = h - bx

where: 6 = In a, y = peak shear force (kg cm-2), and x = swelling ratio.


Regression analyses of the In peak shear force values were therefore
computed against the corresponding values for swelling ratio to quantify
the relationships between peak shear force and swelling ratios illustrated
WHC and tenderness of beef muscles below the ultimate pH 25

°
° (a)
tO.O
i.° (A)
9.0

8.0
'S
.~ 7.0 i°
~ G.O
°
*• • Y : 118.0qe_2.997x y : 22.35e - 2 - 3 7 6 x
~ 5.0 o| •

2.0

1.0

O.G 1.0 1.q 1.8 2.2 0.6 1.0 t.~ 1,8 2.2
~OII irlg ratio Sv~l I i ng r a t i o

Fig. 4. The relationships between cooked meat peak shear force and swelling ratio in
discs of beef M. triceps brachii caput longum. (A) raw meat swelling, (B) cooked meat
swelling.

in Figs 3-5. The derived equations and their statistical significance are
summarised in Table 3. The constants a and b for each of the relationships
between peak shear force and swelling ratio were subsequently used to fit
the curves shown in Figs 3-5. Such results confirm the general trends
illustrated in Fig. 2, and show that quantitative relationships, expressed
by exponential decay equations, exist between cooked meat toughness

10.0 (A) (B)

9.0

r~ 8.0
' Eu
.~ 7.0

~ G .0

,. 5.0

q.0
,•1• Y = 128.38e -3"07fix
°
° \~

-'.~! .
y = 25.71e -2.Gllx

~E 3.0

2.0

1.0
":'~..-

C'.G 1.0 1.q 1.8 2.2 0.6 1.0 1.Zl 1.8 2,2
Svel I i n g r a t i o .~i li ng ratio

Fig. 5. The relationships between cooked meat peak shear force and swelling ratio in
discs of beef M. inJraspinatus. (A) raw meat swelling, (B) cooked meat swelling.
26 N. F. S. Gault

TABLE 3
Regression Analyses of Individual Values of in Peak Shear Force (y) Against the
C o r r e s p o n d i n g Values for Raw Meat Swelling ( R M S ) (x) a n d C o o k e d Meat Swelling
(CMS) (x) of Discs of LD, T B a n d IS Muscles ( N = 112 Per Muscle Type)

Muscle Swelling Regression Percentage variance Level oJ


ratio equation accounted Jbr sign![icanee

LD RMS lny = 4.670- 2.913x 89.1 P < 0.001


CMS lny = 3.125- 2.326x 94.3 P < 0.001
TB RMS lny = 4.771 - 2.997x 94.2 P < 0.001
CMS lny = 3.107- 2.376x 96.3 P < 0.001
IS RMS lny = 4.855 - 3.076x 87.3 P < 0.001
CMS lny = 3.247- 2.61 Ix 90.4 P< 0.001

and both (a) raw meat swelling ratio and (b) cooked meat swelling ratio,
for the LD, TB and IS muscles used in this study.

DISCUSSION

An overall examination of the mean values obtained for titratable acidity


(Table 1) suggests that there was some leaching of weakly acidic buffering
material from the meat discs into the external equilibrating solution. This
was particularly noticeable with the weakest acetic acid solutions, where
the titratable acidity values of the used equilibrating solutions were higher
than those of the fresh solutions. The extensive leaching of nitrogenous
material from the meat discs (Fig. 1) would, in part, account for these
results. However, it is likely that lactic acid, leached from the sarcoplasm
of the fresh meat discs and mainly in the dissociated form, could also have
made a substantial contribution to these observations. Only at acetic acid
concentrations of 0.05M and above were the equilibrating solutions of
sufficient strength to mask this effect.
Despite this buffering effect, it is clear that the apparent distribution of
acetic acid between the meat discs and their equilibrating solutions
changed steadily with increasing acid concentration over the range
studied. This is indicative of a proportionately greater apparent uptake of
available acetic acid by the meat discs with decreasing acid concentration,
showing a greater affinity between the dissociated H + ions of acetic acid
and the meat discs in the weaker equilibrating solutions. Such a trend is
WHC and tenderness of beef muscles below the ultimate pH 27

seen more clearly when the percentage acetic acid contents of the meat
discs and their respective equilibrating solutions are compared. Here, an
apparent distribution balance was reached with the 0.50M (LD) and 0.75M
(TB and IS) acetic acid treatments, beyond which the balance increased in
favour of the equilibrating solutions. Consequently, both the uptake of
acetic acid by the meat discs and their loss of buffering capacity, outlined
above, contributed to the observed pH ranges of the meat discs and their
equilibrating solutions (Table 1).
The nitrogenous material leached from the meat discs (Fig. 1) was most
probably of sarcoplasmic origin, because of the well-established solubility
behaviour of this fraction in contrast to the relative insolubility of
myofibrillar and connective tissue proteins. Lawrie (1979) has also
indicated that non-protein nitrogen and sarcoplasmic protein nitrogen
accou~at for as little as 11 ~o and 26 °/o, respectively of the total nitrogen
content of beef L. dorsi muscle. The maximum losses observed in the
present study could, therefore, account for a substantial proportion of
these components. However, the actual identification of this nitrogenous
material requires further experimental investigation, particularly in the
light of the swelling phenomena discussed later.
The raw meat swelling characteristics of the different muscle types
(Fig. 2) appeared to reflect their differences in connective tissue content,
since the LD and TB samples showed a similar smooth swelling profile
while that of the IS samples was more erratic, exhibiting a pronounced
kink within the pH range 4.3 to 4.0. The swelling behaviour of collagen in
acetic acid solutions is well documented (Gustavson, 1956), and would
account for some of the differences seen over the pH range 4.5 to 4.0
where visible sheets of intramuscular connective tissue in the IS muscle
discs were observed to swell to a greater extent than the myofibrillar
component. Although the diameter of all the meat discs generally
increased to a much greater extent than their thickness, dimensional
changes were too complex to quantify since anisotropic swelling was
regularly observed, particularly with the LD samples, the shape changing
from a circular form to that of an oval.
Despite these differences, the overall raw meat swelling behaviour of
the three muscle types was similar in terms of weight gain. These
relationships (Fig. 2) also bear a striking resemblance to the influence of
acidic pHs on the WHC of ground beef illustrated by H a m m (1960, 1975).
In view of the myofibrillar swelling observed below the IEP by Penny et al.
(1963), it seems reasonable to assume that myofibrillar swelling
28 N. F. S. Gault

contributed extensively to the swelling phenomena observed in the


present study. The extensive loss of nitrogenous material from the meat
discs discussed earlier provides indirect, albeit tentative, support for this
suggestion. Whether or not the hypothesis of Offer & Trinick (1983),
concerning expansion of the myofilament lattices, can explain myofibril-
lar swelling under the more acidic conditions employed here merits
further investigation, particularly in view of the contribution made by
collagen to the swelling observed in the present study. In this context, it is
interesting to note that the maximum swelling observed in the present
study was similar to that observed by Offer & Trinick (1983) for isolated
myofibrils.
Shrinkage of the meat discs on cooking (Fig. 2) was significantly
influenced by the extent of raw meat swelling (Table 2), particularly for
the LD and TB samples. The regression equations in Table 2 also indicate
that greater cooking losses were associated with the lowest raw meat
swelling ratios. This evidence clearly supports the view that the weak
hydration of the myofibrillar proteins at their IEP accounts for greater
losses of water on cooking (Hamm, 1960). At lower pH values, muscle
proteins were more strongly hydrated due to the osmotic pressures
exerted by both the increase in their net positive charge and the presence
of large proportions of acetic acid (see Table 1), thus inhibiting the
characteristic changes associated with heat denaturation of muscle
proteins at the IEP.
With regard to relationships between WHC and meat tenderness, the
present study had certain advantages over previous investigations which
were carried out within the relatively narrow pH range 7.0 to 5.5 normally
associated with meat (Miles & Lawrie, 1970; Bouton et al., 1971;
1972a, b; 1973a, b; McDougall et al., 1979). From Fig. 2 it is clear that
the widest possible range of meat swelling was achieved, not only through
the maximum at the lower pH values, but also through the minimum at
pH values above and below the IEP. With the exception of the control
samples, the use of extraneous ~water' not only enhanced the influence of
pH on ~water' uptake, but also made possible the measurements of WHC
and tenderness on the same samples of meat.
Consequently, the relationships between swelling ratio and peak shear
force illustrated in Fig. 2 could be quantified, as shown in Figs 3 to 5. The
general shape of these curves clearly demonstrates the marked
toughening which occurred in the meat samples at the lowest swelling
ratios and the diminishing benefit of excessive meat swelling on their
WHC and tenderness of beef muscles below the ultimate p H 29

tenderness at the higher swelling ratios. The contribution of the


considerable variability in peak shear force values obtained at the lowest
swelling ratios to the shape of these curves is quite obvious. Such
exponential relationships clearly emphasise the importance of WHC in
relation to cooked meat tenderness as influenced by pH below the pHu.
From the regression analyses summarised in Table 3, it can also be
concluded that the connective tissue content of the meat had a marginal
influence on these relationships.
In the context of normal meat handling, where the pH of post-rigor
meat can theoretically vary from pH 7.0 down to pH 5.4, it has repeatedly
been demonstrated with beef that meat of a higher than normal pH u is
more tender than meat of a lower pH u from similar muscles (Khan &
Lentz, 1973; Fredeen et al., 1974; Fjelkner-Modig & Rud6rus, 1983).
Furthermore, Dransfield (1977) has demonstrated that some of the
observed variability in cooked meat tenderness between eighteen different
types of beef muscle was influenced by differences in their PHu, improved
tenderness again being correlated with higher pHu. Moreover, in those
studie,; where meat covering a wide pH range was used, linear
relationships between cooked meat tenderness and increasing pH have
been established for rabbit (Miles & Lawrie, 1970), mutton (Bouton et al.,
1971 ; i[972a, b), beef(Bouton et al., 1973a, b) and venison (McDougall et
al., 1979).
In comparison with these studies, it is clear from Fig. 2 of the present
study ~Lhat strong linear relationships between cooked meat tenderness
and decreasing pH were achieved within the pH range 4.6 to 4.1. The
c o m m o n feature of both sets of relationships is that they covered pH
ranges either basic (generally pH 55 to pH 7.0) or acidic (pH 4.6 to 4.1) to
the IEP of the myofibrillar proteins. Such similarity of behaviour on
either ~,;ideof the IEP suggests that the influence of a higher than normal
pH u on meat tenderness is primarily and extensively due to an increased
WHC of the meat. Consequently, the results obtained in the present study
strongly support and enhance the traditional opinion that the beneficial
influence of higher pH values on cooked meat tenderness post-rigor is
indirectly a result of the beneficial influence of pH on the WHC of meat
above the IEP. That such relationships, influenced by muscle pH, also
exist in. pre-rigor muscle is strongly supported by the work of Miles &
Lawrie (1970) and even more directly by the detailed study of Currie &
Wolfe (1980) on the mechanical properties of uncooked meat. However,
in view of the renowned bacteriological spoilage problems associated with
30 N. F. S. Gault

meat of pHu > 6 . 0 (Gill & Newton, 1981), only the influence of W H C
below the I E P could be seriously considered for industrial exploitation as
a practical method for tenderizing meat, as the results of the present study
show.

ACKNOWLEDGEMENTS

The author gratefully acknowledges the skilful technical assistance of Mr


E. Gilpin and Mr E. Tolland and the help of Dr D. Kilpatrick and Miss
Eileen Kelly in carrying out the statistical analysis of the results. Some
aspects of this work were presented at the 30th European Meeting of
Meat Research Workers, Session 4:12, l lth September, 1984, Bristol,
Great Britain.

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